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Volume 118, Number 1

March 31, 2004

9912:

CONTENTS ISSN 0028-1344

M. G. Harasewych The deep-sea Buccinoidea (Gastropoda: Neogastropoda) of

Yuri I. Kantor the Scotia Sea and adjacent abyssal plains and trenches.................... I

Lindsey T. Groves New species of Late Cretaceous Cypraeidae (Gastropoda)

from California and British Columbia and new records

from the Pacific slope

OOKMRE VIC WA Ea ORNs keer ed eee inn he welts Me atone aa enn Oe rte ete Sl NED,

THE NAUTILUS 118(1):1-42, 2004

Page |

The deep-sea Buccinoidea (Gastropoda: Neogastropoda) of the

Scotia Sea and adjacent abyssal plains and trenches

M. G. Harasewych Yuri I. Kantor

Department of Systematic Biology

National Museum of Natural History

Smithsonian Institution

Washington, DC 20013-7012 USA

Severtzov Institute

Russian Academy of Sciences

Leninski Prospect 33,

Moscow 117071 RUSSIA

ABSTRACT

Four new genera and species of buccinoidean gastropods, Spi-

kebuccinum stephaniae new genus, new species; Drepanodon-

tus tatyanae new genus, new species; Muffinbuccinum cath-

erinae new genus, new species; and Germonea rachelae new

genus, new species, are described from the Scotia tectonic

plate and adjacent abyssal plains. Only Bathydomus obtectus

Thiele, 1912, Tromina bella abyssicola Clarke, 1961 and T.

abyssorum Lus, 1993, had previously been reported from abys-

sal depths off Antarctica. The latter two species were proposed

in the genus Tromina, subsequently shown to belong to the

family Muricidae. Therefore, a new genus, Lusitromina is pro-

posed for these abyssal and hadal Ihueratmosdleem species. Anal-

yses of the taxonomic placement, geographical and bathymetric

distribution, and diversity of the 29 buccinoidean genera pres-

ently known from Antarctica and the Magellanic Province have

shown that the abyssal (>2200 m) buccinoidean fauna of the

region shares no genera with the sublittoral or bathyal faunas.

None of the six abyssal genera conform readily to the subfam-

ilies represented by the sublittoral or bathyal faunas. Credible

sister taxa and likely origins for some abyssal genera occur on

the adjacent continental ‘slope. For others, alosext relatives may

be found on abyssal plains beyond the Antarctic convergence.

Generic diversity decreases with increasing depth for both the

bathyal and abyssal buccinoidean faunas, while bathymetric

range tends to increase. For abyssal buccinoideans, maximum

generic diversity occurs between 2600 and 3200 meters. The

proportion of monotypic genera in the Antarctic and Magel-

lanic Provinces is extraordinarily high (48.3%), and may ie. an

artefact of low sampling density exacerbated by difficulties in

differentiating closely related species. Neither gigantism nor

dwarfism is auiilont in the abyssal buccinoidean fauna. Rather,

the range in sizes narrows with increasing depth. Genera in-

habiting the base of the continental slope are smaller than

those of either the upper slope or continental rise. In the abys-

sal zone, maximum shell size is reached near the boundary of

the continental rise and abyssal plain, and subsequently de-

creases with increasing depth.

INTRODUCTION

The Buccinoidea are the most geographically wide-

spread and ecologically diverse clade within the Neo-

gastropoda. First appearing during the Early Cretaceous

[Valanginian] (Tracey et al., 1993), these predatory snails

have radiated to occupy most benthic marine habitats

ranging from the tropics to the poles and from the in-

tertidal zone to hadal depths (Clarke, 1962). Several

members of the families Nassariidae and Buccinidae

have even invaded fresh water (Kantor and Kilburn,

2001; Brandt and Temcharoen, 1971).

Buccinoideans are readily distinguished by their usu-

ally weakly sculptured, conical to fusiform shells, their

distinctive rachiglossan radula with multicuspid lateral

teeth, long to very long proboscis, as well as by the ab-

sence of a rectal gland and accessory salivary glands.

Their relationships to other Neogastropoda, however,

have been vee) interpreted, ranging from basal to

derived (e.g., Ponder, 1974: Ponder and Warén, 1988:

Ponder ee Lindberg, 1996; Kantor, 1996; Harasewych

et al., 1997). While a number of authors have attributed

different taxonomic ranks to Buccinoidea and its com-

ponent higher taxa (e.g., Powell, 1929; Thiele, 1929;

Wenz, 1938: Ponder, 1974; Ponder and Warén, 1988),

there is little disagreement as to the monophyly or com-

position of the group. We had earlier briefly reviewed

the history of the higher classification of buccinoideans

(Harasewych and Kantor, 1999), which is based primarily

on differences in shell, opercular and radular morphol-

ogies applied to regional faunas (e.g., Powell, 1929,

1951, Southern Oceans; Habe and Sato, 1973, Northern

Pacific: Bouchet and Warén, 1985, Northeastern Atlan-

tic). We continue to retain provisionally the use of Buc-

cinulidae and its subdivisions, as defined by Powell

(1951), without necessarily endorsing their taxonomic

rank, for the antiboreal members of the Buccinoidea,

pending the availability of sufficient anatomical and/or

molecular data for a meaningful phylogenetic revision of

the higher taxa of Buccinoidea on a global basis. The

subfamilial assignments of presently known buccino-

idean genera that occur south of the Antarctic Conver-

gence, as well as those from the Magellanic Province are

reviewed (Appendix 1) and, in some cases, revised.

Our continuing studies of the Buccinoidea represent-

ed in the collections assembled by the United States

Antarctic Program (USAP) have revealed a number of

D, 9

Page 2

previously undescribed taxa from the abyssal plains and

trenches on and adjacent to the Scotia Plate. These taxa

are described herein, and their affinities to other Ant-

arctic and abyssal buccinoideans are discussed.

MATERIALS AND METHODS

This report is based primarily on buccinoideans sorted

from the abyssal stations sampled by the United States

Antarctic Program (USAP) vessels R/V IsLas OrcADAS

and R/V ELTanin and housed in the collections of the

National Museum of Natural History (USNM). Addi-

tional material, sampled by the German vessel R/V Po-

LARSTERN and in the collection of the Zoological State

Collection, Munich (ZSM) were made available through

the kindness of Enrico Schwabe and Michael Schrédl.

In the material examined sections, “specimen” de-

notes that a preserved animal is present, while “shell”

refers to a record based only on an empty shell. Ana-

tomical descriptions are based on gross dissections of

preserved specimens. Radulae were removed by gross

dissection, cleaned using diluted bleach (NaOCl), coated

with carbon and gold, and examined using a LEO 440

Scanning Electron Microscope. Photographs were taken

using a Nikon D1 Digital Camera with a AF Micro Nik-

kor 60 mm lens. Images were processed using Adobe

Photoshop 6.0.

The following abbreviations are used in the text: AL—

aperture length, D—diameter, FWL—final whorl

length, L—length, L/W—length/width, SCL—Siphonal

canal length, SL—shell length, SW—shell width, W—

width.

SYSTEMATICS

Class Gastropoda Cuvier, 1797

Order Neogastropoda Wenz, 1938

Superfamily Buccinoidea Rafinesque, 1815

Family Buccinulidae Finlay, 1928

Subfamily Buccinulinae Finlay, 1928

Genus Spikebuccinum new genus

Type Species: Spikebuccinum stephaniae new spe-

cies, by original designation.

Description: Protoconch large (to 3.5 mm diameter),

of 274 smooth, evenly rounded, whorls. Teleoconch small

(to 19.9 mm), very thin, translucent, ovate, rounded an-

teriorly, all but final 2-2% whorls eroded. Shell sculpture

of sharp, closely spaced, spiral cords. Columella weakly

concave, shorter than aperture, producing siphonal

notch with weak siphonal fasciole, short pseudoumbili-

cus. Periostracum finely hirsute. Operculum ovate, pau-

cispiral, with subterminal nucleus, spans ~% aperture

length. Eyes absent. Buccal mass, odontophoral cartilag-

es longer than retracted proboscis. Rachidian teeth tri-

cuspid, with outer cusps broader, longer than central

cusp. Lateral teeth with large outer, shorter inner cusp,

with 3-4 denticles between. Salivary glands small, un-

fused. Gland of Leiblein glandular anteriorly, flaccid pos-

THE NAUTILUS, Vol. 118, No. 1

teriorly. Posterior oesophagus forms crop before enter-

ing simple, U-shaped stomach, which has a well-defined

gastric shield.

Etymology: This genus is named after Spike, a Cor-

nish Rex cat that belongs to the senior author's daughter

Stephanie.

Spikebuccinum stephaniae new species

(Figures 1-23, Table 1)

Description: Shell (Figures 1-3, 5-7, 9, 10) small (to

19.9 mm), very thin, translucent, ovate, with rounded

anterior, eroded spire. Protoconch (Figure 11), known

from a single juvenile (Paratype 12), increasing in di-

ameter from 0.4 mm to 3.5 mm, in 2% smooth, evenly

rounded, pitted whorls. Transition to teleoconch distin-

guished by slight change in color, from cream to white,

and by abrupt transition from coarse, irregular axial

growth striae, to finer, regular growth lines. Protoconch

and upper whorls eroded on all other specimens. Ex-

trapolation from growth series suggests that teleoconch

may reach 5-6 whorls, of which all but last 2-2% whorls

eroded. Whorls evenly rounded, with indistinct shoulder,

abutting suture. Axial sculpture limited to very fine,

straight, strongly prosocline growth lines. Spiral sculp-

ture of fine, sharp, uniform, evenly spaced cords (21-29

on final whorl, 11-14 on penultimate whorl). Aperture

large (AL/SL = 0.60-0.67 when using length of eroded

shell; AL/SL = 0.50-0.55, as estimated by linear projec-

tions of apex), broadly oval, deflected from shell axis by

22-25°. Outer lip very thin, not reflected, evenly round-

ed from suture to siphonal notch. Inner lip consists of a

long, straight parietal region that meets the shorter, con-

cavely indented axial portion of columella, ending in

strong siphonal fold. Columella shorter than aperture,

giving rise to a broad siphonal notch. Parietal callus uni-

formly narrow from suture to siphonal fold. Short, weak,

siphonal fasciole and pseudoumbilicus present, often ob-

scured by erosion. Shell color uniformly white. Perios-

tracum very thin, straw yellow in color, with densely

spaced axial lamellae, producing short, fine hairs at in-

tersection with spiral cords, giving shell a finely hirsute

appearance. Operculum (Figures 4, 8) small, spanning

~0.36 AL, yellowish brown, broadly ovate, paucispiral,

with subterminal nucleus rotated relative to opercular

axis.

Anatomy (Holotype): Soft tissues (Figures 14-22)

comprise approximately 2% whorls. Mantle cavity spans

just under % whorl, kidney % whorl, digestive g gland and

gonad just under 1% whorl. Columellar muscle short,

broad, comprising slightly more than one whorl, at-

tached to shell at rear of mantle cavity. Foot large,

broadly rectangular (L/W = 1). Body color yellowish tan,

without pigmentation. Head large with long, thin taper-

ing tentacles (Figure 15, tn), oethoue neck. “Eyes absent.

Nephridium with semi-transparent walls that clearly re-

veal folds. Nephridial gland (Figure 14, ng) small, very

narrow. Pericardium oriented antero-ventrally. Digestive

M. G. Harasewych and Y. I. Kantor, 2004 Page 3

Figures 1-11. Shells and opercula of Spikebuccinum stephaniae new species. 1. Apertural, 2. lateral, 3. and dorsal views of

holotype, USNM 896368, off South Georgia Island, 53°02’ S, 37°40'00" W, in 3056-3102 m [R/V ELTAanIn cruise 9, sta. 735]. 4

Outer view of operculum of holotype. 5. Apertural, 6. lateral, and 7. dorsal views of Paratype 1, USNM 1010626, from the type

locality. 8. Outer view of operculum of Paratype 1. 9. Apertural view, USNM 1010630, E off South Sandwich Islands, 58°27’ S,

29°22" W, in 4643-4645 m [R/V ELTANIN sta. 603]. 10. Apertural view, Paratype 8, USNM 1010629, S off Southern Georgia Island,

58°04’ S, 37°50’ W, in 3255-3166 m [R/V ELTANIN sta. 699]. 11. Scanning electron micrograph of protoconch of Paratype 12, ZSM

(Zoological State Collection, Munich) 20021125, E off South Samebirieln Islands, 58°24.98’ S, 25° 1.00’ W, in 2285.5 m, [R/V

POLARSTERN cruise ANTXIX, sta. PS61/141-8, 22 Mar 2002] Arrow indicates transition to teleoconch. 5 mm scale bar applies to all

shells, 1 mm scale bar applies to opercula, 2 mm scale bar applies to protoconch.

Page 4

THE NAUTILUS, Vol. 118, No. 1

Figures 12-13. Radula of the holotype of Spikebuccinum stephaniae new species. 12. Dorsal, and 13. Left lateral (30°) views

of the central portion of the radular ribbon.

gland lobes (Figure 19, adg, pdg) not fused, separated

by ovary (Figure 19, ov).

Mantle Cavity (Figure 20): Mantle cavity of medium

width (L/W ~0.8), mantle edge slightly serrated. Siphon

long (0.42 AL), free, muscular, extending substantially

bey rand mantle edge. Osphradium (Figure 20, os) small

(~0.4 mantle cavity length) yellowish, bipectinate, with

narrow osphradial nerve. Ctenidium (Figure 20, ct)

large, very wide, spans about %4 of mantle cavity length.

The ctenidial lamellae are low. Hypobranchial gland

lacks distinct folds, covered by thick layer of mucus.

Alimentary System (Figures 14-19): Proboscis (Fig-

ures 16-18, pr) of moderate length when retracted

(~0.36 SL, 0.54 AL), thick (L/D ~3.3), smooth, non-

pigmented. Proboscis retractor muscles (Figures 16-18,

prr) not numerous, but thick, powerful, attached to pro-

boscis sheath at mid-length when proboscis retracted.

Proboscis sheath thin-walled, translucent along anterior

half, thickened posteriorly, but thinner than proboscis

wall. Proboscis wall thick, comprising ~1/10 of retracted

proboscis diameter. Mouth opening dorso-ventrally com-

pressed slit. Buccal mass muscular, large, filling retracted

proboscis and protruding significantly Iyewonnel its rear.

Odontophoral cartilages paired, fused anteriorly, slightly

longer, than retracted proboscis. Radular ribbon longer

adhem retracted proboscis, 5.4 mm long (0.41 AL), about

480 wm wide (0.037 AL), triserial (Figures 12-13), con-

sisting of 58 rows of teeth, posteriormost 6 rows nascent.

Rachidian teeth with 3 short cusps emanating from cen-

tral portion of broad, anteriorly deeply pried basal

plate. Central cusp slightly shorter, narrower than lateral

cusps. Lateral teeth with long basal plate flanked by two

main cusps, outermost near ly twice as long as innermost,

34 smaller, intermediate denticles vary in size and po-

sition from row to row, innermost denticle often abutting

inner cusp.

Salivary glands (Figures 16-18, rsg, Isg) small, acinous,

yellowish, not fused. Right salivary gland (rsg) dorsal to

nerve ring, enveloping most of valve of Leiblein. Left

salivary gland (Isg) lateral to, partially covering nerve

ring. Salivary ducts (Figures 17-18, sd) thick, attached

both to oesophagus and proboscis sheath by numerous

connective tissue fibers. Salivary ducts pass along both

sides of oesophagus, become * ‘embedded” into oesoph-

agus walls immediately after entering the proboscis.

fate of Leiblein (Figure 18, vL) well defined, large,

pyriform, with whitish glandular pad containing ciliary

cone visible through walls of valve.

Gland of Leiblein (Figures 16-18, gL) large, massive,

glandular anteriorly; flaccid, transparent, lacking glan-

dular tissue posteriorly; opens, via short duct, into oe-

sophagus just posterior to nerve ring. Gland yellowish,

only slightly darker than other organs of the cephalic

haemocoel.

Oesophagus thick anterior to nerve ring, becoming

thin-walled, flattened posterior to nerve ring. Oesopha- ~

M. G. Harasewych and Y. I. Kantor, 2004 Page 5

Figures 14-22. Anatomy of Spikebuccinum stephaniae new species. 14-20. Holotype, 21-22. Paratype 1. 14. Right, and 15.

Left lateral views of animal removed from shell. 16. Left, 17. ventral, and 18. right lateral views of anterior alimentary system. 19.

Dorsal view of stomach, digestive glands and ovary. 20. Mantle cavity organs. 21. Lateral view of penis. 22. Ventral view of seminal

vesicle. Scale bars = 5 mm for 14-21, 1 mm for 22. a, anus; adg, anterior duct of the digestive gland; ag, albumen gland; eg,

capsule gland; eme, cut mantle edge; ct, ctenidium; dg, digestive gland; gL, gland of Leiblein; go, female genital opening; hg,

hypobranchial gland; Isg, left salivary gland; nep, nephridium; ng, nephridial gland; nr, circumoesophageal nerve ring; oe, oe-

sophagus; op, operculum; os, osphradium; ov, ovary; pdg, posterior duct of the digestive gland; poe, posterior oesophagus; pr,

proboscis; prr, proboscis retractors; re, rectum; rsg, right salivary gland; s, siphon; sd, salivary duct; st, stomach; tn, cephalic

tentacles: vL, valve of Leiblein.

Page 6

THE NAUTILUS, Vol. 118, No. 1

30°W

a type locality

& examined material

No. of specimens 2 4 6 8 10

Figure 23. Geographic and bathymetric distribution of Spikebuccinum stephaniae new species.

gus widens to form a “crop” (Figure 19, poe) before

entering stomach.

Stomach (Figures 14, 15, 19, st) relatively large,

broadly U-shaped, without posterior mixing area. Due to

poor preservation of holotype it was impossible to ex-

amine the internal morphology of the stomach in detail.

Digestive gland ducts paired. Arrangement of stomach

similar to Lusitromina abyssorum (Figure 132) but dif-

fers in having well developed, highly cuticularized gastric

shield, with crescent-shaped dorsal side that is lifted and

significantly protruded into stomach lumen.

Female Reproductive System: Female reproductive

system typically buccinoidean, with small albumen gland

partially overlapping posterior portion of capsule gland

dorsally. Ingesting gland small, opening between albu-

men and long, broad capsule gland. Small bursa copu-

latrix situated anterior to capsule gland, tapering ante-

riorly to form female opening.

Male Reproductive System: Paratype 1, mature

male. Seminal vesicle (Figure 22) of medium size, spans

> % whorl, formed of numerous loops. Penis (Figure

21) long, narrow, non-pigmented, with slightly folded

walls. Penial papilla long, cylindrical, surrounded by cir-

cular fold around base.

Type Locality: Off South Georgia Island, 53°02’ S,

37°40’ W, in 3056-3102 m [R/V ELTANIN cruise 9, sta.

735, 13 Sep 1963].

Type Material: Holotype, 2, USNM 896368; Para-

types 14, USNM 1010626, all from the type locality.

Paratypes 5-7, USNM 1010628, off South Georgia Is-

land, 53°26.7' S, 36°32.6’ W, in 1967-2186 m, [R/V IsLas

Orcapas sta. 28, 12 May 1976]; Paratypes 8-11, USNM

1010629, S of Southern Georgia Island, 58°04’ S, 37°50’

W, in 3255-3166 m, [R/V ELTANIN cruise 9, sta. 699, 30

Aug 1963]; Paratype 12, ZSM 20021125, E of South

Sandwich Islands, 58°24.98’ S, 25° 1.00’ W, in 2285.5

m, [R/V PoLarsTERN cruise ANTXIX, sta. PS61/141-8,

22, Mar 2002].

Table 1. Spikebuccinum stephaniae new species. Measurements of shell characters. Linear measurements in mm. (n = 7, including

holotype. Juvenile specimens excluded).

Character Mean

Shell length (SL) 15.6

Final whorl length (FWL) 13.9

Aperture length (AL) 10.3

Shell width (SW) 10.1

FWL/SL 0.89

AL/SL 0.65

SW/SL 0.64

Number of spiral cords on penultimate whorl 12.4

Number of spiral cords on final whorl 24.6

o Range Holotype

212, 13.4-19.9 19.9

2.2 11.9-18.0 18.0

ed §.3-13.2 13.2

1.6 8.6-13.2 13.2

0.04 0.81—0.91 0.90

0.03 0.60-0.68 0.66

0.02 0.59—-0.66 0.66

1.0 11-14 14

2.9 21-29 29

M. G. Harasewych and Y. I. Kantor, 2004

Other Material Examined: USNM 896337, E of

South Sandwich Islands, 54°51’ S, 14°54’ W, in 3947—

4063 m, [R/V ELTANIN sta. 1571, 28 Feb 1966], 1 spec-

imen; USNM 1010630, E of South Sandwich Islands,

58°27’ S, 22°92’ W, in 4643-4645 m, [R/V ELTANIN

cruise 9, sta. 603, 5 May 1963], 1 specimen + 1 shell.

Distribution (Figure 23): The species is found in the

Scotia Sea and adjacent abyssal plains at depths of 1967—

4645 m.

Etymology: This species is named for the senior au-

thor’s elder daughter, Stephanie Alexandra Harasewych.

Remarks: The shell morphology of Spikebuccinum

stephaniae superficially resembles that of several species

of Chlanidota, especially C. signeyana Powell, 1971 and

(Pfefferia) invenusta Harasewych and Kantor, 1999,

and, to a lesser extent, an eroded Neobuccinum eatoni

(Smith, 1875). However, it can readily be distinguished

from these taxa by its very short columella, an incon-

spicuous siphonal fasciole, as well as by having a rachi-

dian tooth in which the central cusp is shorter and nar-

rower, rather than longer and stouter than the outer

cusps. The lateral teeth of all species of Chlanidota and

the monotypic Neobuccinum have a strong, single inter-

mediate cusp, rather than the multiple denticles of Spi-

kebuccinum. The shell of Spikebuccinum stephaniae is

also somewhat similar to that of Antarctodomus okutanii

Numanami, 1996, which has a clearly cominelline rad-

ula, with tricuspid rachidian and bicuspid lateral teeth.

The radular morphology of Spikebuccinwm stephaniae

is distinctive, and suffices to distinguish it from all Ant-

arctic and Magellanic bueainatdacmns, The presence of

multiple denticles between the flanking cusps of the lat-

eral teeth would appear to preclude the inclusion of this

genus in the subfamily Buccinulinae, which is defined

on the basis of having tricuspid rachidian and lateral

teeth (Powell, 1951). However, Powell (1951:131) ex-

panded this criterion to include Bathydomus Thiele,

1912, within Buccinulinae, citing the conchological affin-

ities of Bathydomus to Chlanidota. We questionably in-

clude Spikebuccinum within the subfamily Buccinulinae,

noting the possibly pleisiomorphic similarity of its radula

to such boreal and temperate taxa as Neptunea, some

Buccinum, Cantharus and certain Busycotypus.

The strongly paucispiral operculum of Spikebuccinum

stephaniae is a feature it shares with a number of genera,

among them Neobuccinum, Parficulina Powell, 1958,

Falsitromina Dell, 1990, Parabuccinum Harasewych,

Kantor and Linse, 2000, and such boreal genera as

Mohnia Friele in Kobelt, 1878, and Pararetifusus Ko-

suge, 1976. While operculum morphology is undoubt-

edly useful for distinguishing genera, its utility for dis-

cerning phylogenetic relationships among supraspecific

taxa is less clear.

Comparative anatomical data is available for only a

very few buccinulid taxa, among them Chlanidota (Har-

asewych and Kantor, 1999) and Parabuccinum (Hara-

sewych, Kantor and Linse, 2000). Of these, Spikebuccin-

Page 7

um appears most similar anatomically to Chlanidota, but

differs in having proportionally longer odontophoral car-

tilages, salivary glands that are not fused, a valve of Lei-

blein with a ciliary cone, a gland of Leiblein that opens

to the mid-oesophagus via a narrow rather than broad

duct, a broader stomach with a well-defined gastric

shield, and a tapering rather than hemispherical bursa

copulatrix.

Drepanodontus new genus

Type Species: Drepanodontus tatyanae new species,

by original designation.

Description: Protoconch eroded. Shell large (to 56

mm), thin, strongly fusiform, with long, broad, open si-

phonal canal. Early whorls straight to concave, subse-

quently becoming convex. Whorls lacking shoulder, final

whorl evenly convex from suture to distinct peripheral

cord, evenly concave from peripheral cord to tip of si-

phonal canal. Sculpture of axial growth lines, most pro-

nounced on early whorls, and strong, evenly spaced spi-

ral cords. Aperture large, narrow, elliptical. Outer lip not

reflected. Parietal region broad posteriorly, columella

shorter than siphonal fold. Periostracum lamellose.

Operculum large (~0.82 AL), oval, thin, translucent,

with terminal nucleus. Head medium-sized, with short,

stout tentacles, large eyes. Kidney large. Buccal mass

small, odontophoral cartilages short. Rachidian teeth

variable, tricuspid, cusps may be fused to appear mon-

ocuspid, or supplemental denticles or cusps may be pre-

sent in some specimens. Lateral teeth with single, large,

sickle-shaped cusp that may have one or more denticles.

Salivary glands not fused, right gland much larger than

left. Giana of Leiblein long, glandular anteriorly, taper-

ing, flaccid posteriorly. Oesophagus broadens to form

crop, constricts before entering very large, U-shaped

stomach that lacks posterior mixing area.

Etymology: Drepane (Gr.)—sickle, odontos (Gr.)—

tooth. The name reflects the very unusual sickle shape

of the lateral teeth, consisting of a long, narrow basal

plate with a single, long, recurved outer cusp, often with

one or more denticles along its inner edge.

Drepanodontus tatyanae new species

(Figures 24-50, Table 2)

Description: Shell (Figures 24-30, 33-35) large (to

43.1 mm), thin, strongly fusiform. Protoconch eroded on

all specimens. Early whorls of holotype (Figure 27) in-

dicate that protoconch would likely have been ~2-2%

whorls, < 3 mm in diameter. Transition to teleoconch

indistinct in holotype. Teleoconch estimated to consist

of up to 7 whorls. Early teleoconch whorls slightly con-

cave in profile, with narrow spiral cords. By anne whorl,

teleoconch becomes convex, increasingly so in subse-

quent whorls, forming evenly rounded, oval whorls with-

out distinct shoulder. Suture abutting onto pronounced

peripheral cord (Figures 24, 33, arrow). Axial sculpture

Page 8 THE NAUTILUS, Vol. 118, No. 1

Figures 24-35. Shells and opercula of Drepanodontus tatyanae new species. 24. Apertural, 25. lateral, and 26. dorsal views of

the holotype, USNM 1010544, NE of South Shetland Islands, 59°01’ S, 52°00’ W, in 3010-3510 m. [R/V ELTANIN cruise 22, Sta.

1511]. 27. Enlarged upper shell whorls of holotype. 28. Apertural, 29. lateral, and 30. dorsal views of Paratype 1, USNM 881529,

E of South Sandwich Islands, 57°00.24’ S, 26°10.06’ W, in 2740-2757 m. [R/V ELTANIN cruise 575, sta. 38]. 31. Outer, and 32.

inner views of the operculum of Paratype 1. 33. Apertural, 34. lateral, and 35. dorsal views of the shell of paratype 2, USNM

881529. Same locality as Paratype 1.

M. G. Harasewych and Y. I. Kantor, 2004

A _ ta

ee J -

Page 9

Figures 36-40. Radulae of Drepanodontus tatyanae new species. 36. Dorsal, and 37. left lateral (45°) views of the central portion

of the radular ribbon of Paratype 1, USNM 881529. 38. Dorsal, 39. left lateral (45°) views of the central portion of the radular

ribbon, and 40. left lateral (45°) view of the bending plane of a non-type specimen, USNM 1010545, Argentinean Basin, 47°17.3'

S, 47°45.7' W, in 5685-5798 m [R/V IsLas Orcapa, cruise 575, sta. 4].

of pronounced growth lines, opisthocyrt on early whorls,

becoming weaker, orthocline by fourth teleoconch

whorl. Spiral sculpture of broad, sharp cords (9-12 on

penultimate whorl, 10-14 from suture to peripheral

cord, 16-24 from peripheral cord to tip of siphon) nar-

rower than intervening spaces, with 0-4 very fine

threads between adjacent cords, especially near periph-

ery. Peripheral cord, slightly thicker more pronounced

that others, demarcates inflection in curvature of outer

lip, evenly convex abapically, evenly concave from pe-

ripheral cord to tip of siphonal canal. Aperture large

(AL/SL = 0.56), roughly elliptical, deflected from shell

axis by 13-17°. Siphonal canal long (~% shell length),

broad, open, crosses shell axis. Outer lip thin, not re-

flected. Inner lip of weakly concave parietal region,

slightly longer columella, long siphonal fold extending

Page 10 THE NAUTILUS, Vol. 118, No. 1

Figures 41-49. Anatomy of Drepanodontus tatyanae new species. (41-48. Paratype 1. 49. Paratype 2, both USNM 881529). 41—

42. Lateral views of animal removed from shell. 43. Ventral, 44. right lateral, and 45. left lateral views of the anterior alimentary

system. 46. Latero-dorsal view of the anterior oesophagus, showing the duct of gland of Leiblein. Oesophagus twisted counterclockwise.

47. Dorsal view of stomach, dashed line indicates the posterior nephridial border. 48. Mantle complex of organs. 49. Lateral view of

penis. Scale bars = 1 cm for Figures 41-42, 47-49, 5 mm for Figures 43-46. a, anus; adg, anterior duct of the digestive gland; agd,

anterior lobe of digestive gland; ao, aorta; cg, capsule gland; em, columellar muscle; eme, cut mantle edge; et, ctenidium; dgL, duct

of gland of Leiblein; e, eye; gL, gland of Leiblein; go, female genital opening; Isg, left salivary gland; mo, mouth opening; nep,

nephridium; ng, nephridial gland; nr, circumoesophageal nerve ring; oe, oesophagus; op, operculum; os, osphradium; ov, ovary; pdg,

posterior lobe of digestive gland; per, pericardium; poe, posterior oesophagus; pr, proboscis; prp, propodium; prr, proboscis retractors;

re, rectum; rsg, right salivary gland; s, siphon; sd, salivary duct; st, stomach; vL, valve of Leiblein.

M. G. Harasewych and Y. I. Kantor, 2004

Page 11

2000

2200

50° 2400

2600

2800

3000

3200

3400

3600

& examined material

40°W

ty type locality Tana

60° 3800)

5400)

5600

3 0°W 5800

Depth in m

No. of specimens

Figure 50. Geographic and bathymetric distribution of Drepanodontus tatyanae new species.

beyond glossy, translucent parietal region, broad poste-

riorly, tapering toward base of siphonal canal. Neither

siphonal fasciole nor pseudoumbilicus present. Shell uni-

formly white, aperture, parietal callus glossy. Periostrac-

um straw-colored, of widely spaced, short axial lamellae.

Operculum (Figures 31-32, 41-42, op), large (~0.82

AL), oval, thin, brownish yellow, flexible, with terminal

nucleus. Attachment of columellar muscle clearly visible

through operculum, oval, spans about % of operculum

surface.

Anatomy (Paratype 1): Soft tissues (Figures 41-42)

comprise approximately 34% whorls. Mantle cavity spans

slightly less than % whorl. Kidney (Figures 41, 42, nep)

broad, spans slightly less than % whorl, with 12 trans-

verse folds of unequal width visible through wall. Ne-

phridial gland (Figure 41, ng) narrow, situated antero-

dorsally to the nephridium. Digestive glands (Figures 41,

42, adg, pdg) of 2% whorls. Columellar muscle (Figure

42, cm) of 1% whorls, attached to shell at rear of mantle

cavity. Foot medium-sized, short in contracted specimen

(L/W ~1.2), with well-developed narrow propodium

(Figure 42, prp) separated by narrow propodial cleft.

Body color yellowish tan, without pigmentation. Head

medium-sized, with short, stout, conical tentacles, with

small but distinct lobes at their bases. Eyes present (Fig-

ure 42, e), light gray in color and semitransparent, deep-

ly embedded into lobes. Mantle does not cover base of

head.

Mantle Cavity (Figure 48): Mantle cavity deep (L/

W ~1.2). Mantle edge smooth, slightly thickened. Si-

Table 2. Drepanodontus tatyanae new species. Measurements of shell characters. Linear measurements in mm. (n = 3, including

holotype).

Character Mean o Range Holotype

Shell length (SL) 45.1 10.1 36.2—56.0 43.1

Final whorl length (FWL) 33.8 74 28.6-42.2 30.5

Aperture length (AL) 26.7 6.2 22. 2-33.7 94.1

Siphonal canal length (SCL) 11.4 ALT 7.5-16.7 10.1

Shell width (SW) 19.1 2.3 17.5-21.8 18.1

FWL/SL 0.75 0.04 0.71-0.79 0.71

AL/SL 0.59 0.03 0.56—-0.61 0.56

SCL/SL 0.25 0.05 0.21—0.30 0.23

SW/SL 0.43 0.05 (.39-0.48 0.42

Number of spiral cords on penultimate whorl 10.0 15 9-12 10

Number of spiral cords suture to peripheral cord 12.0 2.0 10-14 12

Number of spiral cords peripheral cord to siphon 19.7 4.0 16-24 19

Page 12

THE NAUTILUS, Vol. 118, No. 1

phon (Figure 48, s) short, free, muscular, broad, extend-

ing well beyond mantle edge, with thick siphonal edge

(Figure 48, se), covering anteriormost part of ctenidium.

Osphradium (Figure 48, os) greenish, bipectinate, large,

wide, % as long, 2h as nidle as ctenidium. Ctenidium

(Figure 48, a large, curved, spanning nearly entire

mantle cavity length. Ctenidial lamellae broad, triangu-

lar, with short recurved edges along posterior part (alos.

er to pericardium), gradually becoming narrower, rela-

tively taller anteriorly. Hypobranchial g sland lacks distinct

folds, covered by thick layer of mucus. Rectum long,

spanning ~4/5 of mantle cavity length.

Alimentary System (Figures 41-47): Proboscis non-

pigmented, (Figures 43, 45, pr) short when retracted

(~0.2 SL, 0.58 AL), thick (L/D ~3), with slightly folded

walls. Proboscis retractor muscles (Figures 45, 46, prr)

not numerous, thin, attached to thin- walled, translucent

proboscis sheath at middle-posterior region when pro-

boscis retracted. Proboscis wall thin, ~Il/ 10 proboscis di-

ameter. Anterior part of proboscis flattened to form rim

surrounding mouth opening (Figure 43, mo) in form of

irregular tr angular slit. Anterior “oesophagus very broad,

nearly filling proboscis. Dorsal folds very large, border-

ing deep groove. Buccal mass small, spans slightly more

lhe % of proboscis length. Odontophoral cartilages

paired, fused anteriorly, extend nearly entire length” of

buccal mass, but <% proboscis length. Radular ribbon

(Figures 36-40) equal in length to “cartilages, 6.6 mm

(0.35 AL), about 380 wm wide (0.020 AL), triserial, con-

sisting of 50 rows of teeth, posteriormost 5 rows nascent.

Rachidian teeth of Paratype 1 (Figures 36-37) with 4

cusps, here interpreted as comprising a long central

cusp, flanked by shorter, outer cusps, with an additional,

asymmetrical cusp on the left side. Rachidian teeth of a

second specimen (F igures 38-40) appear monocuspid,

but “central cusp” consists of 3 incompletely fused

cusps, flanked by additional small denticles (Figure 40).

Lateral teeth with distinctive shape, with single large,

recurved outer cusp emanating from long, narrow Ibe

plate. Inner surface of cusp with 1 or more, occasionally

bifid, posteriorly directed denticles, with number, size of

denticles varying from side to side and along radular rib-

bon.

Right salivary gland medium-sized, rounded, partially

covering valve of Leiblein (Figures 43, 44, rsg), latero-

dorsal to nerve ring. Left salivary gland slightly smaller

than right, irregularly shaped, dorsal to nerve ring, ven-

tral to proboscis with its main axis perpendicular to pro-

boscis axis, appears small when viewed from left (Figure

45, Isg). Salivary ducts (Figures 44, 46, sd) short, thick,

enter oesophagus wall shortly after leaving gland. Valve

of Leiblein (Figure 46, vL) well defined, large, pyriform,

with whitish glandular pad visible through welll of valve.

Gland of ienion (Figures 43-45, gL) yellowish,

slightly darker than other organs of cephalic haemocoel,

naeekonn sized, long, tubular, coiled anteriorly. Gland

thin-walled, ascinous anteriorly (Figure 46, agL), opens

into oesophagus slightly posterior to nerve ring via

broad, short duct (Figure 46, dgL), becomes thinner,

more transparent posteriorly (Figure 46, pgL), tapering

to become flaccid, non-glandular (Figure 46, vgL).

Oesophagus thick, broad anterior to nerve ring, nar-

rowing slightly posterior to the ring. Posterior oesopha-

gus expands greatly to form “crop,” (Figure 47, poe)

liven © gradually narrows towards opening into stonmeaclh.

Stomach (Figure 47) very large, spans ~% whorl, from

the posterior border of nephridium, U-shaped, ition

posterior mixing area. Preservation inadequate to dis-

cern internal morphology. Digestive glands ducts (Fig-

ure 47, ddg) large, paired, closely spaced. Posterior duct

close to oesophagus entrance, anterior duct at mid-

length of stomach. Digestive glands clearly separate. An-

terior gland small (Fi igure 41, adg), spans ~% whorl,

posterior gland ~2 whorls (Figures 41, 42, pdg). Glands

meet at the level of the posterior duct to digestive gland.

Rectum long, spans ~4/5 of mantle cavity length. Rec-

tum thin-walled, very broad, filled with polychaete spic-

ules, numerous sand grains of different sizes.

Female Reproductive System: Paratype 1, mature

female. Pallial gonoduct consists of long, tubular, capsule

gland (Figures 42, 48, cg), with a small bursa copulatrix

anterior to it. Genital opening (Figure 48, go) below,

slightly posterior to anus (Figure 48, a).

Male Reproductive System: Paratype 2, male. Penis

(Figure 49) long, very narrow, flattened laterally. Seminal

papilla very small, blunt, surrounded by deep circular

fold around its base.

Type Locality: NE of South Shetland Islands, 59°01’

S, 52°00’ W, in 3010-3510 m. [R/V ELTANIN cruise 22,

Sta. 1511, 26 Jan 1966].

Type Material: Holotype, USNM 1010544, from the

type locality. Paratype 12, Paratype 2 3d, USNM

881529, E of South Sandwich Islands, 57°00.24’ S,

26°10.06’ W, in 2740-2757 m. [R/V ELTANIN cruise 575,

Sta. 38, 22 May 1975].

Other Material Examined: USNM 1010545, South

Atlantic Ocean [Argentine Abyssal Plain], 47°17.3' S,

47°45.7' W, in 5685-5798 m. [R/V Istas Orncapas Cruise

575, Sta. 4, 8 May 1975], 2 bodies without shells (radula

illustrated, Figures 38-40); USNM 1010546, Scotia Sea,

S of South Georgia Island, 58°04’ S, 37°50’ W, 3255—

3166 m. [R/V ELTANIN cruise 9, sta. 699, 30 Aug 1963],

1 dead poorly preserved juvenile. USNM 1013084, Sco-

tia Sea, SW of South Georgia Island, 55°56’ S, 44°56’

W, 3742-3614 m. [R/V LAN cruise 575, Sta. 472, 13

Feb 1963], 1 body and fragments of the shell.

Distribution (Figure 50): This species occurs in the

Scotia Sea and adjacent Argentine Abyssal Plain, at

depths of 2740-5798 m.

Etymology: This species is named in honor of the ju-

nior author's wife, Tatyana Steiker, an ichthyologist and

illustrator at the P. P. Shirsov Institute of Oceanology.

Remarks: The large, elongate, fusiform, siphonate,

M. G. Harasewych and Y. I. Kantor, 2004

Page 13

spirally corded shell of Drepanodontus tatyanae easily

distinguishes this species from most Antarctic buccino-

ideans. Conchological similarity is limited to relatively

few large taxa, notably Antarctoneptunea aurora (Hed-

ley, 1912) and Cavineptunea monstrosa Powell, 1951,

both members of continental shelf and upper slope fau-

nas. Drepanodontus tatyanae is most easily distinguished

from the former by having a distinctive peripheral spiral

cord that demarcates a change in the direction of cur-

vature in the outer lip, coshalle the monotypic Cavinep-

tunea is most easily distinguished by its unique, cylin-

drical, flat-sided, indented “protoconch. Both Antarcto-

neptunea (Dell, 1972: fig. 6) and Cavineptunea (Powell,

1951: 145) have radulae with tricuspid rachidian teeth

and lateral teeth with 3 (or 4) cusps, quite unlike the

distinctive radula of Drepanodontus.

The radula of Drepanodontus tatyanae most closely

resembles that of Kapala bathybius Bouchet and Warén,

1986 (Bouchet and Warén, 1986: fig. 8), a species in-

habiting the Cape Basin off southwestem Africa at

depths of 3550 m. Like Drepanodontus, K. bathybius

has rachidian teeth that may appear to be monocuspid

in some individuals, with anteriorly indented, squarish

basal plates, and lateral teeth characterized by a single,

large, sickle-like cusp with secondary denticles that vary

in number and prominence from side to side and from

tooth to tooth. The shell of K. bathybius, and the related

K. bonaespei (Barnard, 1963), also from the Cape Basin

in 2504-3103 m, are comparable in size, and also elon-

gate, fusiform and spirally corded, but broader (Bouchet

and Warén, 1986: figs. 42, 43), and lack the distinctive

peripheral cord of Drepanodontus. Barnard (1963: 432,

fig. 6b) illustrates and describes the radula of K. bon-

aespei (which he described as a Neptunea) to have rect-

angular rachidian teeth with a “median cusp, sometimes

a minute denticle on one side or on both sides” and

lateral teeth “unequally bicuspid, with 2-5 tiny denticles

between the two cusps, the denticles not always sym-

metrical.”

Barnard (1963) assigned this species to the genus

Neptunea Réding, 1798, because of the similarity of its

lateral teeth with those of boreal buccinoideans, despite

striking differences in the morphology of the shell and

rachidian teeth. Bouchet and Warén, (1986: 464) also

commented on lateral tooth similarities of Kapala (in-

cluding its type species, the southern Australian bathyal

species K. kengrahami Ponder, 1982) with the type spe-

cies of the boreal genera Volutopsius Mérch, 1857, Neo-

beringius Habe and Ito, 1965, Ancistrolepis Dall, 1895,

and certain representatives of Japelion Dall, 1918. These

boreal taxa have elongated, spatulate lateral teeth with a

large outer cusp, a significantly smaller inner cusp, and

a variable number of smaller denticles or cusps between

them. We interpret the lateral teeth of Drepanodontus

to be different, in that they have a single, large outer

cusp, but lack the shorter inner cusp of the boreal taxa.

While some of the denticles that frequently emerge

from the inner edge of the cusp of Drepanodontus may

be large enough to be confused with an inner cusp, the

lateral teeth of the boreal species are fundamentally bi-

cuspid, while those of Drepanodontus are fundamentally

monocuspid. Interestingly, the radula of K. bathybius iL

lustrated by Bouchet and Warén (1986: fig. 8) has mon-

ocuspid lateral teeth lacking denticles distill but de-

veloping denticles proximally along the left side of the

radula, while the lateral teeth on Bhe right side have 1

large and 1-3 smaller denticles along the inner edge of

the single cusp.

The operculum of Drepanodontus is large, ovate, and

has a terminal nucleus. While all species of Kapala share

this opercular morphology, it is not distinctive, but wide-

spread throughout Buccinoidea.

Although the presence of large eyes is not surprising

in the bathyal type species of Kapala (Ponder, 1982: fig.

2), their occurrence in the abyssal taxa K. bonaespei

(Barnard, 1963: 432) and Drepanodontus tatyanae is

noteworthy. Other anatomical features that are congru-

ent between Drepanodontus tatyanae and Kapala ken-

grahami include a small buccal mass and odontophore,

a large kidney, a large, well-developed valve of Leiblein,

crop, and a simple stomach.

Drepanodontus tatyanae co-occurs with Muffinbuc-

cinum catherinae at the type locality of that species.

Subfamily Prosiphiinae Powell, 1951

Muffinbuccinum new genus

Type Species: Muffinbuccinum catherinae new spe-

cies, by original designation.

Description: Protoconch, early whorls eroded. Teleo-

conch of moderate size (to ~30 mm) very thin, chalky,

with evenly rounded whorls, tapering anterior, short,

broad siphonal canal. Sculpture of sharp, narrow axial

ribs, crossing uniform, closely spaced spiral cords, oc-

casionally forming finely reticulate surface sculpture. Ap-

erture large, elongate, elliptical. Parietal region broader

posteriorly. Columella short, not axial. Periostracum of

very fine axial lamellae, not hirsute. Operculum, large

(~0.56 aperture length), elongated, claw-like, with ter-

minal nucleus and growth lines nearly perpendicular to

long axis. Head large, with tapering tentacles lacking

eyes. Columellar muscle short. Nephridium very narrow.

Rachidian teeth broadly rectangular, with 3 cusps con-

centrated near center of tooth, central cusp longest. Lat-

eral teeth as broad as rachidian tooth, roughly rectan-

gular in outline, with 7-8 cusps of nearly equal length,

outermost cusp, slightly shorter, stouter, weakly to

strongly serrated along outer margin. Salivary glands

large, fused ventrally. Gland of Leiblein long, narrowly

tubular, highly coiled. Oesophagus muscular, does not

widen before opening into stomach. Stomach very small,

broadly U-shaped, lacks posterior mixing area. Rectum

short, narrow, with terminal papilla.

Etymology: This genus is named after Muffin, a do-

mestic shorthair cat that belongs to the senior author's

daughter Catherine.

Page 14

THE NAUTILUS, Vol. 118, No. 1

Muffinbuccinum catherinae new species

(Figures 51-76, Table 3)

Description: Shell (Figures 51-56) of moderate size

(to 26.6 mm), very thin, chalky, ovate, with tapering an-

terior, eroded spire. Protoconch and upper whorls erod-

ed on all available specimens, with no more than last 2%

whorls remaining. Extrapolation suggests that un-eroded

specimens might reach 32 mm, with 5-6 teleoconch

whorls. Whorls evenly rounded, with indistinct shoulder,

abutting suture. Axial sculpture of fine, sigmoidal, weakly

prosocline growth lines, narrower than spiral cords, vary-

ing in prominence, producing reticulate pattern in some

areas of the shell, especially on penultimate whorl (Fig-

ures 51, 52, arrows). Spiral sculpture of fine, sharp, uni-

form, evenly spaced cords (43-50 on final whorl, 18-21

on penultimate whorl), weaker, more broadly spaced be-

tween suture and shoulder as well as near siphon. Ap-

erture large (AL/SL = 0.64-0.71 when using length of

eroded shell: AL/SL = 0.54-0.60 as estimated by linear

projection of apex), elongate, ovate, tapering anteriorly,

deflected from shell axis by 17—18°. Outer lip very thin,

not reflected, evenly round from suture to siphonal

notch. Inner lip of broad parietal region, shorter, non-

axial columellar region, long siphonal fold and short,

broad, axial siphonal canal. Parietal callus consisting of

thin glaze, slightly broader posteriorly, narrower or in-

dented near siphonal fold, may form barely perceptible

pseudo-umbilicus-like indentation. Shell color unifor mly

white. Periostracum of very thin axial blades, straw yel-

low in color. Operculum (Figures 57-58) large, spanning

~0.56 AL, translucent amber yellow in color, with dis-

tinctive, claw-like shape, tapering toward nucleus, which

is eroded. Outer surface with numerous growth lines

nearly perpendicular to long axis. Inner surface with out-

er rim slightly thickened, glazed, except for long, narrow,

triangular attachment area.

Anatomy (Holotype): Body (Figures 65-66) compris-

es approximately 2% whorls. Mantle cavity spans ~%

whorl, mantle edge thin, does not cover head or penis

base. Nephridium very narrow (Figure 65, nep), spans

<1/6 whorl, brown-grey. Border between mantle cavity

and nephridium marked by deep cleft. Digestive gland

(Figures 65, 66, dg) spans 1% whorl, ovenetia by testis.

Columellar muscle very short, spanning <1 whorl,

broad, attached to shell anterior at rear of mantle cavity.

Foot large, narrowly oval (L/W ~2.3 for holotype, up to

2.8 in other specimens), not folded during fixation. Body

light yellowish, without pigmentation. Head large, with

broad, tapering cylindrical tentacles (Figure 65, 66, tn)

lacking eyes.

Mantle Cavity (Figure 67): Mantle cavity short,

broad (L/W ~0.63). Siphon very short, thin, weakly

muscular, extending slightly beyond mantle edge. Os-

phradium (Figure 67, os) greenish, bipectinate, with

broadly curved axis, spanning < 0.4 mantle cavity length.

Ctenidium (Fi igure 67, ct) large, narrower than osphra-

dium, strongly curved, spanning % mantle cavity length.

Hypobranchial gland formed of numerous, indistinct,

closely spaced oblique folds covered by thick layer of

mucus that partly overlays the rectum.

Alimentary System (Figures 70-75): Proboscis (Fig-

ure 71, pr) short (~0.46 AL in holotype), thick, smooth-

walled, non-pigmented, not coiled, occupies entire ce-

phalic haemocoel. Proboscis sheath extremely thin-

walled, anterior half translucent, thicker posteriorly, but

<%, thickness of proboscis wall. Mouth opening (Figure

73) rounded, lumen of buccal tube triangular immedi-

ately posterior to mouth. Buccal mass muscular, medi-

um-sized, slightly shorter than retracted proboscis.

Odontophoral cartilages paired, fused anteriorly, span-

ning nearly entire length of buccal mass. Radular ribbon

slightly longer than cartilages (~0.30 AL in holotype),

~480 xm wide (holotype), triserial (Figures 59-64),

consisting of about 70 (holotype) to 85 (paratype 1) rows

of teeth, posteriormost.5 teeth nascent. Oldest several

rows of teeth with dark brown pigment that faded when

radula cleaned with bleach (NaOCl). Rachidian teeth

with broad, rectangular, slightly arched base with

straight lateral sides, 3 large, robust, closely spaced

cusps. Central cusp slightly longer than lateral cusps.

Lateral teeth as broad as rachidian teeth, roughly rect-

angular, with 7-8 cusps nearly equal in length, outermost

cusp shortest, stouter than others, weakly (Figure 61) to

strongly (Figure 64) serrated along outer margin. Sali-

vary glands (Fi igures 70-72, sg) large in comparison to

proboscis, acinous, yellowish, fused ventrally, dorsal to

oesophagus. Right salivary gland covers valve of Leiblein

(Figure 70, vL), part of nerve ring and anterior of pro-

boseis. Left salivary gland shifted posteriorly. Salivary

glands tightly attached to proboscis sheath by connective

Hose. Salivary ducts (Figures 70, 72, sd) thick, free

along both sides of oesophagus, becoming embedded in

oesophagus walls shortly after entering retracted pro-

boscis. Valve of Leiblein (Figures 70, 79 74, vL) well

defined, large, pyriform, with ciliary cone, yellow, slightly

darker than other organs of cephalic haemocoel. Gland

of Leiblein (Figures 70-72, 74, gL) long, dark grey, nar-

rowly tubular, glandular, highly coiled, covered by con-

nective tissue sheath, opens into middle oesophagus pos-

terior to nerve ring via short, narrow duct (Figure 74,

dgL). Oesophagus muscular, nearly constant in diameter

along most of its length, does not widen before opening

into stomach. Stomach (Figure 75) very small, broadly

U-shaped, lacks posterior mixing area. Preservation was

inadequate to document sniermul morphology of stom-

ach. Rectum (Figure 67, re) short, narrow, thin-walled,

with terminal papilla, spans slightly more than half the

length of mantle cavity.

Male Reproductive System: Holotype, Paratypes 1

and 2, males. Testis large, anterior margin at posterior-

most part of stomach extends posteriorly along digestive

gland for 1% whorls. Testis similar in color and texture

to digestive gland. Seminal vesicle small, formed of very

few loops. Penis (Figures 66, p; 68, 69) non- -pigmented,

spans entire length of mantle cavity, flattened in middle

M. G. Harasewych and Y. I. Kantor, 2004 Page 15

Figures 51-58. Shells and operculum of Muffinbuccinum catherinae new species. 51. Apertural, 52. lateral, 53. dorsal, and 54.

apical views of the holotype, USNM 1010623, South Atlantic Ocean [Argentine Abyssal Plain], 47°17.3' S, 47°45.7' W, in 5685—

5798 m [R/V Istas Orcapas Cruise 575, Sta. 4]. 55. Apertural, and 56. lateral views of the shell of Paratype 1, USNM 1010624,

from the type locality. 57. Inner, and 58. outer views of operculum of holotype. Arrows indicate regions of finely reticulate surface

sculpture.

THE NAUTILUS, Vol. 118, No.

50 u m

Figures 59-64. Radulae of Muffinbuccinum catherinae new species. 59-61. Paratype 1. 62-64. Holotype. 59, 62. Dorsal, and

60, 63. right lateral (45°) views of the central portion of the radular ribbon. 61, 64. Enlarged right lateral views of lateral teeth

showing the denticles along the outer cusp.

M. G. Harasewych and Y. I. Kantor, 2004

Page 17

Figures 65-69. Anatomy of Muffinbuccinum catherinae new species, holotype. 65. Left, and 66. right lateral views of animal

removed from shell. 67. Mantle cavity organs. 68. Dorsal, and 69. right lateral views of the penis. Scale bars = 5 mm. eme, cut

mantle edge; colm, columellar muscle; ct, ctenidium; dg, digestive gland; hg, hypobranchial gland; nep, nephridium; op, oper-

culum; os, osphradium; p, penis; per, pericardium; prp, propodium; re, rectum; s, siphon; st, stomach; tes, testis; tn, cephalic

tentacles.

part, widens anteriorly. Long, cylindrical papilla at distal

end of penis surrounded by circular fold around its base.

Type Locality: South Atlantic Ocean [Argentine

Abyssal Plain], 47°17.3’ S, 47°45.7’ W, in 5685-5798 m.

[R/V Istas Orcapas Cruise 575, Sta. 4, 8 May 1975].

Type Material: Holotype, d, USNM 1010623; Para-

type 1, d, shell partially broken, Paratype 2, 6, shell

fragmented, USNM 1010624, all from the type locality.

Other Material Examined: USNM 1010625, frag-

ments and soft tissues of 3 specimens from the type

locality. One was dissected to illustrate the anatomy of

the anterior foregut.

Distribution (Figure 76): This species is known only

from the type locality, the Argentine Abyssal Plain, off

the northern slope of the Falkland Plateau, at depths of

5685-5798 m.

Etymology: This species is named for the senior au-

thors younger daughter, Catherine Laura Harasewych.

Remarks: Of the Antarctic buccinoideans, this new

species appears conchologically most similar to Spike-

Page 18

THE NAUTILUS, Vol. 118, No. 1

prr

wD. oe eL

Figures 70-75. Digestive system of Muffinbuccinum catherinae new species. 70-74. Female specimen without a shell, USNM

1010625. 75. Holotype. 70. Left lateral, 71. right lateral, and 72. ventral views of anterior alimentary system. 73. Anterior view of

the proboscis tip. 74. Right lateral view of the anterior oesophagus showing the opening of the duct of gland of Leiblein and valve

of Leiblein. Salivary glands removed. 75. Dorsal view of stomach. Scale Bare = 5 mm for Figures 70-73, 75, and 2 mm for Figure

74. dgL, duct of gland of Leiblein; gL, gland of Leiblein; Isd, duct of left salivary gland; nr, circumoesophageal nerve ring: oe,

oesophagus; poe, posterior oesophagus; pr, proboscis; prr, proboscis retractors; rsd, “aunt of right salivary gland; sd, salivary aes

sg, salivary gland; vL, valve of Leiblein.

buccinum stephaniae and, to a lesser extent, to Chlani-

dota (Pfefferia) invenusta. It differs from both in having

a more elongated shell, with a long siphonal fold and a

short, broad, anteriorly rounded siphonal canal. The sur-

face sculpture of Muffinbuccinum catherinae is finer, and

the occasional, weak reticulate sculpture of Muffinbuc-

cinum is absent in Spikebuccinum and Chlanidota.

The distinctive radular morphology of Muffinbuccin-

um catherinae, with its broad, tricuspid rachidian and

pectinate lateral teeth, readily distinguishes this species

from other buccinoidean taxa. Based on the morphology

of the radula, Muffinbuccinum catherinae appears to be

related to Proneptunea Thiele, 1912 (Thiele, 1904, pl. 9,

fig. 58; Powell, 1951, fig. K 66-67), a circumantarctic

genus that occurs from sublittoral (Proneptunea fenes-

trata Powell, 1951, 17-27 m) to bathyal (Proneptunea

rossiana Dell, 1990, 369-870 m) depths. Muffinbuccin-

um and Proneptunea both have tricuspid rachidian teeth

and multicuspid lateral teeth that may bear serrations

along their outer edge. However, the rachidian teeth of

Muffinbuccinum are as broad as the lateral teeth, shorter

than wide, and have the cusps concentrated in the mid-

dle portion of the tooth, while the rachidian teeth of

species of Proneptunea are narrower, with the three

cusps spanning much or their width. The longest and

most prominent of the 7-8 cusps on the lateral teeth of

Muffinbuccinum is the innermost cusp, while the largest

and most prominent of the 5 cusps on the lateral teeth

M. G. Harasewych and Y. I. Kantor, 2004

Page 19

South Georgi

55°

Depth in m

40°W

® type locality

160°

5200

5400)

5600)

° 5800

30°W 12345

No. of specimens

Figure 76. Geographic and bathymetric distribution of Muffinbuccinum catherinae new species.

of the 3 (of 6) species of Proneptunea for which the

radular morphology is known, is invariably the outermost

cusp. Based on the presence of a tricuspid rachidian and

multicuspid lateral teeth serrated along their outer edge

in Muffinbuccinum, and their overall resemblance to the

radulae of Proneptunea, we include Muffinbuccinum in

the subfamily Prosiphiinae.

Similarities also exist between Muffinbuccinum cath-

erinae and several bathyal, abyssal, and hadal buccino-

ideans of uncertain affinities, notably Calliloncha Lus,

1978, Costaria Golikoy, 1977, Thalassoplanes Dall, 1908,

and Troschelia Morch, 1876. The radula of Calliloncha

[as Calliloconcha] knudseni Bouchet and Warén, 1986

(Bouchet and Warén, 1986: fig. 1), a species from the

Kermadec Trench (in 5480 m) resembles that of Muffin-

buccinum catherinae in having a tricuspid rachidian

tooth that is nearly as broad as the lateral teeth, and

lateral teeth with 5 cusps of equal length, but lacking

serrations along their outer margin. Bouchet and Warén

(1986:484) noted similarities in the radulae of Bayerius

Olsson, 1971 [Olsson, 1971: fig. 101 for B. fragilissimus

(Dall, 1908) (type species), and Warén and Bouchet,

2001: figs. 39b,c) for B. arnoldi (Lus, 1981) and B. pe-

ruvianus Warén and Bouchet, 2001, respectively] and

Calliloncha Lus, 1978 [Lus, 1978: figs. 2 (7-10) for C.

solida Lus, 1978 (type species), and Lus, 1989: figs. 5

(1-2) for C. iturupi Lus, 1989], and speculated that

these genera may prove to be synonyms. However, the

type species of Calliloncha, which dwells in the Izu-Bo-

nin Trench (6770 m), C. iturupi, from the Kurile-Kam-

chatka Trench (8240 m), Bayerius fragillisimus (Dall,

1908), known from off Panama and Ecuador (2877-3200

m), B. arnoldi, from the Japan, Kuril, and Aleutian

Trenches (4800-6135 m), and B. peruvianus from 5385—

5996 m off Peru all share tricuspid lateral teeth that

differ in dentition and proportion from those of Calli-

Table 3. Muffinbuccinum catherinae new species. Measurements of shell characters of Holotype and Paratype 1. Paratype 2

fragmented. Linear measurements in mm.

Character

Shell length (SL)

Final whorl length (FWL)

Aperture length (AL)

Siphonal canal length (SCL)

Shell width (SW)

FWL/SL

AL/SL

SCL/SL

SW/SL

Number of spiral cords on penultimate whorl

Number of spiral cords on final whorl

Holotype Paratype 1

26.4 26.6

24.6 23.7

19.1 20.8

2.4 3.6

16.3 15.6

0.93 0.89

0.72 0.78

0.09 0.14

0.62 0.59

18 21

43 50

Page 20

THE NAUTILUS, Vol. 118, No. 1

loncha knudseni. While Calliloncha may prove to be a

synonym of Bayerius, it seems unlikely that C. knudseni

is referable to this genus. The radulae of Costaria

(Bouchet and Warén, 1986: fig. 6), Thalassoplanes

(Bouchet and Warén, 1986: fig. 5) and Troschelia

(Bouchet and Warén, 1985: figs. 454455) all have lateral

teeth that are roughly rectangular and multicuspid.

However, these three genera have rachidian teeth that

are very narrow. In Costaria and Thalassoplanes the cen-

tral of three cusps may be reduced or absent, while in

Troschelia the flanking cusps are absent, resulting in

monocuspid rachidian teeth.

The operculum of Muffinbuccinum catherinae is rath-

er distinctive in being claw-shaped, with growth lines

nearly perpendicular to the long axis, and a long, narrow,

triangular attachment area along its inner surface. A sur-

vey of the literature shows that this operculum type also

occurs in Proneptunea fenestrata (Powell, 1951: fig.

N124), as well as in Thalassoplanes moerchi (Bouchet

and Warén, 1986: fig. 25), Calliloncha knudseni (Bouch-

et and Warén, 1986: fig. 32), but not C. solida, the type

of the genus, (Lus, 1978: fig. 2, 4-5) both from the abys-

sal to hadal zones of the Pacific Ocean, and in Liomesus

ovum (Turton, 1825) (Bouchet and Warén, 1985: fig.

440) a North Atlantic species inhabiting outer continen-

tal shelf depths. Muffinbuccinum catherinae co-occurs

with Drepanodontus tatyanae new genus, new species

(see above).

Germonea new genus

Type Species: Germonea rachelae new species, by

original designation.

Description: Protoconch large (to 4.4 mm), of ~2%

whorls, first 2 whorls smooth, rounded, followed by on-

set broad, rounded spiral cords. Transition to teleoconch

distinct, marked by axial indentation, change in shell col-

or. Teleoconch large (to 67.8 mm), of up to 5 smooth,

evenly ovate whorls, with indistinct shoulder, abutting

suture. Axial sculpture of fine growth lines, irregular ru-

gae near suture. Spiral sculpture of closely spaced cords.

Aperture large, narrowly elliptical, with short, broad, si-

phonal canal. Outer lip may be slightly reflected, thick-

ened along margin. Inner lip with glossy parietal callus,

long columella, siphonal fold. Neither siphonal fasciole,

nor pseudoumbilicus present. Periostracum thin, tightly

adherent. Operculum large (~0.65 AL), long, narrow,

with terminal nucleus at end of strongly curved anterior

margin. Columellar muscle very short (~% whorl). Foot

with deep propodial cleft. Head large, with short neck,

large black eyes at base of broad, tapering tentacles.

Buccal mass large, filling proboscis. Odontophore short-

er, radula longer than buccal mass. Rachidian teeth long,

narrow (L/W ~2.5), Y-shaped, with 3 stout, radially ori-

ented cusps, central cusp longest. Lateral teeth with 3—

4 cusps of similar length along innermost % of basal

plate. Salivary glands large, right larger than left. Gland

of Leiblein large, ascinous. Oesophagus thick, broad.

Stomach U-shaped, lacking posterior mixing area.

Etymology: The genus is named after Mrs. Rachel

(Raye) N. Germon, in recognition of her years of work

in managing the collections of Antarctic Mollusca at the

National Museum of Natural History, Smithsonian In-

stitution.

Germonea rachelae new species

(Figures 77-98, Table 4)

Description: Shell (Figures 77-79, 82-86) large (to

67.8 mm), thin to moderately thick, fusiform. Proto-

conch (Figure 87) increasing from 1.1 mm to 4.4 mm in

diameter in ~2% whorls, first 2 whorls smooth, rounded,

followed by onset of weak, broad, rounded spiral cords.

Transition to teleoconch marked by distinct axial inden-

tation, change in shell color, with axial growth lines, spi-

ral cords markedly more prominent. Teleoconch of up

to 5 smooth, evenly ovate whorls, with indistinct shoul-

der, abutting suture. Axial sculpture limited to fine,

straight, weakly prosocline growth lines, some more pro-

nounced, giving rise to irregularly spaced, weak surface

rugae near suture. Spiral sculpture of numerous (18-30

on penultimate whorl; 48-81 on last whorl), closely

spaced cords, broader than intervening spaces. Finer

threads may occur between adjacent cords, especially

along anterior portion of final whorl. Aperture large (AL/

SL = 0.49-0.57), narrowly elliptical, deflected from shell

axis by 14-17°. Siphonal canal short (~% aperture

length), broad, open, crosses shell axis. Outer lip may be

slightly reflected, weakly thickened along margin, evenly

rounded from suture to base of siphonal canal. Inner lip

of short, weakly convex parietal region, long, slightly

concave columella, long siphonal fold extending beyond

narrow glossy parietal callus. Neither siphonal fasciole,

nor pseudoumbilicus present. Protoconch ivory colored,

teleoconch uniformly white, aperture, parietal callus

glossy. Periostracum amber yellow to brown, tightly ad-

herent to shell. Operculum (Figures 80, 81) large (~0.65

AL), translucent yellow to amber in color, long, narrow,

with terminal nucleus at end of strongly curved anterior

margin. Outer surface of operculum with fine, regular

growth lines. Inner surface with small, oval attachment

area along posterior, adaxial region, broad, strongly

thickened, heavily glazed free outer and anterior mar-

gins.

Anatomy (Holotype): Based on rehydrated animal,

soft tissues (lacking upper part of kidney, digestive

gland) comprise approximately 14% whorl. Mantle cavity

spans <¥% whorl, mantle edge thick, does not cover head

or base of penis. Columellar muscle very short, com-

prising ~% whorl. Foot of medium size, strongly con-

tracted, broadly oval (L/W ~1.4), with narrow propo-

dium separated by deep propodial cleft. Body yellowish

tan, non-pigmented. Head large, with short neck, broad,

tapering tentacles, large black eyes at base of tentacles.

M. G. Harasewych and Y. I. Kantor, 2004 Page 21

Figures 77-87. Shells and opercula of Germonea rachelae new species. 77. Apertural, 78. Lateral, and 79. dorsal views of the

holotype, USNM 896594, off South Georgia Island, 58°02’ S, 37°57’ W, in 3197-3239 m [R/V ELTANIN cruise 9, sta. 698]. 80.

Outer and 81. Inner views of the operculum of the holotype. 82. Apertural and 83. Dorsal views of paratype 1, USNM 1010620,

N of South Orkney Islands, 58°06’ S, 44°56’ W, in 2800 m [R/V ELTantn, cruise 7, sta. 480]. 84. Apertural and 85. dorsal views

of paratype 2, USNM 898828, off South Georgia Island, 55°02’ S, 44°21’ W, in 3623-3714 m [R/V ELTANIN cruise 7, sta. 469]. 86.

Apertural and 87. apical views of paratype 3, USNM 1010621, off South Georgia Island, 54°59’ S, 38°13’ W, in 2379-2196 m

[R/V ELTANN, cruise 9, sta. 686].

THE NAUTILUS, Vol. 118, No. 1

Figures 88-92.

100 um

Radula of holotype of Germonea rachelae new species. 88. Dorsal, and 89. right lateral (45°) views of the central

portion of the radular ribbon. 90. Single lateral tooth (partially broken) to show concentration of cusps along inner portion of basal

plate. 91. Dorsal view of the bending plane, showing wom teeth. 92. Right lateral (75°) view of the central portion of the radular

ribbon.

Siphon very short, free, muscular, extends slightly be-

yond mantle edge.

Mantle Cavity: Mantle of female specimen, although

damaged, was partially present. Mantle of moderate

width (L/W ~0.8), very thin, mantle edge evenly thick-

ened. Osphradium yellowish, bipectinate, symmetrical,

narrow, spanning ~0.7 mantle cavity length. Ctenidium

equal in width to osphradium, slightly longer, curved.

Lamellae uniform in shape along entire ctenidium

length, tall, triangular, with short curved tip, similar to

that in posterior part of Lusitromina abyssorum (Lus,

1993) (Figure 129). Hypobranchial gland poorly devel-

oped, lacks distinct folds. Rectum spans ~% mantle cav-

ity length, with terminal anus, distinct anal papilla.

Alimentary System (Figures 95-97): Proboscis (Fig-

ures 95, 97, pr) short when retracted (~0.30 AL), nar-

row (L/D ~4.0), with folded walls, non-pigmented. Pro-

boscis retractors (Figures 95, 96, prr) powerful, arranged

in symmetrical bundles attached to posterior part of pro-

boscis sheath when proboscis retracted. Anterior % of

proboscis sheath very thin-walled, translucent, thicken-

ing posteriorly until as thick as proboscis wall, which

comprises ~0.1 proboscis diameter. Mouth opening tri-

angular slit. Buccal mass muscular, large, filling, but not

extending beyond retracted proboscis. Odontophoral

cartilages paired, fused anteriorly, ~% of buccal mass

length. Radular ribbon of holotype ( (Figures 88-92) lon-

ger her proboscis (14 mm, 0.42 AL), narrow, (~420

ym), triserial, consisting of 110 rows, anteriormost 28

M. G. Harasewych and Y. I. Kantor, 2004

Figures 93-97. Anatomy of Germonea rachelae new species. 93-94. Holotype, male. 95-97. Female, [R/V ELTANtn, sta. 695].

93. Anterior view of animal. 94. Head and penis. 95. Right lateral, 96. left lateral, and 97. ventral views of anterior alimentary

system. Scale bars =

5 mm. gL, gland of Leiblein; Isg, left salivary gland; nr, cireumoesophageal nerve ring; oe, oesophagus; p,

penis; poe, posterior oesophagus; pr, proboscis; prp, propodium; prr, proboscis retractors; rsg, right salivary gland; s, siphon; sp,

seminal papilla; tn, cephalic tentacles.

rows below bending plane in sublingual pouch, posteri-

ormost 7 rows nascent. Radula strongly sclerotized, dark

yellow. Rachidian teeth (Figures 88, 89) long (~185

um), very narrow (~75 wm), with anteriorly arched, lat-

erally indented basal plate, 3 stout, radially oriented

cusps, central cusp longest. Lateral teeth (Figures 90-

92) with long, stout basal plates attached at acute angle

(~22-28°) to axis of radular ribbon, with 3-4 robust,

roughly parallel cusps of similar length emanating from

innermost ~% of basal plate, outermost ~% of basal

plate beneath cusps of adjacent lateral tooth. Teeth along

bending plane (Figure 91) badly worn. Salivary glands

(Figures 95-97, rsg, Ise) large, seemingly fused, acinous.

Right salivary gland (Figures 95, 97, 188) completely cov-

Page 24

THE NAUTILUS, Vol. 118, No. 1

ee 3

~|60°s

40°W

30°W

1234

No. of specimens

uy type locality

@® examined

material

Figure 98. Geographic and bathymetric distributions of Germonea rachelae new species.

ers valve of Leiblein, lies dorsal to nerve ring, flanks

right, ventral, anterior part of proboscis sheath. Left sal-

ivary gland (Figures 96, 97, Isg) smaller than right. Sal-

ivary ducts short, thick, Iewornne embedded in “esopha-

geal wall shortly after leaving salivary glands. Valve of

Leiblein large, well defined, pyrifor m, Beommeath in pre-

served specimens, with ciliary cone, whitish glandular

pad visible through walls of valve. Gland of Leiblein

(Figures 95, 96, oL) large, bulky, brownish, envelopes

pr oboscis ventro- laterally, extends posteriorly along oe-

sophagus, opens into oesophagus via narrow dine well

posterior to small circumoesophageal nerve ring (Fig-

ures 96, 97, nr). Anterior, mid-, and posterior oesopha-

gus thick, broad, narrowing slightly when passing

through nerve ring. Stomach unknown for holotype.

Non-type specimen (2) had portion of stomach pre-

served, indicating stomach U-shaped, lacking posterior

mixing area.

Male Reproductive System (Figures 93-94): Sperm

duct runs anteriorly along right side of body to enter

Table 4. Germonea rachelae new species. Measurements of shell characters. Linear measurements in mm. (n =

base of long, narrow, nearly cylindrical penis (Figure 94)

that extends length of mantle cavity. Seminal papilla

(Figure 94, sp) very short, surrounded by circular fold.

Type Locality: Off South Georgia Island, 58°02’ S

37°57’ W, in 3197-3239 m. [R/V ELTANIN cruise 9, sta.

698, 30 Aug, 1963].

Type Material: Holotype, ¢, USNM 896594, from

the type locality; Paratype 1, ¢, USNM 1010620, N of

South Orkney Islands, 58°06’ S, 44°56’ W, in 2800 m.

[R/V ELTANIN, cruise 7, sta. 480, 15 Feb 1963]; Paratype

2, USNM 898828, off South Georgia Island, 55°02’ S

44°21’ W, in 3623-3714 m. [R/V ELTANIN cruise 7, sta.

469, 12 Feb 1963]; Paratype 3, USNM 1010621, off

South Georgia Island, 54°59’ S, 38°13’ W, in 2379-2196

m. [R/V ELTANIN cruise 9, sta. 686, 25 Aug 1963].

Other Material Examined: USNM 1013061, off

South Georgia Island, 56°53’ S, 37°33’ W, 3144-3138 m.

[R/V ELTANIN cruise 9, sta. 695, 28 Aug 1963], 2 frag-

4, including

holotype).

Character Mean o Range Holotype

Shell length (SL) 57.0 111 41.7-67.8 61.8

Final whorl length (FWL) 41.5 8.0 30.2-47.9 46.1

Aperture length (AL) 30.4 5.8 99,.9-35.5 S155)

Shell width (SW) 24.7 5.3 18.2-30.3 30.3

FWL/SL 0.73 0.02 0.71-0.75 0.75

AL/SL 0.54 0.03 0.49-0.57 0.57

SW/SL 0.43 0.04 0.40-0.49 0.49

Number of spiral cords on penultimate whorl 24.5 5.2 18-30 30

Number of spiral cords on final whorl 58.0 14.4 48-S1 81

M. G. Harasewych and Y. I. Kantor, 2004

mentary bodies (12 + 16) without shells (anatomy ex-

amined).

Distribution (Figure 98): This species is known only

from the abyssal plain of the Scotia Sea, at depths of

2196-3714 m.

Etymology: The species is named after Mrs. Rachel

(Raye) N. Germon, in recognition of her years of work

in managing the collections of Antarctic Mollusca at the

National Museum of Natural History, Smithsonian In-

stitution.

Remarks: Germonea rachelae may be easily identified

by its large, fusiform shell, with short siphonal canal and

deep amber periostracum. Of the few Antarctic species

that reach comparable size, only Cavineptunea monstro-

sa Powell, 1951, from outer continental shelf depths off

South Georgia Island, is superficially similar, but differs

in having a unique, cylindrical protoconch with indented

apex, a broader aperture, and in lacking the axial rugae

of Germonea near the suture.

The radula of Germonea rachelae, with its narrow, tri-

cuspid rachidian teeth, and lateral teeth in which the 3—

4 cusps are concentrated along the inner half of the basal

plate, has clear affinities with several genera assigned to

the subfamily Prosiphiinae (Appendix 1). It is most sim-

ilar to the radula of several species of Prosipho |i.e., Pro-

sipho spiralis Thiele, 1912, Numanami, 1996: figs.

116D-E; Prosipho crassicostatus (Melvill and Standen,

1907), see Hain, 1989: pl. 24, fig. 1] and Anomacme smi-

thi Strebel, 1905 (Powell, 1951: fig. K57). The shells of

these sublittoral to bathyal genera, while similar in gen-

eral shape and proportion to those of Gemonea, rarely

exceed 10 mm in length. The rachidian teeth of Ger-

monea rachelae are far narrower, more elongated, and

deeply indented anteriorly (producing a tooth with a Y-

shaped outline) than any species of Prosiphiinae. While

the opercula of relatively few Prosiphiinae are illustrat-

ed, Numanami (1996: fig. 110) shows the operculum of

Prosipho hunteri Hedley, 1916 to be elongated, slightly

smaller than the aperture, with a round attachment area

and a terminal nucleus. The distal, free portion of the

operculum, containing the terminal nucleus, is not as

strongly curved as it is in Germonea rachelae.

Subfamily Cominellinae Gray, 1857

Lusitromina new genus

Type Species: Tromina abyssorum Lus, 1993, by orig-

inal designation.

Description: Protoconch increasing in diameter from

470 pm to 2.2 mm in 2 whorls. First whorl smooth,

followed by onset of fine, sharp axial lamellae, then, %

whorl later, by broad, shallow spiral cords. Transition to

teleoconch distinct, marked by onset of coarser sculp-

ture. Teleoconch of moderate size (to 29.2 mm), up to

3% whorls, with strong shoulder on first 2 whorls, in-

creasingly rounded thereafter. Axial sculpture prominent

on protoconch, reduced on first teleoconch whorl, lim-

Page 25

ited to fine growth lines on subsequent whorls. Spiral

sculpture of broad cords that increase in number, but

decrease in prominence with increasing shell size. Ap-

erture large, ovate, columella long, axial, with short si-

phonal fold. Neither siphonal fasciole nor pseudoumbil-

icus present. Periostracum of fine axial lamellae. Oper-

culum D-shaped, paucispiral, large. Eyes absent. Kidney

narrow. Foot with pronounced propodium. Buccal mass

larger than retracted proboscis. Radula with tricuspid

rachidian teeth, with central cusp longer, wider that

flanking cusps, basal plate broadly rectangular. Lateral

teeth with two long, sharp, curved cusps that join above

basal plate. Stomach small, U-shaped, without posterior

mixing area. Penis with long, cylindrical papilla sur-

rounded by fold around base.

Etymology: The genus is named after the late Dr. Val-

entina Lus of the P.P. Shirsov Institute of Oceanology,

Russian Academy of Sciences, Moscow, for her work on

abyssal and hadal Buccinoidea.

Remarks: Dall (1918) proposed the genus Tromina

without discussion, designating as type species Fusus

unicarinatus Philippi, 1868, (from subtidal depths of the

Magellanic Province), a taxon that he had previously

(Dall, 1902: 536) questionably included in the genus

“?Trophon”. Later, Dall (1919: 336) commented that this

genus “has the nucleus [protoconch] of a Trophon and

is probably related to the austral Trophons, but the soft

parts are not known.” Later still, Dall (1925: 28, plate

21, fig. 7) illustrated the type species.

Several species from the South Atlantic have since

been attributed to the genus Tromina, including the sub-

littoral to bathyal Magellanic species: T. fenestrata Pow-

ell, 1951, T. simplex Powell, 1951, T. tricarinata Powell,

1951, and T. bella Powell, 1951; the abyssal T. bella abys-

sicola Clarke, 1961, from the Cape Basin off SW Africa,

and T. traversiensis Clarke, 1961, from off the South

Sandwich Islands, and the hadal T. abyssorum Lus, 1993,

from the Orkney Trench. Clarke (1961, 1962) also rec-

ognized, but did not name three additional abyssal taxa,

referring to them as Tromina a, b, c.

Cernohorsky (1977: 110) was the first to note that Fu-

sus unicarinatus Philippi, 1868 was preoccupied by Fu-

sus unicarinatus Deshayes, 1835. He considered Trom-

ina tricarinata to be conspecific with F unicarinatus

Philippi, and suggested that it might serve as a replace-

ment name for the type species. Bouchet and Warén

(1985: fig. 328, 330) subsequently illustrated the shell

(fig. 328) and radula (fig. 330) of T. wnicarinatus, con-

firming Dall’s (1919) belief that this taxon was a muricid

closely “related to the Austral Trophons.” Since the orig-

inal description of Tromina tricarinata includes an illus-

tration of its radula that clearly shows this taxon to be a

buccinoidean, it could not serve as a replacement name

for a species of muricid.

Dell (1990: 208) reviewed the nomenclatural history

of Tromina and its type species, and proposed Tromina

dispectata as a new name for Fusus unicarinatus Philip-

pi, 1868, non Deshayes, 1835. Recognizing that several

Page 26

buccinoidean taxa previously described in Tromina re-

quired a new generic allocation, he proposed the new

genus Falsitromina (type species: Tromina bella Powell,

1951), characterized by a distinctive, complex proto-

conch morphology, small (to 15 mm) shell, paucispiral

operculum, and radula with tricuspid rachidian teeth and

lateral teeth with two cusps set close together. In addi-

tion to the type species, he included F. simplex, F. tri-

carinata, F. fenestrata, and proposed an additional spe-

cies F. powelli, all from bathyal depths of the Magellanic

Province, but did not include or mention Clarke’s (1961)

abyssal species.

The bathyal genus Antarctodomus Dell, 1972 (type

species: Bathydomus thielei Powell, 1958) shares the un-

usual protoconch morphology (Numanami, 1996: fig.

96C) and obesely fusiform shell shape (Numanami,

1996: fig. 96A-B) of Falsitromina, but differs in reaching

a much larger size (to 35 mm, Dell, 1990: 169), in having

a triangular, sharply tapering operculum (Arnaud, 1972:

fig. 20B), and in having a radula with tricuspid rachidian

teeth that have a more narrowly rectangular basal plate

with a broadly indented anterior edge, and bicuspid lat-

eral teeth with both cusps appearing blunt and nearly

cylindrical (Arnaud, 1972: fig. 20C; Numanami, 1996:

fig. 96D).

The new genus Lusitromina is proposed to contain

the abyssal and hadal taxa that were originally described

in Tromina, namely T. abyssicola Clarke, 1961 (as T. bel-

la abyssicola), and T. abyssorum Lus, 1993. Tromina

traversiensis Clarke, 1961, was shown to be a cancellar-

iid of the genus Iphinopsis Dall, 1924, by Bouchet and

Warén (1985: 261).

Lusitromina shares many of the distinctive features

characteristic of both Falsitromina and Antarctodomus,

including the complex protoconch morphology, obesely

fusiform shell shape, and radulae with tricuspid rachi-

dian teeth and bicuspid lateral teeth. However, the

rounded paucispiral operculum of Lusitromina serves to

distinguish it from Antarctodomus, which has a tapering,

triangular operculum. The rachidian teeth of Lusitrom-

ina, while tricuspid, differ from those of both Falsitrom-

ina and Antarctodomus in have a central cusp that is

larger than, rather than equal in size to the flanking

cusps. The basal plate of the rachidian tooth is broadly

rectangular, intermediate between the squarish basal

plate of Falsitromina, and the very narrow and deeply

indented basal plate of Antarctodomus. The lateral teeth

of Lusitromina have two sharp, curved cusps of equal

size that fuse above the basal plate, while the cusps of

both Falsitromina and Antarctodomus are not equal in

length or width.

Lusitromina abyssorum (Lus, 1993)

(Figures 99-137, Table 5)

Synonymy—Tromina abyssorum Lus, 1993:178

Description: Shell (Figures 99-100, 103-109) of

moderate size (to 29.2 mm), thin, small specimens trans-

THE NAUTILUS, Vol. 118, No. 1

lucent, largest specimens opaque along last whorl. Shell

shouldered, biconical when small, ovate when large. Pro-

toconch well preserved, especially in small specimens

(Figures 115-117). Protoconch increasing from 470 pm

to 2.2 mm in diameter in 2 whorls, first 0.9 whorl

smooth, slightly pitted, followed by onset of fine, sharp

axial threads, and % later, by broader spiral cords. Tran-

sition to teleoconch distinct (Figures 116, 117 arrows),

followed by onset of coarser axial sculpture. Teleoconch

of up to 3% whorls. Shoulder pronounced on first 2

whorls, marked by thick spiral cord, producing a stepped

spire, becoming progressively less conspicuous in sub-

sequent whorls, entirely rounded following third whorl

(Figures 99-100). Axial sculpture of strong, evenly

spaced cords dominant on protoconch, becoming more

widely spaced, less prominent following transition to te-

leoconch, disappearing entirely after first 4% teleoconch

whorl. Subsequent axial sculpture confined to very fine,

weakly prosocline growth lines. Spiral sculpture of broad

cords that usually increase in number, but decrease in

prominence with increasing shell size, 4-20 on penulti-

mate whorl, 2-20 between suture and shoulder, 19-53

on last whorl. Aperture large (AL/SL = 0.62-0.66),

broadly oval, deflected from shell axis by 24-27°. Outer

lip very thin, not reflected, evenly rounded from shoul-

der to base of short, broad, axial siphonal canal. Shell

composed of three layers (Figure 121), outermost layer

(~2 wm) of columnar calcitic crystals, middle layer,

thickest (~26 jm) of collabrally oriented crossed-la-

mellar crystals, innermost layer (~4 jm) of crossed la-

mellar crystals oriented perpendicular to middle layer.

Inner lip of wide, weakly convex parietal region, straight,

weakly indented axial columellar region of equal length,

with long siphonal fold defining abaxial portion of si-

phonal canal. Neither siphonal fasciole nor pseudoum-

bilicus present. Shell color uniformly white. Periostrac-

um (Figure 120) very thin, straw yellow in color, with

extremely fine axial lamellae. Operculum (Figures 101—

102, 118-119) D-shaped, large, spanning ~0.61 AL,

thin, yellowish, broadly ovate, paucispiral, with nucleus

near adaxial, abapical edge (Figures 118-119).

Anatomy (Figures 122-124): Soft tissues comprise

approximately 3% whorls. Mantle cavity spans just under

¥% whorl, kidney narrow, spans about 1/7 whorl. Colu-

mellar muscle short, comprising slightly less than 1

whorl, attaching to shell at rear of mantle cavity. Foot

large, long (L/W ~2.2), with well developed, crescent-

shaped propodium. Propodial cleft very wide, containing

conspicuous propodial gland with rounded opening (Fig-

ures 123, 124, pg). Body color yellowish tan, without

pigmentation pattern. Kidney, digestive system, testis all

greenish. Head large, with long, conical tentacles (Fig-

ure 123, tn), without discernable neck. Eyes absent. Ne-

phridium brown, folds lighter in color, visible through

wall. Mantle covers base of head and most of penis. Peri-

cardium (Figure 122, per) ventral to narrow nephridial

gland.

Mantle Cavity (Figure 128): Mantle cavity short,

Page 27

M. G. Harasewych and Y. I. Kantor, 2004

Figures 99-109. Shells and opercula of Lusitromina abyssorum (Lus, 1993). 99-102. USNM 1010536, off Saunders Island,

South Sandwich Islands, 57°39'00" S, 26°00'24" W, in 2380-2609 m [R/V Istas Orcapas, sta. 54]. 99. Apertural, and 100. dorsal

views of shell. 101. Outer, and 102. inner views of eperculum. 103. Apertural view, USNM 896533, off South Georgia Island,

56°04’ S, 33°59’ W, in 3138-3239 m [R/V Exranin, sta. 722]. 104. Apertural view, USNM 896785, N of South Orkney Islands,

60°06" S, 45°26’ W, depth not recorded (estimated > 5000 m), [R/V Exranin, sta. 488]. 105. Apertural and 106. Dorsal views,

USNM 1010535, E of Candlemas Island, South Sandwich Islands, 57°00'24” S, 26°10'06” W, in 2740-2757 m [R/V IsLas Orcapas,

sta. 38]. 107. Apertural, 108. dorsal, and 109. apical views, USNM 896525, N of South Georgia Island, 53°02’ S, 37°40’ W, in

3056-3102 m [R/V ELTantn, sta. 735].

Page 28

THE NAUTILUS, Vol. 118, No. 1

Figures 110-114. Radulae of Lusitromina abyssorum (Lus, 1993). 110. Dorsal, and 111. Right lateral (30°) views of the central

portion of the radular ribbon of specimen in Figures 99-100, USNM 1010536. 112-113. Dorsal views of the radular ribbon of

specimen in Figures 107-109, USNM 896525. 112. Dorsal view of central portion of radula. 113. Anteriormost part of radular

ribbon, showing wom teeth. 114. Enlarged right lateral teeth, showing bifurcation of outer cusp.

broad (L/W ~0.6). Mantle edge smooth. Siphon very

short, broad, muscular, extending slightly beyond mantle

edge. Osphradium (Figure 128, os) yellowish, bipectin-

ate, large (~% mantle cavity length), very wide, with

wide, curved osphradial nerve. Ctenidium (Figure 128,

ct) large, narrower than osphradium, strongly curved,

spanning nearly entire mantle cavity length. Shape of

ctenidium lamellae varies with position along organ. La-

mellae tall, triangular posteriorly (Figure 129), gradually

becoming narrower anteriorly (Figure 130). Hypobran-

chial gland without distinct folds, aovenad by thick layer

of mucus. Rectum short, spans <¥ mantle cavity length.

Anus terminal, without papilla.

Alimentary System (Figures 131-136): Proboscis

(133, pr) short when retracted (~0.36 AL), not thick (L/

D ~3.3), smooth, non-pigmented. Proboscis retractors

(Figures 133, 134, prr) not numerous, but thick, pow-

erful, attached to proboscis sheath at mid-length, when

proboscis retracted. Proboscis sheath very thin-walled,

anterior half translucent, thickened posteriorly to be-

come as thick as proboscis wall. Proboscis wall thin

~1/10 proboscis diameter. Mouth opening triangular

slit. Buccal mass muscular, large, filling retracted pro-

boscis, slightly protruding from its posterior end (Figure

136, bm). Odontophoral cartilages paired, fused anteri-

orly, but connected only by very thin layer of tissue,

comprise ~%4 of buccal mass length. Radular ribbon

equal in length to retracted proboscis (4.3-5.0 mm,

0.26-0.28 AL), 300-330 jum wide, triserial (Figures 110—

114), with 65-76 rows of teeth, posteriormost 5-6 rows

M. G. Harasewych and Y. I. Kantor, 2004

Page 29

Figures 115-121.

Lusitromina abyssorum (Lus, 1993). 115. Apertural, 116. apical, and 117. latero-dorsal views of the shell of

a juvenile specimen, USNM 896800, Scotia Ridge, 56°02’ S, 61°56’ W, in 4008 m, [R/V Eltanin, sta. 112]. Arrows indicate transition

from protoconch to teleoconch. 118-119. Operculum of the same specimen. 118. Outer surface of operculum. 119. Enlarged

nucleus of operculum. 120. Periostracum, and 121. shell ultrastructure of specimen shown in Figures 107-109, USNM 896525.

nascent. Rachidian teeth with 3 cusps on posterior por-

tion of broad, anteriorly arched basal plate. Central cusp

slightly longer, wider than lateral cusps. Lateral teeth

with 2 cusps of approximately equal length. Outer cusp

with bifurcated tip in one specimen (Figure 114). Sali-

vary g glands (Figures 133-135, Isg, rs¢) small, not fused,

acinous. Right salivary gland completely covers valve of

Leiblein, lies dorsally to the nervous ring (Figures 134,

135, rsg). Left salivary gland more rounded, situated lat-

erally, just posterior to the nerve ring (Figure 133, 134,

Isg). Salivary ducts (Figure 134, 136, sd) short, thick,

become embedded in oesophageal wall shortly after

leaving the gland. Valve of Leiblein (Figure 136, vL) well

defined, large, pyriform, with ciliary cone, whitish glan-

dular pad visible through walls of valve. Gland of Lei-

blein (Figures 133, 134, 136, gL) small, yellowish, short,

tubular, not coiled, opening into oesophagus via short

duct (Figure 136, dgL) slightly posterior to the nerve

ring. Oesophagus thick, broad anterior to nerve ring,

narrower posterior to nerve ring (Figures 134, 135, poe),

a not widen before entering stomach (Figures 131,

2, poe). Stomach (Figures ‘131, st; 132) “waielll U-

Page 30 THE NAUTILUS, Vol. 118, No. 1

—- 128

Figures 122-130. Anatomy of Lusitromina abyssorum (Lus, 1993), specimen shown in Figures 107-109, USNM 896525. 122—

123. Lateral views of the animal removed from shell. 124. Anterior view of the foot showing the pedal gland. 125. Cephalic

haemocoel, with organs of the digestive system removed to show the enlarged seminal duct. 126. Dorsal view of penis. 127. Seminal

vesicle. 128. Mantle cavity organs. 129-130. Shape of ctenidium lamellae in different regions of ctenidium. Scale bars = 2 mm.

cme, cut mantle edge; ct, ctenidium; dg, digestive gland; hg, hypobranchial gland; nep, nephridium; op, operculum; os, osphra-

dium; p, penis; pe, propodium; per, pericardium; pg, propodial gland; re, rectum; rhs, rhynchostome; s, siphon; sdt, glandular

seminal duct (prostate gland); sem.d, seminal duct; st, stomach; tes, testis; tn, cephalic tentacles.

M. G. Harasewych and Y. I. Kantor, 2004 Page 31

Sea . \ Sa g

7 NS Se AN WENN

i eo a AVR NY WO =

Psy

135

Figures 131-136. Anatomy of digestive system of Lusitromina abyssorum (Lus, 1993). 131-132. Specimen shown in Figures 107—

109, USNM 896525. 131. Dorsal view of stomach, embedded in digestive gland. 132. Stomach, opened dorsally along incision shown

by dashed line in Figure 131. 133-136. Specimen shown in Figures 99-100, USNM 1010536. 133. Left lateral, 134. ventral, and

135. right lateral views of anterior alimentary system. 136. Latero-dorsal view of the anterior oesophagus to show the opening of the

duct of gland of Leiblein. Scale bars = 2 mm for Figures 131, 133-135, 1 mm for Figures 132, 136. a, anus; addg, anterior duct of

the digestive gland; ao, anterior aorta; bm, buccal mass; dg, digestive gland; dg, duct of gland of Leiblein; gL, gland of Leiblein;

If, longitudinal fold; Is, lateral sulcus; Isg, left salivary gland; nep, nephridium; ng, nephridial gland; nr, cireumoesophageal nerve

ring; pddg, posterior duct of digestive gland; per, pericardium; poe, posterior oesophagus; pr, proboscis; prr, proboscis retractors;

rsg, right salivary gland; sd, salivary duct; ss, style sac; st, stomach; T1, T2, major and minor typhlosoles; vL, valve of Leiblein.

Page 32

THE NAUTILUS, Vol. 118, No. 1

2000

50° 2200

2400

2600

2800

3000 |

3200

tie Peninsula

50°W

Figure 137.

shaped, without posterior mixing area, with numerous,

fine, semicircular dorso-ventral folds. Transition from

oesophagus to stomach marked by change from tall lon-

gitudinal folds to much finer, curved folds that line stom-

ach. Posterior duct of digestive gland (Figure 132, pddg)

large, situated near entrance of oesophagus. Longitudi-

well (ol (Figure 132, If) narrow, distinct, originates at

entrance of oesophagus, runs length of stommachn, be-

comes obsolete near intestine. Antietor duct of digestive

gland (Figure 132, addg) small. Lateral sulcus (Figure

132, Is) shallow. Typhlosoles (Figure 132, T1, T2) more

prominent in intestine, bordering, deep, narrow intesti-

nal groove. Posterior part of style sac (Figure 132, ss)

lined with transverse folds. Folds lining stomach very

low, ventral channel of gastric chamber poorly separated

from dorsal chamber.

Male Reproductive System: Seminal vesicle of me-

dium size (Figure 127), spans less than % of whorl,

formed of few large loops. Seminal duct descends to

floor of mantle cavity at its rear, becomes thickened, sin-

40°W

us type locality

& examined material

4000

= 4200

i=)

5000

30°W

5200

5400

600

No. of specimens 2 4 6 8 10

seographic and bathymetric distribution of Lusitromina abyssorum (Lus, 1993).

uous, running to base of penis, where it enters cephalic

haemocoel (Figure 125) expands greatly, makes a long

loop posteriorly, before entering penis. Penis shorter

than mantle cavity, flattened, non-pigmented, with long,

cylindrical terminal papilla surrounded by circular fold

at its base (Figure 126).

Type Locality: Lorie Trench, off South Orkney Is-

lands, 60°12'9” S, 43°59'0" W, in 5450-5480 m. [R/V

AKADEMIK KurcHaToy, sta. 909, 10 Dec 1971].

Type Material: Holotype and 2 paratypes, from the

type locality; 2 paratypes, Orkney Trench, 60°50'5” S

41°11'7" W, in 5063-5470 m, [R/V Dorirriy MENDED

Ey, sta. 4089, 4 Mar, 1989]. Type material could not be

located in the collections of the Institute of Oceanology

of Russian Academy of Sciences, Moscow. The type se-

ries consists entirely of juvenile specimens. The shell of

the holotype (the only intact shell) measured 7.4 mm in

length, 5.5 mm in width.

Other Material Examined: SOUTH GEORGIA IS-

Table 5. Lusitromina abyssorum (Lus, 1993). Measurements of shell characters. Linear measurements in mm. (n = 5).

Character

Shell length (SL)

Final whorl length (FWL)

Aperture length (AL)

Siphonal canal length (SCL)

Shell width (SW)

FWL/SL

AL/SL

SCL/SL

SW/SL

Number of spiral cords on penultimate whorl

Number of spiral cords suture to peripheral cord

Number of spiral cords peripheral cord to siphon

Mean o Range

20.9 6.5 12.3-29.2

16.7 49 10.3-23.2

13.4 4] §.6-18.9

2.5 0.4 1.9-3.3

13.4 42, 7.9-19.4

0.80 0.02 0.78—0.84

0.65 0.03 0.61—0.70

0.12 0.02 0.10-0.16

0.64 0.04 0.60-0.70

12.8 6.4 4-20

9.6 7.4 2-2()

35.0 13.1 19-53

M. G. Harasewych and Y. I. Kantor, 2004

LAND: USNM 896525, 53°02’ S, 37°40’ W, in 3056—

3102 m. [R/V ELTANIN, sta. 735, 13 Sep 1963], 6 spec-

imens (1 adult + 5 juveniles), (anatomy examined);

USNM 896533, 56°04’ S, 33°59’ W, in 3138-3239 m,

[R/V Evranin, sta. 722, 8 Sep 1963], 1 specimen.

SOUTH SANDWICH ISLANDS: USNM 1010535, E

of Candlemas Island, 57°00'24” S, 26°10'06” W, in 2740—

2757 m. [R/V IsLas Orcapas, sta. 38, 22 May 1975], 1

specimen; USNM 1010536, Saunders Island, 57°39’00"

S, 26°00'24” W, in 2380-2609 m. [R/V IsLas Orcapas,

sta. 54, 27 May 1975] 1 specimen (radula and anterior

foregut anatomy studied), SCOTIA RIDGE: USNM

896800, 56°02’ S, 61°56’ W, in 4008 m. [sta. 112, 20 Jul

1962], 4 specimens (juveniles). SOUTH ORKNEY IS-

LANDS: USNM 896785, 60°06’ S, 45°26’ W, depth not

recorded, but > 5000 m according to the bathymetric

Atlas GEBSCO97], [R/V Exranin, sta. 488, 18 Feb

1963] 1 specimen; USNM 1010537, 60°07’ S, 45°14’ W,

5285 m. [R/V ELTANIN, sta. 485, 18 Feb 1963] 8 speci-

mens (juveniles).

Distribution (Figure 137):

margins of the Scotia Sea, in 2

Within and around the

2380-5480 m.

Remarks: Lus’s (1993) original description of Tromina

abyssorum is undoubtedly the most extensive of any

Antarctic or abyssal buccinoidean in terms of concho-

logical and anatomical detail. She was familiar with

Clarke’s (1961) work on the abyssal mollusks from the

South Atlantic Ocean, and regarded T. abyssorum to be

closely related to T. bella abyssicola Clarke, 1961. She

was apparently unfamiliar with Dell’s (1990) clarification

of the taxonomic affinities of Tromina, as discussed in

the remarks under Lusitromina (above).

DISCUSSION

While exploration of Antarctic Seas began during the

second half of the eighteenth century (Conrad, 1999 for

chronology), significant additions to our knowledge of

their molluscan fauna originated with the cruises of the

H.M.S. CHaLLeNcerR (Numanami, 1996: Table 1) and

continue to this day. Several of the research cruises dur-

ing the late 19th and early 20th centuries sampled at

ledst some stations at abyssal depths, yet only a single

buccinoidean genus (Bathydomus Thiele, 1912) had

been described from these depths off Antarctica. Clarke

(1961) reported on the abyssal mollusks collected during

the cruises of the R/V VeMa in the South Atlantic (1957—

1958). He recognized five buccinoidean taxa, but attri-

buted them all to the subtidal genus Tromina. Following

his example, Lus (1993) described a related abyssal to

hadal species in the genus Tromina, trawled by the Rus-

sian vessels R/V AKADEMIK Kurcuaroy (1971) and R/V

Dmitriy MENDELEEV (1989) during their explorations of

the abyssal benthic faunas of the Weddell and Scotia

Seas. The present study, based on material collected by

the United States Antarctic Program (USAP) vessels

R/V Istas Orcapas, R/V ELTANIN and supplemented by

samples from the German vessel R/V POLARSTERN, has

Page 33

increased the number of abyssal buccinoidean genera

from the seas surrounding Antarctica to six. ;

In the course of our study, we have reviewed the tax-

onomic placement, geographic and bathymetric distri-

bution, diagnostic characters, and diversity of each of the

29 genera that have thus far been proposed for Antarctic

ancl Magellanic Buccinoidea (Appendix 1). The bathy-

metric ranges of these genera are plotted in Figure 138.

We analysed the bathymetric distribution of these taxa

by subdividing depth into 200 meter increments, and

scoring each genus as present or absent within each in-

crement. Employing the Correlation Distance Measure

and Ward’s Method for Group Linkage, PC-ORD

(McCune and Mifford, 1999) was used to cluster the

depth increments based on similarities of their bucci-

noidean fauna. The resulting dendrogram (Figure 138,

Ward’s Method) reveals a primary andl profound dichot-

omy in generic composition of the faunas at depths

above and! below 2200 m, roughly corresponding to the

transition from the bathyal to the abyssal zone (Gage and

Tyler, 1991). This differentiation of slope and abyssal

faunas is absolute, with the two faunas having no genera

in common. The apparent minor overlap in the ranges

of Parabuccinum and Spikebuccinum shown in Figure

138 is spurious, as all records for Parabuccinum from

depths greater than 866 m are based on dead shells

(Harasewych, Kantor and Linse, 2000). On a finer scale,

the continental shelf (0-200 m: Fi igure 138: A) and up-

per continental slope (200-1,000 m; Figure 138: B) fau-

nas are differentiated from the lower slope fauna (1000—

2200 m; Figure 138: C), while the abyssal fauma is par-

titioned into continental rise (2200-3800 m; F igure 138:

D), abyssal plain (3800-4800 m; Figure 138: E), and

hadal (>4800 m: Figure 138: F) faunas. The pattern of

clustering agrees generally with that obtained by Rex

(1977: fig. 1) for the deep-sea gastropod fauna of the

western North Atlantic, except that, for Antarctic and

Magellanic Buccinoidea, the upper continental slope

fauna is more similar to the continental shelf fauna than

to the lower continental slope fauna.

When assessing the taxonomic placement of genera,

we observed that those genera occurring at continental

shelf and slope depths conform readily to Powell: s (1951)

criteria for the subfamilies Buccinulinae, Prosiphiinae

and Cominellinae. However, taxa from abyssal and hadal

depths do not. It is interesting to note that Powell (1951:

131) had to modify his criteria for Buccinulinae to ac-

commodate Bathydomus, the only abyssal genus previ-

ously included in his classification. We admit to having

similarly modified or expanded subfamilial criteria in or-

der to fit our newly described abyssal taxa into an exist-

ing classification for the sake of taxonomic expediency.

While the sublittoral and bathyal buccinoideans of Ant-

arctica and the Magellanic Province are likely the prod-

uct of one, or very few in sitw evolutionary radiations,

the origins and relationships of the abyssal fauna are

more complex. For some abyssal genera (e.g., Spikebuc-

cinum, Germonea, Lusitromina), credible sister taxa in-

habit adjacent continental slopes. For others (e.g., Dre-

Page 34

THE NAUTILUS, Vol. 118, No. 1

Buccinulinae

= £ LZ

— S BSB E& 5S BS

Ward's @ =5528238558

S Sa

BEssssesessae

Method SBSESsSSER SS

Seta ShSGSFLBSA

Prosiphiinae Cominellinae

= a

Ea~j

o = S&S = =>

= 2 ee Se = S

SEs5sS22888 SESSESEE

ao 8S OB2Z2Fae Les = [0 = Ss

SSe5 5558585 se5azess

Sesce2X Sesrtaus DS Co oS = se S

Qe ereetsscggpFtZs et tote ezs

Figure 138.

Bathymetric distributions of buccinoidean genera occurring in the Magellanic Province and south of the Antarctic

convergence. The dendrogram clusters 200 m increments of depth on the basis of shared buccinoidean genera. A, continental shelf;

B, upper continental slope; C, lower continental slope; D, continental rise; E, abyssal plain, F; hadal depths.

Depth (m)

(eo) 5 10

Number of Genera

Figure 139. Diversity of buccinoidean genera vs. depth. A,

continental shelf; B, upper continental slope; C lower conti-

nental slope; D, continental rise; E, abyssal plain; F, hadal

depths.

panodontus, Muffinbuccinum), we have noted closer af-

finities with abyssal or hadal taxa from austral seas be-

yond the Antarctic convergence. Lipps and Hickman

(1982) reviewed various hypotheses regarding origin and

age of deep-sea and Antarctic faunas. They concluded

that both faunas have evolved primarily in place since at

least the Mesozoic, but were supplemented by subse-

quent minor immigration. Our findings are qualitatively

consistant with this hypothesis, indicating that, at least

for the Buccinoidea, which have origins in the late Me-

sozoic, the abyssal Antarctic fauna consists of a combi-

nation of taxa, some with nearest relatives, and presum-

ably origins, on the adjacent continental slope, and oth-

ers with nearest relatives in neighboring ocean basins.

The diversities of both the slope and the abyssal buc-

cinoidean faunas decrease with increasing depth (Figure

139). While the shallower components of each fauna

tend to have narrow bathymetric ranges and the deeper

dwelling genera tend to have broader bathymetric rang-

es, most genera span two or three bathymetric regions.

Only 4 of 19 sublittoral genera (21.1%) are confined to

the continental shelf, while 3 of 16 (18.8%) upper slope

genera and 0 of 4 lower slope genera (0%) are limited

to a single region. Within the abyssal zone, 2 of 5 genera

(40.0%) are known only from the continental rise, while

none of the 3 genera that occur on the abyssal plain are

restricted to it. Of the 3 hadal genera, only one (33.3%),

Muffinbuccinum, is restricted to this zone. The genus

Muffinbuccinum is presently known from a single sta-

M. G. Harasewych and Y. I. Kantor, 2004

fe)

200 A

400

600 B

800

1000

1200

1400

1600

1800 C

2000

2200

2400

2600

2800

3000 D

3200

3400

3600

3800

4000

4200 E

4400

4600

4800

5000

5200

5400 F

5600

5800

6000

Depth (m)

(e) 25 sO 7S 100

Percent of Genera Monotypic

Figure 140. Proportion of monotypic genera vs. depth. A

continental shelf; B, upper continental slope; C, lower conti-

nental slope; D, continental rise; E, abyssal plain; F, hadal

depths.

tion. Drepanodontus, which co-occurs with Muffinbuc-

cinum at this station, has a much broader bathymetric

range. For the abyssal buccinoideans, the maximum ge-

neric diversity occurs from 2600 to 3200 m, which close-

ly corresponds to the depths for peak of gastropod spe-

cies richness reported by Rex (1981: fig. 1).

The buccinoidean fauna of the Antarctic and Magel-

lanic Provinces contains an extraordinarily high propor-

tion (14 out of 29, 48.3%) of genera known only from

their type species. In comparison, of 33 genera of boreal

Buccinidae, only 5 (15.2%) are monotypic. When plotted

by depth (Figure 140), it is evident that the abyssal fauna

has a much higher fraction of such genera. The bathy-

metric distribution of monotypic genera at aby ssal

depths in the Southern Ocean appears to be the inverse

of that predicted by a plot of species per genus ratios

against depth (Rex, 1983: fig. 4). Based on samples from

the NW Atlantic Ocean, the number of species per ge-

nus actually peaks between 2000 and 3000 m. Data are

insufficient to determine if this pattern represents a real

biological phenomenon (true monotypy as the result of

an early adaptive radiation into the deep-sea around Ant-

arctica without subsequent diversification), or is an ar-

tefact of low sampling density exacerbated by difficulties

in differentiating closely related species. As noted by

Gage and Tyler (1991: 204), the few large data sets that

are available for deep-sea faunas have produced rarefac-

tion curves that do not even approach their asymptote,

the point at which additional sampling will no longer

discover new taxa. While buccinoidean genera have his-

Depth (m)

10) 25 50 75 100

Maximum Shell size (mm)

Range and Mean

Figure 141. The range and mean of maximum shell size

within a genus vs. depth. A, continental shelf; B, upper con-

tinental slope; C, lower continental slope; D, continental rise;

E, abyssal plain; F, hadal depths.

torically been based on features of the shell, operculum

and radula, species within genera tend to have very sim-

ilar opercula and radulae, and are usually differentiated

primarily on shell morphology. Increases in latitude and

depth each contribute to a reduction in the amount of

calcium carbonate in seawater, which, in turn, has pro-

found effects on shell thickness, geometry and the pres-

ence of sculpture (Graus, 1974). High latitude and deep-

water gastropods often have shells that are thin, roughly

spherical (minimizing the ratio of surface area to vol-

ume), and lack pronounced surface sculpture. There are

numerous examples in the Antarctic fauna of such con-

vergent ecophenotypic similarities in shell form obscur-

ing phylogenetic relationships, even at the level of fam-

‘es and superfamilies (e.g., discussions of Tromina and

Notoficula in Appendix 1; Kantor and Harasewych, 1999,

2000). Some of the genera now regarded as monotypic

will likely be discovered to contain additional cryptic or

sibling species with similar shell and/or radular mor-

phologies when investigated using molecular or multi-

variate morphometric techniques.

A number of authors (e.g., Lipps and Hickman, 1982;

Gage and Tyler, 1999) have noted the prevalence of both

gigantism and small body size in organisms from the

Antarctic and the deep-sea. Rex et al. (1999) reported

that size increases significantly with depth from upper

bathyal region to the abyssal plain for the related neo-

gastropod family Turridae. They suggested that larger

size may be favored at greater depths because of its met-

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THE NAUTILUS, Vol. 118, No. 1

abolic and competitive advantages, and noted that the

density of polychaete worms, a major food source for

Turridae as well as buccinoideans (Taylor, 1978), de-

creases exponentially with depth.

We plotted the range and mean of maximum shell

sizes of all genera occurring within each 200 m bathy-

metric increment (Figure 141). The continental shelf

and upper continental slope support the largest as well

as the smallest buccinoidean genera, with the range in

size narrowing as depth increases. The mean of the max-

imum sizes if genera increases from the continental

shelf to the lower slope, before decreasing abruptly at

the base of the continental slope. The trend of decreas-

ing size range with increasing depth is repeated in the

abyssal fauna, with the largest genera and the broadest

range in size occurring along the continental rise. De-

spite the differences in taxa, ocean basins, and sample

sizes between the work of Rex et al. (1999: fig. 2) and

the present study, both indicate that taxa inhabiting the

base of the continental slope are smaller than those of

either the upper slope or continental rise, and that max-

imum shell size is reached near the boundary of the

continental rise and abyssal plain, and subsequently de-

creases with increasing depth.

ACKNOWLEDGMENTS

We are grateful to Enrico Schwabe and Michael Schrédl

of the Zoological State Collection, Munich, for calling

our attention to, and making available material in their

care that was collected by the R/V Polarstern. We thank

Ms. Yolanda Villacampa for her assistance with scanning

electron microscopy. The thoughtful comments of Dr.

Philippe Bouchet and an anonymous referee are much

appreciated.

This research was supported by a grant from the NSF-

USAP United States Antarctic Program [Contract num-

ber OPP-9509761].

LITERATURE CITED

Arnaud, P. M. 1972. Invertébrés marins des XIJéme et XVeme

expéditions antarctiques frangaises en Terre Adélie 8.—

Gastéropodes Prosobranches. Téthys Supplement 4: 105—

134.

Bamard, K. H. 1963. Deep sea Mollusca from west of Cape

Point, South Africa. Annals of the South African Museum

46: 407-452.

Bouchet, P. and A. Warén. 1985. Revision of the Northeast

Atlantic Bathyal and Abyssal Neogastropoda, excluding

Turridae (Mollusca, Gastropoda). Bolletino Malacologico,

Supplement 1: 121-196.

Bouchet, P. and A. Warén. 1986. Mollusca: Gastropoda: Tax-

onomical notes on tropical deep water Buccinidae with

descriptions of new taxa. Mémoires Muséum national

d Histoire naturelle, Sér A, Zool. 133: 457-499, pls. 1-18.

Brandt, R. A. M. and P. Temcharoen. 1971. The molluscan

fauna of the Mekong at the foci of Schistosomiasis in

South Laos and Cambodia. Archiv fiir Molluskenkunde

101: 111-140.

Cernohorsky, W. O. 1977. The taxonomy of some Southern

Ocean Mollusca (Gastropoda) mainly Antarctic and sub-

antarctic. Records of the Auckland Institute and Museum

14: 105-119.

Clarke, A. H. 1961. Abyssal mollusks from the South Atlantic

Ocean. Bulletin of the Museum of Comparative Zoology

125(12): 345-387, pls. 1-4.

Clarke, A. H. 1962. Annotated List and Bibliography of the

Abyssal Marine Molluses of the World. National Museum

of Canada Bulletin No. 181, pp. vi + 114.

Conrad, L. J. 1999. Bibliography of Antarctic Exploration. Ex-

pedition Accounts from 1768-1960. L. J. Conrad, Wash-

ougal, Washington, xv + 424 pp.

Cossmann, M. 1906. Essais de paléontologie comparée. Liy-

raison 4, Chez LAuteur, Paris. 293 pp., 10 pls.

Dall, W. H. 1902. Illustrations and descriptions of new, unfi-

gured, or imperfectly known shells, chiefly American, in

the U. S. National Museum. Proceedings of the United

States National Museum 24: 499-566, pls. 27-40.

Dall, W. H. 1918. Changes in and additions to molluscan no-

menclature. Proceedings of the Biological Society of

Washington 31: 137— 138.

Dall, W. H. 1919. Descriptions of new species of Mollusca

from the North Pacific Ocean in the collection of the

United States National Museum. Proceedings of the Unit-

ed States National Museum 56(2295): 293-371.

Dall, W. H. 1925. Illustrations of unfigured types of shells in

the collection of the United States National Museum. Pro-

ceedings of the United States National Museum 66(2554):

1-41, pls. 1-36.

Dell, R. K. "9 72. A new genus of Antarctic buccinid gastropod.

Records of the Dominion Museum §(7): 115-119.

Dell, R. K. 1990. Antarctic Mollusca, with special reference to

the fauna of the Ross Sea. Royal Society of New Zealand,

Bulletin 27, 311 pp.

Finlay, H. J. 1928. The Recent Mollusca of the Chatham Is-

lands. Transactions of the New Zealand Institute 59: 232—

286.

Gage, J. D. and P. A. Tyler. 1991. Deep-sea biology, a natural

history of organisms at the deep-sea floor. Cambridge Uni-

versity Press, Cambridge, xvi + 504 pp.

GEBSCO97. 1997. Digital Bathymetric Atlas. British Ocean-

ographic Data Centre, Birkeenhead. CD-ROM.

Graus, R. R. 1974. Latitudinal trends in the shell characteristics

of marine gastropods. Lethaia 7: 303-314.

Habe, T. and J. Sato. 1973. A classification of the family Buc-

cinidae from the North Pacific. Proceeding of the Japa-

nese Society of Systematic Zoology 8: 1-8, pls. 1, 2).

Hain, S. G. 1990. Beitriige zur Biologie der beschalten Mol-

lusken (KI. Gastropoda und Bivalvia) des Weddellmeeres,

Antarktis. Berichte zur Polarforschung 70: 1-181.

Harasewych, M. G., S. L. Adamkewicz, J. A. Blake, D. Saudek,

T. Spriggs and C. J. Bult. 1997. Neogastropod Phylogeny:

A Molecular Perspective. Journal of Molluscan Studies 63:

327-351.

Harasewych, M. G. and Yu. I. Kantor. 1999. A revision of the

Antarctic genus Chlanidota (Gastropoda: Neogastropoda:

Buccinulidae). Proceedings of the Biological Society of

Washington 112: 253-302.

Harasewych, M. G., Yu. I. Kantor and K. Linse. 2000. Para-

buccinum, a new genus of Magellanic Buccinulid (Gastro-

poda: Neogastropoda), with a description of a new species.

Proceedings of the Biological Society of Washington 113:

542-560.

M. G. Harasewych and Y. I. Kantor, 2004

Hutton, F. W. 1880. Manual of the New Zealand Mollusca, a

systematic and descriptive catalogue of the marine and

land shells, and of the soft mollusks and polyzoa of New

Zealand and the adjacent islands. James Hughes, Welling-

ton, pp. xvi + iv + 224.

Kantor, Yu. I. 1996. Phylogeny and relationships of Neogastro-

poda. In: J. D. Taylor (ed.): Origin and evolutionary ra-

diation of Mollusca. Oxford University Press, Oxford, pp.

221-230.

Kantor, Y. I. and M. G. Harasewych. 1999. Rediscovery of the

Antarctic species Sipho g gaint Lamy, 1910 (Gastropoda:

Neogastropoda) with Temmarks on its taxonomic position.

Antarctic Research 11: 431—436.

Kantor, Yu. I. and M. G. Harasewych. 2000. Obscwranella pa-

pyrodes, a new genus and species of abyssal Tonnoidean

Gastropod from Antarctica. The Nautilus 114: 103-111.

Kantor, Yu. I. and R. N. Kilburn. 2001. Rediscovery of Canidia

dorri Wattlebed, 1886, with a discussion of its systematic

position (Gastropoda: Neogastropoda: Nassariidae: Nas-

sodonta). The Nautilus 115: 99-104,

Linse, K. 2002. The Shelled Magellanic Mollusca: with special

reference to biogeographic relations in the Southern

Ocean. Theses Zoologicae 34: vii + 252 pp.

Lipps, J. H. and C. S. Hickman. 1982. Origin, age, and evo-

lution of Antarctic and deep-sea faunas. In: W. G. Ernst

and J. G. Morin (eds). The Environment of the Deep Sea.

Prentice-Hall, Englewood Cliffs, pp. 324-356.

Lus, V. Ya. 1978. New genus and species of Buccinidae (Mol-

lusca: Prosobranchia, Buccinidae) from the lower abyssal

zone of Idzu-Bonin Trench in the Pacific Ocean. Pro-

ceedings of the P. P. Shirsov Institute of Oceanology 113:

147-156, [In Russian, English summary].

Lus, V. Ya. 1989. The deepest gastropods Buccinacea (Neo-

gastropoda: Buccinacea). Proceedings of the P. P. Shirsov

Institute of Oceanology 123: 151-164. [In Russian, En-

glish summary].

Lus, V. Ya. 1993. New species of Tromina (Neogastropoda,

Buccinulidae, Tromina) from low abyssal of Antarctic

trenches, Lorie and Orkney. Proceedings of the P. P. Shir-

sov Institute of Oceanology 127: 176-197. [In Russian,

English summary].

McCune, B. and M. J. Mefford. 1999. PC-ORD, Multivariate

Analysis of Ecological Data, Version 4. MjM Software De-

sign, Gleneden Beach, Oregon.

Numanami, H. 1996. Taxonomic study on Antarctic gastropods

collected by Japanese Antarctic Research Expeditions.

Memoirs of National Institute of Polar Research Series E

(Biology and Medical Science) 39: 1-244.

Oliver, P. G. 1983. Notoficula Thiele, a neotenous genus of

Eratoid gastropod from Antarctica. British Antarctic Sur-

vey Bulletin 61: 1-6.

Olsson, A. A. 1971. Mollusks from the Gulf of Panama col-

lected by the B/V John Elliott Pillsbury, 1967. Bulletin of

Marine Science 21: 35-92.

Ponder, W. F. 1973. A review of the Australian species of Pen-

ion Fischer (Neogastropoda: Buccinidae). Journal of the

Malacological Society of Australia 2: 401-428.

Ponder, W. F. 1974. The origin and evolution of the Neogas-

tropoda. Malacologia 12: 295-338.

Ponder, W. F. 1982. A new genus and species of Buccinidae

(Mollusca: Gastropoda) from the continental slope of east-

em and southern Australia. Journal of the Malacological

Society of Australia 5: 201-207.

Ponder, W. F. and D. L. Lindberg. 1996. Gastropod Phyloge-

Page 37

ny—Challenges for the 90's. In: J. Taylor (ed). Origin and

Evolutionary R Radiation of the Mollusca. Oxford University

Press, Oxford, pp. 185-154.

Ponder, W. F. and A. Warén. 1988. Classification of the Caen-

ogastropoda and Heterostropha—a list of the family group

names and higher taxa. Malacological Review Supplement

4: 288-326.

Powell, A. W. B. 1929. The Recent and Tertiary species of the

Genus Buccinulum in New Zealand, with a review of re-

lated genera and families. Transactions of the New Zea-

land Institute 60: 57-98.

Powell, A. W. B. 1951. Antarctic and Subantarctic Mollusca:

Pelecypoda and Gastropoda. Discovery Reports 26: 47—

196, pls. 5-10.

Powell, A. W. B. 1958. Mollusca from the Victoria-Ross Quad-

rants of Antarctica. B.A.N.Z. Antarctic Research Expedi-

tion (1929-1931). Report Series B. (Zoology & Botany)

6(9): 165-215.

Powell, A. W. B. 1960. Antarctic and Subantarctic Mollusca.

Records of the Auckland Institute and Museum 5 (3 and

4): 117-193.

Rex, M. A. 1977. Zonation in deep- sea gastropods: the impor-

tance of biological interactions to rates of zonation. In: B.

F. Keegan, P. O Ceidigh, and P. J. S. Boaden (eds.). Bi-

ology of Benthic Organisms. Pergammon Press, New

York. pp. 521-530.

Rex, M. A. 1981. Community structure in the deep-sea ben-

thos. Annual Review of Ecology and Systematics 12: 331—

353.

Rex, M. A. 1983. Geographical pattems of species diversity in

the deep-sea benthos. In: The Sea G. T. Rowe (ed). John

Wiley, New York. Volume 8, pp. 453-472.

Rex, M. A., R. J. Etter, A. J. Cain and M. S. Hill. 1999, Bathy-

metric patterns of body size in deep-sea gastropods. Evo-

lution 53: 1298-1301,

Suter, H. 1913. Manual of the New Zealand Mollusca. John

Mackay, Wellington. xvi + 1120 pp.

Taylor, J. D. 1978. The diet of Buccinum undatum and Nep-

tunea antiqua (Gastropoda: Buccinidae). Journal of Con-

chology 29: 309-318.

Thiele, J. 1904. Die beschalten Gastropoden der deutschen

Tiefsee-Expedition 1898-1899. B. Anatomisch-systematis-

che Untersuchungen einiger Gastropoden. Wissenschaf-

tliche Ergebnisse der deutschen Tiefsee-Expedition auf

dem Dampier: Valdivia” 7: 147-180, pls. 6-9.

Thiele, J. 1912. Die antarktischen Schnecken und Muscheln.

Deutschen Sudpolar-Expedition 1901-1903. Vol. 13: 183—

285, pls. 11-19.

Thiele, J. 1929. Handbuch der systematischen Weichtierkunde.

G. Fischer, Stuttgart, 1-376.

Tomlin, J. R. Le B. 1932. Reports on the marine Mollusca in

the collections of the South African Museum. VI-VIII.

Annals of the South African Museum 30(2): 157-169.

Tracey, S., J. A. Todd and D. H. Erwin. 1993. Mollusca: Gas-

tropoda. In: M. J. Benton (ed), The Fossil Record 2.

Chapman and Hall, London, pp. 137-167.

Warén, A. and P. Bouchet. 2001. Gastropoda and Monopla-

cophora from hydrothermal vents and seeps; new taxa and

records. The Veliger 44: 116-231.

Wenz, W. 1938-1943. Gastropoda. Allgemeiner Teil und Pro-

sobranchia. In: O. H. Schindewolf (ed). Handbuch der

Paliozoologie. Gebriider Borntraeger, Berlin, pp. 1—1506.

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Appendix 1. A review of the taxonomic placement of

the genera of Antarctic and Magellanic Buccinulidae.

The family Buccinulidae was proposed by Finlay

(1928: 250) to unite a number of genera from the region

of New Zealand that had previously been included in

Muricidae, Neptuniidae (as Chrysodomidae), and Buc-

cinidae by earlier workers (Hutton, 1880; Cossmann,

1906; Suter, 1913). Shortly thereafter, Powell (1929: 58)

expanded and revised Finlay’s classification based mainly

on the morphology of the radula, sorting primarily aus-

tral genera into the families:

Buccinulidae. Recognized by having a radula with 3

cusps on rachidian teeth and 3 cusps on lateral teeth,

and an operculum with a terminal nucleus. This family

was further subdivided into groups A-E based on mor-

phology of the siphonal canal, parietal tubercle, and pro-

toconch.

Neptuniidae. Recognized by having a radula with 4

cusps on rachidian teeth and 3 cusps on lateral teeth.

The operculum also has a terminal nucleus.

Buccinidae. Characterized by having a radula with

about 6 cusps on rachidian teeth and 4 cusps on lateral

teeth. The operculum has a median, submarginal nucle-

us.

Cominellidae. Distinguished by having a radula with

3 cusps on rachidian teeth, 2 cusps on lateral teeth, and

an operculum with a terminal nucleus.

Published the same year, Thiele’s (1929) treatment of

the Stenoglossa interpreted the family Buccinidae far

more broadly, subsuming all of the families treated by

Powell (1929) without recognizing any natural groupings

or subdivisions. Powell (1951: 131) proposed a revised

classification of the southern whelks, reatfirming his ear-

lier (Powell, 1929) narrower interpretation of Buccinidae

(limited to the genera Buccinum Linné, 1758 and Bur-

nupena Iredale, 1918), and stating that the Buccinulidae

were more closely related to the northern Neptuniidae

than to the Buccinidae. He subdivided the Buccinulidae

into the subfamilies Buccinulinae, Prosiphiinae and

Cominellinae. In subsequent works, Powell (1960) again

elevated Cominellinae to family status. In the section

below, we list chronologically the genera originally as-

signed to each subfamily by Powell (1951), update the

systematics for Antarctic and Magellanic genera, and add

genera from this region that were published subsequent-

ly.

Family Buccinulidae Finlay, 1928

Subfamily Buccinulinae Finlay, 1928.

Defined on the basis of a radula with three cusps on the

rachidian teeth and three cusps on the lateral teeth.

Originally included genera:

New Zealand: Buccinulum Swainson, 1837: Aeneator

Finlay, 1926; Verconella Iredale, 1915.

Australia: Austrosipho Cossmann, 1906; Berylsma

Iredale, 1924.

California: Kelletia Bayle, 1884.

Antarctic and Subantarctic: Chlanidota Martens,

1878; Pfefferia Strebel, 1908 (reduced to a subgenus of

Chlanidota by Harasewych and Kantor, 1999); Neobuc-

cinum Smith, 1877; Probuccinum Thiele, 1912: Cavi-

neptunea Powell, 1951; Bathydomus Thiele, 1912. [ad-

ditional genera: Chlanificula Powell, 1958; Antarctonep-

tunea Dell, 1972; Parabuccinum Harasewych, Kantor

and Linse, 2000; Spikebuccinum Harasewych and Kan-

tor, herein; Drepanodontus Harasewych and Kantor,

herein].

Neobuccinum Smith, 1877

Type Species: Buccinopsis eatoni Smith, 1875 (by

monotypy).

Distribution: Circum-Antarctic, Kerguelen and

Heard Islands, in 6-1335 m.

Remarks: Shell large (to 86 mm). Operculum large,

occupying nearly entire aperture, oval, paucispiral. Rach-

idian teeth with 3 strong cusps, broad basal plate. Lateral

teeth with 3 cusps, middle cusp small.

Diversity: Currently a monotypic genus. Numanami

(1996: 147) noted some geographical differences in shell

morphometry.

Chlanidota (Chlanidota) Martens, 1878

Type Species: Cominella (Chlanidota) vestita Mar-

tens, 1878 (by monotypy).

Distribution: Circum-Antarctic. Kerguelen Island,

the Crouzets, Herald Island, South Georgia Island, Ant-

arctica, in 38-1100 m.

Remarks: Shell medium-sized (to 43 mm). Opercu-

lum very small (less than 0.4 AL), coiled, with terminal

nucleus. Rachidian teeth with 3 strong cusps, broad, an-

teriorly indented basal plate, lateral teeth with 3 cusps,

middle cusp small.

Diversity: 5 species (recent revision—Harasewych

and Kantor, 1999).

Chlanidota (Pfefferia) Strebel, 1908

Type species: Pfefferia palliata Strebel, 1908 (by sub-

sequent designation, Wenz, 1943).

Distribution: South Georgia Island, in 45-1600 m.

Remarks: Shell medium-sized (to 38 mm). Opercu-

lum large (0.5-0.8 AL), leaf-shaped, coiled, with termi-

nal nucleus. Posterior edge of operculum with tall ridge

of feathered lamellae. Rachidian teeth with 3 strong

cusps, broad, anteriorly indented basal plate. Lateral

teeth with 3 cusps, middle cusp small.

Diversity: 3 species (recent revision—Harasewych

and Kantor, 1999).

Probuccinum Thiele, 1912

Type species: Neobuccinum tenerum Smith, 1907 (by

subsequent designation, Wenz, 1943).

M. G. Harasewych and Y. I. Kantor, 2004

Page 39

Distribution: Circum-Antarctic, in 50-590 m.

Remarks: Shell medium-sized (to 30 mm). Opercu-

lum large (~% AL), with terminal nucleus. Rachidian

teeth ila 3 cusps, central cusp largest, may bear den-

ticles (Numanami, 1996: fig. 104H). basal plate broad,

rectangular. Lateral teeth with 3 cusps of approximately

equal size.

Diversity: 8 species (Powell, 1960).

Bathydomus Thiele, 1912

Type species: Bathydomus obtectus Thiele, 1912 (by

original designation).

Distribution: Antarctic, Marion Island and Crozets, in

2500-3400 m.

Remarks: Shell medium-sized (to 30 mm). Opercu-

lum large (~0.5 AL) long, narrow, wedge-shaped, with

terminal nucleus. Rachidian teeth with 3 strong cusps,

middle cusp largest, flanked by 1 weak denticles on each

side. Basal plate broad, squarish, anteriorly indented.

Lateral teeth with 2 large cusps with 2-3 smaller cusps

between them (Thiele, 1912: pl. 16, fig. 23).

Diversity: 3 species (Dell, 1990: 198).

Cavineptunea Powell, 1951

Type species: Cavineptunea monstrosa Powell, 1951

(by original designation).

Distribution: South Georgia Island, in 90-700 m.

Remarks: Shell large (to 70 mm), with characteristic

protoconch, “like a tall, spirally wound collar, and sur-

rounds a deep apical cavity” (Powell, 1951: fig. N105).

Operculum large (~% AL), with terminal nucleus. Rach-

idian teeth with 3 cusps, central cusp largest, broad, an-

teriorly indented basal plate. Lateral teeth usually with

3, occasionally 4 cusps, outermost cusps largest.

Diversity: A monotypic genus.

Chlanificula Powell, 1958

Type species: Chlanificula thielei Powell, 1958 (by

original designation).

Distribution: Weddell Sea to off Enderby Land, Ant-

arctica, in 220-660 m.

Remarks: Shell medium-sized (to 30 mm). Opercu-

lum large, occupying nearly entire aperture, narrow,

wedge-shaped, with terminal nucleus. Rachidian teeth

with strong central cusp, flanked by one smaller cusp

and one denticle on each side. Basal plate squarish, shal-

lowly indented anteriorly. Lateral teeth with 3 cusps,

middle cusp smaller, additional denticles may appear be-

tween cusps (Hain, 1990: pl. 23, fig. 2). Powell (1958)

regarded this genus to be part of the “Chlanidota com-

plex” closest to Notoficula. The genus Notoficula Thiele,

1917, has been transferred to Eratoidae by Oliver (1983)

based on radular morphology.

Diversity: A monotypic genus.

Antarctoneptunea Dell, 1972

Type species: Fusitriton aurora Hedley, 1916 (by

original designation).

Distribution: Ross Sea to off Enderby Land, Antare-

tica, 15-603 m.

Remarks: Shell large (to 102 mm). Operculum large

(~% AL), with terminal nucleus. Rachidian teeth with

three cusps. Basal plate broad, deeply indented anteri-

orly. Lateral teeth with 3 cusps, middle cusp smaller.

This genus is hardly distinguishable from Penion Fischer,

1884 in general shell shape, radula and operculum (Pon-

der, 1973).

Diversity: A monotypic genus.

Kantor and Linse, 2000

Type species: Chlanidota bisculpta Dell, 1990 (by

original designation).

Distribution: Magellanic region, in 247-2165 m.

Remarks: Shell small (to 16.5 mm). Operculum large

(> 0.5 AL), oval, paucispiral. Rachidian teeth with 3

strong cusps, broad, anteriorly indented basal plate. Lat-

eral teeth with 3 cusps, middle cusp small. Included in

Buccinulinae by Harasewych, Kantor and Linse (2000).

Diversity: 4 species (recent revision—Harasewych,

Kantor and Linse, 2000).

Parabuccinum Harasewych, K

Spikebuccinum Harasewych and Kantor, herein.

Type species: Spikebuccinum stephaniae Harasewych

and Kantor, herein (by original designation).

Distribution: Scotia Sea and adjacent abyssal plains,

in 1967-4645 m.

Remarks: Shell small (to 19.9 mm). Operculum small

(~0.36 AL), broadly ovate, paucispiral, with subterminal

nucleus. Rachidian teeth with 3 cusps, central shorter,

weaker that outer cusps. Basal plate broad, deeply in-

dented. Lateral teeth with large outer, smaller inner

cusp, with 3-4 smaller denticles between them.

Diversity: A monotypic genus.

Drepanodontus Harasewych and Kantor, herein.

Type species: Drepanodontus tatyanae Harasewych

and Kantor, herein (by original designation).

Distribution: Scotia Sea and adjacent Argentine

Abyssal Plain, in 2740-5798 m.

Remarks: Shell large (to 43.1 mm). Operculum large

(~0.82 AL), oval, with terminal nucleus. Rachidian teeth

usually with 3 cusps, central cusp large, may or may not

be flanked by one lateral cusp and aaléktionall denticles.

Basal plate narrow, rectangular, weakly indented. Lateral

teeth with 1 cusp, which may bear 1 or more denticles,

near inner edge of basal plate.

Diversity: A monotypic genus.

Subfamily Prosiphiinae Powell, 1951

Defined on the basis of a radula with 3 cusps on the

rachidian teeth, and lateral teeth with multiple cusps.

Page 40

THE NAUTILUS, Vol. 118, No. 1

Contains taxa with heterogeneous lateral tooth mor-

phologies, that may be divided into: Proneptunea type

lateral teeth, with multiple cusps spanning the width of

the basal plate, usually fused above the basal plate, may

have one or more denticles along the outer edge; Pro-

sipho type lateral teeth, with cusps confined to inner

portion of basal plate, often rotated to form an acute

angle with the basal plate, the outer portion of which is

thin and lacks teeth. Prosiphiinae appear to be restricted

to Antarctic and subantarctic seas. Originally included

genera:

Antarctic and subantarctie: Proneptunea Thiele,

1912; Meteuthria Thiele, 1912; Prosipho Thiele, 1912:

Anomacme Strebel, 1905; Fusinella Thiele, 1917 (re-

placement name for Buccinella Thiele, 1912, non Perry,

1811); Chlanidotella Thiele, 1929. [additional genera:

Savatieria Rochebrune and Mabille, 1885; Crenatosipho

Linse, 2002; Muffinbuccinum Harasewych and Kantor,

herein; Germonea Harasewych and Kantor, herein].

Savatieria Rochebrune and Mabille, 1885

Type species: Savatieria frigida Rochebrune and Ma-

bille, 1885 (by monotypy).

Distribution: Magellanic region, in 100 m.

Remarks: Shell smell (usually < 10 mm). Operculum

large, oval, with terminal nucleus. Radula Prosipho type.

TPs Fam teeth narrow, with 3 cusps. Lateral teeth with

4 short, broad cusps, basal plate prolonged anteriorly.

Diversity: 7 species (Powell, 1960: 149).

Anomacme Strebel, 1905

Type species: Anomacme smithi Strebel, 1905 (by

monotypy).

Distribution: Magellanic region, 220-250 m.

Remarks: Shell small (< 10 mm). Operculum not de-

scribed. Radula Prosipho type. Rachidian teeth with 3

cusps, basal plate squarish, broader than long, weakly

indented anteriorly. Lateral teeth with long, narrow basal

plate, with 6 cusps concentrated toward the inner por-

tion of the radular tooth, 4th cusp longest (Thiele, 1912:

pl. 16, fig. 14). This genus was included in Cominellidae

by Pomel (1960: 149), presumably because of its simi-

larity i in shell shape to Glypteuthria, despite its different

radular morphology.

Diversity: A monotypic genus.

Proneptunea Thiele, 1912

Type species: Proneptunea amabilis Thiele, 1912 (by

original designation).

Distribution: Kerguelen, South Georgia Island, South

Orkney Islands, Ross Sea, Antarctica, in 12-870 m.

Remarks: Shell small (< 15 mm). Operculum medi-

um-sized, oval, with terminal nucleus. Radula Pronep-

tunea type. Rachidian teeth with 3 cusps. Basal plate

longer than wide, shallowly indented anteriorly. Lateral

teeth broad, spatulate, with 5 long cusps fused above

basal plate. Outer edge may have multiple denticles

(Powell, 1951: fig. K 66-67).

Diversity: 5 species (Dell, 1990: 199).

Meteuthria Thiele, 1912

Type species: Euthria martensi Strebel, 1905 (by

original designation).

Distibution: Magellanic region, and Ross Sea, Ant-

arctica, in 57-870 m.

Remarks: Shell very small (< 5 mm). Operculum un-

described. Radula Prosipho type, without rachidian

teeth. Lateral teeth with multiple (4-5) cusps concen-

trated toward inner portion of tooth. Outermost cusp

may have denticles (Powell, 1951: fig. K62; Dell, 1990:

fig. 296). This genus was included in the subfamily Com-

inellidae by Powell (1960: 149), presumably because of

its similarity in shell shape to Glypteuthria, despite its

different radular morphology.

Diversity: 4 species, 1 subspecies (Powell, 1960: 149;

Dell, 1990: 173).

Prosipho Thiele, 1912

Type species: Prosipho gaussianus Thiele, 1912 (by

subsequent designation, Thiele. 1929).

Distribution: ‘(Grain Antarctic, in 12-800 m.

Remarks: Shell small, rarely > 10 mm. Operculum

large (~% AL), oval, with coiled nucleus. Radula Pro-

sipho type. Rachidian teeth with 3 cusps, basal plate

squarish to longer than broad, weakly indented anteri-

orly. Lateral teeth with long, narrow basal plate, with 2

or more cusps concentrated toward the inner portion of

the radular tooth.

Diversity: About 40 species (Powell, 1951; Dell, 1990;

Numanami, 1996).

Fusinella Thiele, 1917 (replacement name for

Buccinella Thiele, 1912, non Perry, 1811)

Type species: Buccinella jucunda Thiele, 1912 (by

monotypy).

Distribution: Kerguelen Island, 0-100 m.

Remarks: Shell very small (< 6 mm). Cperculum

oval, with terminal nucleus. Radula Prosipho type. Rach-

idian teeth with 3 cusps of equal size near center, with

an additional cusp at each posterior corner of the squar-

ish basal plate with a deep, V-shaped indentation ante-

riorly. Lateral teeth with long, narrow basal plate, with

multiple (5-6) cusps concentrated along inner half of

basal plate and directed parallel to its long axis (Thiele,

1912: pl. 16, fig. 13).

Diversity: A monotypic genus.

Chlanidotella Thiele, 1929

Type species:

monotypy).

Cominella modesta Martens, 1885 (by

M. G. Harasewych and Y. I. Kantor, 2004

Distribution: South Georgia, 0-18 m.

Remarks: Shell medium-sized (< 15 mm). Opercu-

lum medium-sized (~0.5 AL), oval, with terminal nu-

cleus. Radula Proneptunea type. Rachidian teeth with 3

cusps, central cusp slightly longer than flanking cusps.

Basal plate broader than long, deeply indented anteri-

orly. Lateral teeth with 4 cusps roughly equal in length,

spanning the entire width of the basal plate. Thiele’s il-

lustration (1912: textfig. 11) of the radula shows a den-

ticle along the outer edge along one side of the radula,

while Powell’s illustration (1951: fig. L 80) does not.

Diversity: A monotypic genus.

Crenatosipho Linse, 2002

Type species: Crenatosipho beaglensis Linse, 2002

(by original designation).

Distribution: Magellanic region, in 67-200 m.

Remarks: Shell small (to 9 mm). Operculum large,

oval with eccentric nucleus. Radula Prosipho type. Rach-

idian teeth very narrow, long, without cusps. Lateral

teeth with up to 6 cusps long concentrated toward the

inner margin of the tooth, with long, cuspless outer por-

tion of the basal plate. (Linse, 2002: fig. 9.1.1-112).

Diversity: A monotypic genus.

Muffinbuccinum Harasewych and Kantor, herein

Type species: Muffinbuccinum catherinae Harasew-

ych and Kantor, herein (by original designation).

Distribution: Argentine Abyssal Plain, off the north-

em slope of the Falkland Plateau, in 5685-5798 m.

Remarks: Shell of moderate size (to 27 mm). Oper-

culum large (~0.56 AL), triangular, tapering toward ter-

minal nucleus. Radula Proneptunea type. Rachidian

teeth with 3 cusps, central cusp slightly longer than

flanking cusps. Basal plate broad, short, shallowly in-

dented anteriorly. Lateral teeth with 7-8 cusps spanning

basal plate, innermost cusp longest, outermost shortest,

with or without denticles along outer edge.

Diversity: A monotypic genus.

Germonea Harasewych and Kantor, herein

Type species: Germonea rachelae Harasewych and

Kantor, herein (by original designation).

Distribution: Abyssal plain of the Scotia Sea, in 2196—

3714 m.

Remarks: Shell large (to 68 mm). Operculum large

(~0.65 AL), narrow, recurved, with terminal nucleus.

Radula Prosipho type. Rachidian teeth with 3 cusps, very

long, narrow, deeply indented anteriorly, appearing Y-

shaped. Lateral teeth with 3-4 broad, stout cusps con-

centrated toward the inner margin of the tooth, with

short, cuspless outer portion of the basal plate.

Diversity: A monotypic genus.

Page 41

Subfamily Cominellinae Gray, 1857

Defined on the basis of a radula with 3 cusps on the

rachidian tooth, and lateral teeth with 2 cusps. Originally

included genera:

New Zealand and Australia:

Fax Iredale, 1925.

Tropical Pacific: Phos Montfort, 1810.

Northwest Pacific: Searlesia Harmer, 1914.

Antarctic and Subantarctic: Pareuthria Strebel,

1905; Tromina Dall, 1918; Notoficula Thiele, 1917 [now

in Eratoidae]; Falsimohnia Powell, 1951; Glypteuthria

Strebel, 1905. [additional genera: Antistreptus Dall,

1902; Parficulina Powell, 1958; Antarctodomus Dell,

1972; Lusitromina Harasewych and Kantor, herein].

Cominella Gray, 1850;

Antistreptus Dall, 1902

Type species: Antistreptus magellanicus Dall, 1902

(by original designation).

Distribution: Magellanic region, in 30-600 m.

Remarks: Shell very small (< 5 mm), sinistral. Oper-

culum described but not figured by Linse (2002: 100) as

“small, horny, brown, nine, eccentric, with a large oval

foot muscle scar,” The radula is unknown. Apart from

being sinistral, shell shape is similar to Glypteuthria and

Anomacme. This genus was described within Muricidae

and compared to Trophon by Dall (1902). Powell (1951)

included it in Buccinulidae, but later placed it in Com-

inellidae (Powell, 1960).

Diversity: 2 species, Antistreptus magellanicus Dall,

1902, A. rolani Castellanos, 1985.

Pareuthria Strebel, 1905

Type species: Fusus plumbeus Philippi, 1844 (by sub-

sequent designation, Tomlin, 1932).

Distribution: Magellanic region, Cireum-Antarctic, in

0-549 m.

Remarks: Shell small (< 10 mm). Operculum ovate,

with terminal nucleus. Despite shell similarities between

Magellanic and Antarctic species, there are conspicuous

differences in radular dentition between species inhab-

iting these regions. Magellanic species, including the

type species of the genus, have broad rachidian teeth

with 3 subequal cusps. Antarctic species have rachidian

teeth with squarish basal plates and a large triangular

central cusp that may or may not be flanked by one or

rarely more (Numanami, 1996: fig. 125D), smaller den-

ticles on each side. Lateral teeth with long, stout basal

plates that give rise to 2 long, recurved cusps.

Diversity: About 18 species (Powell, 1960; Numana-

mi, 1996).

Glypteuthria Strebel, 1905

Type species: Euthria meridionalis Smith, 1881 (by

subsequent designation, Tomlin, 1932).

Page 42

Distribution: Magellanic region, Southern Africa, in

6-250 m.

Remarks: Shell medium-sized (< 29 mm). Opercu-

lum ovate, with terminal nucleus. Rachidian teeth with

3 cusps on squarish, posteriorly indented basal plate.

Lateral teeth with broad basal plate, 3 cusps, inner and

outer cusps long, intermediate cusp short. Powell (1951:

138) regarded Glypteuthria to be a strongly sculptured

relative of Pareuthria, with the intermediate cusp a bi-

furcation of the inner cusp, and not comparable to the

3rd cusp of Probuccinum.

Diversity: 7 species, 4 Magellanic, 3 South African.

Notoficula Thiele, 1917 (New name for Ficulina

Thiele, 1912, non Gray, 1867)

Type species: Ficulina bouveti Theile, 1912 (by mon-

otypy). The anatomy, radular morphology, and opercu-

lum of the type species remain unknown. Oliver (1983)

described an additional species of Notoficula, and, based

on its radular morphology and anatomy, transferred the

genus to Eratoidae. Powell (1958: 192) reconsidered the

affinities of the species he originally described as Noto-

ficula problematica Powell, 1951, and erected the genus

Parficulina Powell, 1958 to accommodate it (see below).

Tromina Dall, 1918

Type species: Fusus unicarinatus Philippi, 1868 (by

original designation). The type species of this genus was

shown to belong to the family Muricidae. Dell (1990)

proposed the name Falsitromina to include the bathyal

Magellanic buccinoidean taxa previously included in

Tromina. The genus Lusitromina is proposed herein for

the abyssal species that had been assigned to Tromina.

For a detailed review of the taxonomy of Tromina, see

the “Remarks” section under the description of Lusi-

tromina.

Falsimohnia Powell, 1951

Type species: Buccinwm albozonatum Watson, 1881

(by original designation).

Distribution: South Georgia, Kerguelen Island, in

18—250 m.

Remarks: Shell small (< 10 mm), operculum small,

with blunt, terminal nucleus. Rachidian tooth with sin-

gle, triangular cusp on squarish basal plate with shallow

anterior indentation. Lateral teeth with long basal plate

giving rise to 2 teeth, outermost broader, longer. Powell

(1951: 137) regarded Falsimohnia to be derived from

Pareuthria.

Diversity: A monotypic genus (see comments under

Antarctodomus diversity).

THE NAUTILUS, Vol. 118, No. 1

Parficulina Powell, 1958

Type species: Notoficula problematica Powell, 1951

(by original designation).

Distribution: Magellanic region, in 545 m.

Remarks: Shell small (to 7 mm). Operculum medium

sized (~0.50 AL), D-shaped, paucispiral. Rachidian

teeth with 3 equal cusps on a squarish basal plate. Lat-

eral teeth with 2 cusps, the inner cusp much broader

than the outer cusp. See remarks under Notoficula

(above).

Diversity: A monotypic genus.

Antarctodomus Dell, 1972

Type species: Bathydomus thielei Powell, 1958 (by

original designation).

Distribution: Eastern hemisphere, from the Ross Sea

to off Queen Maud Land, Antarctica, 110-420 m.

Remarks: Shell medium-sized (to 30 mm). Opercu-

lum long, triangular, with terminal nucleus (Dell, 1972).

Rachidian tooth with 3 cusps, basal plate broad, short,

deeply indented anteriorly. Lateral teeth with 2 long,

blunt, cusps, basal plate narrower than rachidian teeth.

Diversity: 2 species (Numanami, 1996: 147. In our

view, Antarctodomus okutanii Numanami, 1996 differs

in opercular and radular morphology from the type spe-

cies of Antarctodomus, and may be referable to Falsi-

mohnia).

Falsitromina Dell, 1990

Type species: Tromina bella Powell, 1951 (by original

designation).

Distribution: Magellanic region, in 81-878 m [Dell

(1990: 175) reported but did not illustrate or provide

catalog numbers for two specimens he attributed to F

bella from the Ross Sea, at depths of 1565-1674 m.]

Remarks: Shell small (to 14 mm). Operculum medi-

um-sized (~0.50 AL), oval, paucispiral. Rachidian teeth

with 3 cusps, central cusp may be slightly longer, on a

squarish basal plate, shallowly indented anteriorly. Lat-

eral teeth with two broad curved cusps that join above

the basal plate.

Diversity: 5 species (Dell, 1990).

Lusitromina Harasewych and Kantor, herein

Type species:

inal designation).

Distribution: Scotia Sea, and adjacent abyssal plains,

Cape Basin, in 2380-5480 m.

Remarks: Shell medium sized (to 29 mm). Opercu-

lum D-shaped, large (~0.61 AL), broadly ovate, paucis-

piral. Rachidian teeth with 3 cusps on posterior portion

of broad, anteriorly arched basal plate. Central cusp

slightly longer, wider than lateral cusps. Lateral teeth

with 2 cusps of approximately equal length.

Diversity: 2 species.

Tromina abyssorum Lus, 1993 (by orig-

THE NAUTILUS 118(1):43-51, 2004

Page 43

New species of Late Cretaceous Cypraeidae (Gastropoda) from

California and British Columbia and new records from the

Pacific slope

Lindsey T. Groves

Natural History Museum of Los

Angeles County

Malacology Section

900 Exposition Boulevard

Los Angeles, CA 90007 USA

[email protected]

ABSTRACT

Cretaceous cypraeids are uncommon in North American strata

and comprise 15 recognized species, seven of which are from

the Pacific slope of North America (Groves, 1990). Four new

species are described herein from localities in southern and

northern California and British Columbia, Canada: Palaeocy-

praea (Palaeocypraea) wilfredi new species and Bernaya (Ber-

naya) jeanae new species, both from the Upper Cretaceous

(lower Campanian) Chico Formation, Butte County, California;

Bernaya (Bernaya) beardi new species from the Upper Cre-

taceous (uppermost Santonian to lowermost Campanian) upper

Haslam Formation, Vancouver Island, British Columbia, Can-

ada; and Bernaya (Protocypraea) popenoei new species from

the Upper Cretaceous (lower Campanian) Ladd Formation,

Orange County, California. The late Campanian to early Maas-

trichtian was the Mesozoic peak of cypraeids, in terms of num-

bers of species and geographic distribution both in North

America and worldwide (Groves, 1994). New paleogeographic

and chronologic records of previously described species and

indeterminate species are listed as well.

INTRODUCTION

Four new Late Cretaceous species of cypraeid gastro-

pods, uncommon in strata of that geologic age in North

America, are described from localities in Orange and

Butte counties, California, and Vancouver Island, British

Columbia (Figure 1). Two are from the lower Campan-

ian Chico Formation, Butte County, California; the third

is from the uppermost Santonian to lowermost Campan-

ian upper Haslam Formation, near Brannen Lake, Van-

couver Island, British Columbia, Canada; and the fourth

new species is from the lower Campanian Holz Shale

Member of the Ladd Formation, Santa Ana Mountains,

Orange County, California. Cypraeid records from the

Chico and Ladd formations are recorded here for the

first time.

STRATIGRAPHY AND GEOLOGIC AGE

The formations listed below, from oldest to youngest, are

those from which the new cypraeid taxa are described.

Squires and Saul (2001) recently described several new

species of gastropods from these formations and dis-

cussed their geologic age, stratigraphy, and paleoenvi-

ronment. Therefore, only a brief overview of stratigraph-

ic nomenclature and age of the units will be dksewesed

and readers are referred to additional sources for de-

tailed descriptions.

HasLaM FORMATION

The Haslam Formation of Clapp (1912) was described

for outcrops on southeastern Vancouver Island, British

Columbia. Based on ammonite biostratigraphy, Muller

and Jeletzky (1970) cited the age of the formation as late

Santonian to early Campanian, as did Ward (1978) and

Haggart (1991). Squires and Saul (2001) concurred with

a late Santonian to early Campanian age for the forma-

tion and indicated that magnetostratigraphic analysis

could more precisely refine the stage boundary. Mag-

netostratigraphic work by Enkin et al. (2001 and pers.

commun.) indicated that the formation was entirely

Campanian. However, most recently Mustard et al.

(2003 and pers. commun.) concluded that the formation

was diachronous and contained both late Santonian and

early Campanian fossils.

PENTZ ROAD MEMBER OF THE CHICO FORMATION

Russell et al. (1986) described the informal Pentz Road

member of the Chico Formation for outcrops near

“Pence’s Ranch” (= Pentz), Butte County, northern Cal-

ifornia. Based on the presence of the ammonites Sub-

mortoniceras chicoense (Trask, 1856) and Baculites chi-

coensis (Trask, 1856), they assigned an early Campanian

age to these outcrops. Interestingly, Haggart et al. (1997)

Page 44

THE NAUTILUS, Vol. 118, No. 1

Figure 1.

Index map showing type localities for new taxa and

other pertinent geographic areas mentioned in the text (mod-

ified with permission from Squires and Saul, 2001). 1 = Van-

couver Island, British Columbia; 2 = Sucia Island, San Juan

County, Washington; 3 = Gualala, Mendocino County, Cali-

fornia; 4 = Pentz, Butte County, Califomia; 5 = Santa Ana

Mountains, Orange County, California; 6 = Carlsbad, San Di-

ego County, California.

informally named a Pentz member for the same beds

described by Russell et al. (1986) and interpreted these

facies as a very nearshore shallow-marine environment.

Squires and Saul (1997; 2001) concurred with the early

Campanian age; based on the presence of the soft-bot-

tom dwelling gastropod Boggsia tenuis (Gabb, 1864),

however, they also interpreted the paleoenvironment as

shallow marine rather than the estuarine environment

reported by Russell et al. (1986).

Houz SHALE MEMBER OF THE LADD FORMATION

Popenoe (1942) described the Holz Shale Member of

the Ladd Formation for lower Campanian outcrops in

Ladd Canyon, Santa Ana Mountains, Orange County,

California. The fossiliferous upper part of the member

is dominated by sandstone beds deposited in a deep-

shelf environment (Squires and Saul, 2001).

ABBREVIATIONS

Abbreviations used for institutional catalog and/or local-

ity numbers are as follows: CAS, California Academy of

Sciences, San Francisco; CIT, California Institute of

Technology (collections now at LACMIP); LACMIP,

Natural History Museum of Los Angeles County, Inver-

tebrate Paleontology Section; RBCM, Royal British Co-

lumbia Museum, Vancouver; SC, Sierra College, Rock-

lin, California; SDSNH, San Diego Society of Natural

History; UCLA, University of California, Los Angeles

(collections now at LACMIP); UCMP, University of Cal-

ifornia, Museum of Paleontology, Berkeley; USGS, Unit-

ed States Geological Survey, Menlo Park, California (col-

lections now at UCMP): and VIPM, Vancouver Island

Paleontological Museum, Qualicum Beach, Vancouver

Island, British Columbia, Canada. Measurement param-

eters are defined as follows: length = greatest distance

between anterior and posterior ends; width = greatest

distance between lateral margins; and height = greatest

distance between base and dorsum. The systematic clas-

sification herein follows that of Schilder and Schilder

(1971).

SYSTEMATIC PALEONTOLOGY

Superfamily Cypraeoidea Rafinesque, 1815

Family Cypraeidae Rafinesque, 1815

Subfamily Bernayinae Schilder, 1927

Tribe Archicypraeini Schilder, 1927

Genus Palaeocypraea Schilder, 1928

Type Species: Cypraeacites spiratus Schlotheim,

1820 by original designation. Early Paleocene (Danian),

Faxe, Denmark.

Diagnosis: Shell small to medium in size, elongated,

spire broad and partially covered, aperture wide with

deep terminal canals and fine dentition; fossula broad,

concave and smooth.

Remarks: Schilder and Schilder (1971) recognized

L. T. Groves, 2004

Page 45

Pa M9 indi

Figures 2, 3. Palaeocypraea (Palaeocypraea) wilfredi new species, holotype, LACMIP 13065, from LACMIP loc. 17611, 35.8

mm length. Figures 4,5. Bernaya (Bernaya) jeanae new species, holotype, LACMIP 13067, from LACMIP loc. 17611, 42.4

mm length. Figures 6-9. Bernaya (Bernaya) beardi new species, holotype RBCM.EH2003.008.0001, from Vancouver Island,

British Columbia, 36.7 mm length. Figures 10, 11. Bernaya (Protocypraea) popenoei new species, holotype UCMP 154951,

from UCMP loc. A3404, 31.4 mm length.

nine species and 10 subspecies of Cretaceous Palaeo-

cypraea s.s. Of these, six are from North America and

two are from the Pacific slope (Groves, 1990). Palaeo-

cypraea (Palaeocypraea) fontana (Anderson, 1958) from

the Lower Cretaceous (uppermost lower Albian), Bud-

den Canyon Formation, Shasta County, California, is the

earliest known cypraeoidean from the Western Hemi-

sphere. Other North American species are from San

Juan County, Washington, Navarro County, Texas, New

Castle County, Delaware, and Dawson County, Montana

(Groves, 1990).

Subgenus Palaeocypraea Schilder, 1928

Palaeocypraea (Palaeocypraea) wilfredi new species

(Figures 2-3)

Diagnosis: A Palaeocypraea of medium size, elongate

shell, broad spire, fine dentition, fossula concave and

smooth.

Description: Shell medium in size, slightly constrict-

ed anteriorly; maximum height and width near center;

spire of medium height, partially covered by successive

whorls; dorsum slightly flattened; aperture narrow, fairly

Page 46

THE NAUTILUS, Vol. 118, No. 1

straight; denticulation fine with smooth interstices, outer

lip with 18 teeth, inner lip with six teeth; outer lip with

prominent anterior terminal ridge, forming slight mar-

ginal callus.

Comparison: The new species is most similar to Pa-

laeocypraea (Palaeocypraea) suciensis (Whiteaves, 1895:

127-128, pl. 3, fig. 5) from the Upper Cretaceous (lower

Campanian) Cedar District Formation, Nanaimo Group,

Sucia Island, San Juan County, Washington. Palaeocy-

praea (Palaeocypraea) wilfredi is larger, has finer aper-

tural dentition, a narrower aperture, shallower anterior

and posterior canals, and a more cylindrical shape than

P. (P_) suciensis.

Discussion: Although post-burial crushing has dam-

aged part of the posterior dorsum, preservation is ade-

quate enough for unequivocal generic and subgeneric

assignments. Palaeocypraea (P.) wilfredi is the first cy-

praeoidean reported from the Chico Formation.

Material: The new species is represented by two spec-

imens. The holotype is slightly crushed, with minor

amounts of original-shell material missing. The paratype

exhibits small amounts of original-shell material but

prominently displays the spire. An anterior outer lip

fragment that exhibits original shell material from LAC-

MIP loc. 24081 is also attributable to the new species.

Type Material: Holotype LACMIP 13065, paratype

LACMIP 13066. Holotype measures 35.8 mm in length,

20.7 mm in width, and 17.0 mm in height. Paratype

measures 34.8 mm in length, 20.6 mm in width, and 16.6

mm in height.

Type Locality: LACMIP loc. 17611, along Dry Creek,

near Pentz, Butte County, California. Upper Cretaceous

(lower Campanian), informal Pentz Road member, Chi-

co Formation.

Etymology: Named after Wilfred Géhre (father of

Eric Gohre, who collected and donated the type material

to LACMIP) of Oroville, California.

Tribe Bernayini Schilder, 1927

Genus Bernaya Jousseaume, 1884

Type Species: Cypraea media Deshayes, 1835, by

original designation. Upper middle Eocene (Bartonian

Stage), Auvers-sur-Oise, Val-d’Oise, France.

Diagnosis: Shell medium to large in size, anterior end

somewhat carinate, dorsum smooth, spire of medium

height and partially covered by successive whorls, aper-

ture wide, sides rounded, anterior and posterior canals

deep, fossula smooth, concave, wide.

Remarks: Schilder and Schilder (1971) recognized six

species and two subspecies of Cretaceous Bernaya s.s.

Only one of these is from North America; Bernaya (Ber-

naya) burlingtonensis (Schilder, 1932) from the Upper

Cretaceous (upper Campanian), Mt. Laurel-Navesink

Formation, Burlington County, New Jersey. Groves

(1990) described B. (B.) crawfordcatei, the first reported

Bernaya s.s. from the Pacific slope, from the Upper Cre-

taceous (uppermost Campanian to lowermost Maastrich-

tian), Point Loma Formation, near Carlsbad, northern

San Diego County, California.

Bernaya (Bernaya) jeanae new species

(Figures 4—5)

Diagnosis: A Bernaya of medium size, anterior and

posterior canals deep, spire of medium height, fossula

smooth, concave, anterior and posterior terminal ridges

prominent extending to margins.

Description: Shell medium in size, constricted ante-

riorly, maximum height and width posterior to center;

spire of medium height, partially covered by successive

whorls; aperture wide, straight; denticulation faint, outer

lip with 13 teeth, teeth absent from inner lip; outer lip

with prominent anterior and posterior terminal ridges

extending to margins forming slight marginal callus.

Comparison: The new species is most similar to Ber

naya (Bernaya) crawfordcatei Groves, 1990: 278, figs.

17-18, from the Upper Cretaceous (uppermost Cam-

panian to lowermost Maastrichtian), Point Loma For-

mation, San Diego County, California. Bernaya (Ber-

naya) jeanae is smaller in size, has finer apertural den-

tition, less prominent anterior and posterior basal ridges,

and a less sinuous aperture than B. (B.) crawfordcatei.

Discussion: Post-burial crushing has damaged parts of

the aperture and dorso-ventrally distorted the type ma-

terial. Generic and subgeneric assignment are based on

the wide aperture, deep anterior and posterior canals,

and medium-height spire. Along with Palaeocypraea (P.)

wilfredi (described above), this is the second cypraeo-

idean described from the Chico Formation.

Material: The new species is represented by the well

preserved holotype and paratype, both of which exhibit

original-shell material. Topotypic material includes 14

specimens in the collection of Eric Géhre, Oroville Cal-

ifornia, with varying amounts of original shell material.

A single outer lip fragment from LACMIP loc. 24081 is

assigned to the new species. A poorly preserved internal

mold from the Chico Formation at the Granite Bay sub-

division, Placer County, California (SC MG135) is at-

tributable to the new species. An unusually large, poorly

preserved specimen from Dry Creek near Pentz that

measures 115.2 mm in length, 72.3 mm in width, and

38.7 mm, is tentatively identified as the new species.

Type Material: Holotype LACMIP 13067, paratype

LACMIP 13068. Holotype measures 42.4 mm in length,

29.4 mm in width, and 19.8 mm in height. Paratype

measures 47.1 mm in length, 30.6 mm in cordlan and 21.5

mm in height.

Type Locality: LACMIP loc. 17611, along Dry Creek,

near Pentz, Butte County, Califomia. Upper Cretaceous

L. T. Groves, 2004

(lower Campanian), informal Pentz Road member, Chi-

co Formation.

Etymology: Named after Jean Géhre (mother of Eric

Gohre, who collected and donated the type material to

LACMIP) of Oroville, California.

Bernaya (Bernaya) beardi new species

(Figures 6-9)

Bernaya crawfordcatei Groves, 1990: Ludvigsen and Beard,

1994: 93, fig. 58 (left 2 figs.). Ludvigsen and Beard, 1997:

113, fig. 69 (left 2 figs.).

Bernaya (Bernaya) n. sp.: Groves, 1997: 7.

Diagnosis: A Bernaya of medium size, spire of me-

dium height, aperture wide; fossula smooth, concave, an-

terior terminal canal deep.

Description: Shell medium in size; maximum height

and width slightly posterior to center; spire of medium

height, partially covered by successive whorls; aperture

wide, straight; denticulation coarse with smooth inter-

stices, outer lip with 13 teeth, teeth obsolete from inner

lip; outer lip with weak posterior terminal ridge forming

slight callus; marginal callus extends toward dorsum

from outer and inner lip margins, forming coarse den-

ticular pattern.

Comparison: The new species is unlike any known

species of Bernaya (B.) in the Western Hemisphere al-

though it superficially resembles B. (B.) azevedoi (Oliv-

eira, 1957: 20, pl. 2, figs. 1, 3) from Upper Cretaceous

(Maastrichtian) strata, Pernambuco State, Brazil and, B.

(Protocypraea) argonautica (Anderson, 1958: 177, pl. 21,

figs. 44a) from Upper Cretaceous (Cenomanian to Tu-

ronian) Hornbrook Formation, Osburger Gulch Sand-

stone Member (of Nilsen, 1984), Jackson County,

Oregon. However, both species are markedly smaller

than B. (B.) beardi and both are poorly preserved inter-

nal molds with little original-shell material preserved.

Discussion: Post-burial processes have removed much

of the original-shell material from the dorsal surface of

the holotype and the anterior terminal canal area is miss-

ing due to mechanical breakage. Generic and subgeneric

assignments are based on the wide aperture, deep-pos-

terior terminal canal, and spire of medium height. The

unusual coarse marginal denticular pattern could be nat-

ural or an artifact of erosional processes. If this denti-

cular pattern is indeed natural, it is unprecedented

amongst cypraeids. Only species of the Eocene to Re-

cent genus Nucleolaria Oyama, 1959, some members of

the Recent genus Cypraeovula Gray, 1824, the Miocene

to Recent species Ipsa childreni (Gray, 1825), and the

Pleistocene to Recent species Erosaria guttata (Gmelin,

1791), have any outwardly similar marginal sculpture.

Bernaya (B.) beardi appears to represent the northern-

most record for a Cretaceous cypraeid worldwide. How-

ever, recent paleomagnetic paleolatitudinal studies by

Kodama and Ward (2001) indicate that deposition of Na-

naimo Basin sediments may have occurred at or around

Page 47

40° N latitude (northern California) and transported

northward in the post Late-Cretaceous. Enkin et al.

(2001) concluded that the Nanaimo Basin was deposited

near the present day California-Mexico border also

based on paleomagnetic evidence. They also noted that

this interpretation conflicts with sedimentologic and pa-

leontologic evidence established by Elder and Saul

(1993) and Haggart (2000) that the Nanaimo Basin was

deposited near its present northern position.

Material: Represented by a well preserved holotype

and three slightly juvenile topotypic specimens (VIPM

144, 146, and 147), all of which exhibit varying amounts

of original shell material.

Type Material: Holotype RBCM.EH2003.008.0001

(ex VIPM 148), 36.7 mm in length, 28.9 mm in width,

and 21.4 mm in height.

Type Locality: Near Brannen Lake, Vancouver Is-

land, British Columbia, Canada, Upper Cretaceous (up-

permost Santonian to lowermost Campanian), upper

Haslam Formation, Nanaimo Group.

Etymology: Named after Graham Beard, founder of

the Vancouver Island Paleontological Museum, Quali-

cum Beach, Vancouver Island, British Columbia, Cana-

da.

Subgenus Protocypraea Schilder, 1927

Type Species: Eocypraea orbignyana Vredenburg,

1920 by original designation. Upper Cretaceous (Turon-

ian through Santonian), Trichinopoly Group, Kullygoody,

southern India.

Diagnosis: Shell small to medium in size, shape mod-

erately pyriform, somewhat constricted anteriorly; fos-

sula smooth, concave, wide.

Remarks: Schilder and Schilder (1971) recognized

eight species and seven subspecies of Cretaceous Ber-

naya (Protocypraea). Two of their species [B. (P.) argo-

nautica and B. (P.) berryessae both (Anderson, 1958)]

and one subspecies, now recognized as a full species [B.

(P.) gualalaensis (Anderson, 1958)], are from the Pacific

slope of North America. Groves (1990) described B. (P.)

rineyi from the Upper Cretaceous (uppermost Campan-

ian to lowermost Maastrichtian), Point Loma Formation,

near Carlsbad, northern San Diego County, California.

The only other North American species, Bernaya (Pro-

tocypraea) mississippiensis Groves (1990), is from the

Upper Cretaceous (Campanian), Coffee Formation, Lee

County, Mississippi.

Bernaya (Protocypraea) popenoei new species

(Figures 10-11)

Bernaya (Protocypraea) n. sp: Groves, 1997: 7.

Diagnosis: Pyriform Protocypraea, posterior terminal

ridges forming slight marginal callus; fossula concave,

smooth.

Page 48

THE NAUTILUS, Vol. 118, No. 1

Description: Shell of medium size, moderately inflat-

ed, elongate and somewhat constricted anteriorly; spire

partially covered by successive whorls; dorsum moder-

ately arched; maximum height and width slightly poste-

rior of center; aperture somewhat straight, narrow; teeth

absent from both outer and inner lips; fossula concave,

smooth, wide; posterior basal terminal ridges forming

slight marginal callus; anterior and posterior terminal ca-

nals shallow.

Comparison: The new species is most similar to Ber-

naya (Protocypraea) gualalaensis (Anderson, 1958: 176,

p 62, figs. 88a) fon the Upper Cretaceous (upper

Campanian to lower Maastrichtian) Gualala Formation,

informal Anchor Bay member of Wentworth (1966) (see

also Elder et al., 1998), Mendocino County, northern

California. Bernaya (Protocypraea) popenoei is smaller

than B. (P.) gualalaensis and has a narrower and straight-

er aperture, and more globose shape.

Discussion: Good preservation of the holotype per-

mits unequivocal generic and subgeneric assignments.

Although the Upper Cretaceous =e of the Santa Ana

Mountains: Orange County, California contain abundant

mollusks (Packard, 1922; Popenoe, 1937, 1942; Saul,

1982, 1996), B. (P.) popenoei is the only cypraeoidaen so

far described from the Ladd Formation.

Material: Represented by a single well preserved

specimen that exhibits original-shell material.

Type Material: Holotype, UCMP 154951, measures

31.4 mm in length, 20.2 mm in width, and 17.8 mm in

height.

Type Locality: UCMP loc. A3404, Lucas Canyon,

Santiago Peak quadrangle, Santa Ana Mountains,

Orange County, California, Upper Cretaceous (lower

Campanian), Ladd Formation, Holz Shale Member.

Etymology: Named for the late Willis Parkinson

(“Parky”) Popenoe (University of California, Los Ange-

les), in recognition of his numerous significant contri-

butions to Cretaceous paleontology and str: atigraphy of

the Santa Ana Mountains, Orange County, California.

NEW RECORDS OF PACIFIC SLOPE

CRETACEOUS CYPRAEIDS

Bernaya (Bernaya) crawfordcatei Groves, 1990

New Record: LACMIP loc. 17198, west side of Bee

Canyon, El Toro quadrangle (1949 ed.), Santa Ana

Mountains, Orange County, California. Upper Creta-

ceous (upper lower Campanian), Williams Formation,

Pleasants Sandstone Member. Poorly preserved internal

mold.

Distribution: Formerly restricted to the B. (B.) craw-

fordcatei type locality (SDSNH loc. 3392), Upper Cre-

taceous ( uppermost Campanian to lowermost Maastrich-

tian), Point Loma Formation, near Carlsbad, northern

San Diego County, California.

Bernaya (Protocypraea) gualalaensis (Anderson, 1958)

New Record: USGS Mesozoic loc. M8829 north side

of Haven'’s Neck, Mendocino County, California. Upper

Cretaceous (upper Campanian to lower Maastrichtian),

Gualala Formation, informal Anchor Bay member. Two

fairly well preserved internal molds were illustrated by

Elder et al. (1998: 152, 163, pl. IL figs. 2-3, 6).

Distribution: Type locality (CAS loc. 61918), near

Gualala, Mendocino County, to the Carlsbad area, north-

erm San Diego County, California (SDSNH locs. 3162,

3162-A, 3162-B, 3162-M, 3392, 3405, and 3454), Upper

Cretaceous (uppermost Campanian to lowermost Maas-

trichtian), Point Loma Formation.

Palaeocypraea sp.

Y) I

New Record: LACMIP loc. 10441 (ex CIT loc. 1396).

Sucia Island, San Juan County, Washington. Upper Cre-

taceous (lower Campanian), Cedar District Formation.

Single, fairly well preserved, slightly dorso-ventrally

crushed, juvenile specimen.

Cypraeidae, undetermined genus and species.

New Record: LACMIP loc. 17421, Palmer Way,

Carlsbad, San Diego County, California. Upper Creta-

ceous (uppermost Campanian to lowermost Maastrich-

tian) Point Loma Formation. Single poorly preserved in-

ternal mold.

ACKNOWLEDGMENTS

Eric Gohre, Oroville, California, generously loaned spec-

imens from his personal collection for this project, do-

nated type material to LACMIP, and arranged for access

to and accompanied the author on private property along

Dry Creek, near Pentz, Butte County, California. With-

out his collecting skills, enthusiasm, and generosity, a

major portion of this paper would not have been possi-

ble. Richard P. Hilton (Sierra College, Rocklin, Califor-

nia), LouElla R. Saul (LACMIP), and Richard L. Squires

(California State University, Northridge, Geological Sci-

ences) also accompanied the author in the field. Special

thanks to Graham Beard (VIPM) and David Lindberg

(UCMP) for specimen loans from their respective insti-

tutions. LouElla R. Saul (LACMIP) expertly removed

matrix from several specimens, discussed stratigraphic

problems, and allowed access to her personal library. Pe-

ter S. Mustard (Simon Fraser University, Burnaby, Brit-

ish Columbia) and Randy J. Emkin (Geological Survey

of Canada, Sydney, British Columbia) are thanked for

reprints and detailed discussions of Haslam Formation

age and paleomagnetism of the Nanaimo Basin. I thank

Tom Cockburn, Victoria Palaeontological Society, British

Columbia for his assistance in acquiring key references

and making important contacts. Ellen Kim (LACM Mal-

acology volunteer) assisted with digital photography and

L. T. Groves, 2004

Angel Valdés (LACM Malacology) kindly assisted with

digital image manipulations.

Don McNamee and Mali Griffin (Natural History

Museum of Los Angeles County, Research Library) ex-

pertly assisted in locating obscure references. Many

thanks to LouElla R. Saul (LACMIP), Richard L.

Squires (CSUN), and James H. McLean and Angel Val-

dés (LACM Malacology) for reviewing an early dient of

the manuscript. The evaluations of James G. Haggart

(Geological Survey of Canada, Vancouver, British Co-

lumbia) and an anonymous reviewer greatly enhanced

this paper.

LITERATURE CITED

Anderson, F. M. 1958. Upper Cretaceous of the Pacific Coast.

Geological Society of America, Memoir 71, 378 pp., 75

Is.

oun C. H. 1912. Geology of Nanaimo Sheet, Nanaimo coal-

field, Vancouver Island, British Columbia. Summary Re-

port of the Geological Survey Branch of the Department

of Mines for 1911: 91-105.

Deshayes, G. P. 1824-1837. Description des coquilles fossiles

des environs de Paris. F-G. Levrault, Paris, 1 [Conchi-

fers]: 1-392 [1824]; 2 [Mollusques]: 1-783 [1824-1835];

Atlas, pls. 1-101 [1837].

Elder, W. P. and L. R. Saul. 1993. Paleogeographic implications

of molluscan assemblages in the Upper Cretaceous (Cam-

panian) Pigeon Point Formation, California. In: Dunn, G.

and McDougall, K. (eds.) Mesozoic paleogeography of the

western United States—II. Society of Economic Paleon-

tologists and Mineralogists, Pacific Section Guidebook 71:

171-186, figs. 1-4, pls. 1-2.

Elder, W. P., L. R. Saul and C. L. Powell Il. 1998. Late Cre-

taceous and Paleogene fossils of the Gualala Block and

their paleogeographic implications. In: Elder, W. P. (ed.),

Geology and tectonics of the Gualala Block, northern Cal-

ifornia. Society of Economic Paleontologists and Miner-

alogists, Pacific Section Guidebook 84: 149-168, figs. 1-

B, jak i.

Enkin, R. ]., J. Baker and P. S. Mustard. 2001. Paleomagnetism

of the Upper Cretaceous Nanaimo Group, southwestern

Canadian Cordillera. Canadian Journal of Earth Sciences

38: 1403-1422, figs. 8),

Gabb, W. M. 1864. Description of the Cretaceous fossils. Geo-

logical Survey of California, Palaeontology 1: 57-217, pls.

9-32,

Gmelin, J. F. 1791. Caroli a Linné Systema naturae per regna

tria naturae. Editio decimo tertia. 1(6): 3021-3910, Leip-

zig, Germany.

Gray, J. E. 1824-1828. Monograph on the Cypraeidae, a Fam-

ily of Testaceous Mollusca. Zoological Journal 1: 71-80,

137-152 [1824]; 1: 367-391 [1824]: 1: 489-518 [1825]; 3:

363-371 [1827]; 3: 567-576; 4: 66-88 [1828].

Groves, L. T. 1990. New species of Late Cretaceous Cyprae-

acea (Mollusca: Gastropoda) from California and Missis-

sippi, and a review of Cretaceous cypraeaceans of North

America. The Veliger 33: 272-285, figs. 1-34.

Groves, L. T. 1994. Jurassic and Cretaceous cypraeacean bio-

geography and paleontology with an annotated list of the

species. The Cowry, new series 1: 25-41, figs 1-20.

Groves, L. T. 1997. Fossil and Recent species of eastern Pacific

Cypraeacea (Cypraeidae and Eocypraeinae [Ovulidae]):

Page 49

An update [extended abstract]. Western Society of Mala-

cologists, Annual Report 29: 7-10.

Haggart, J. W. 1991. Biostratigraphy of the Upper Cretaceous

Nanaimo Group, Gulf Islands, British Columbia. In:

Smith, PL. (ed.), A field guide to the paleontology of

southwestern Canada. University of British Columbia,

Vancouver, pp. 223-257.

Haggart, J. W., L. R. Saul, R. Watkins and E. S. Gohre. 1997.

Cretaceous shallow marine strata at Pentz, California. In:

Erskine, M. and D. Lawler (eds.) Northern Sierra Nevada

Region Geological Field trip Guidebook, Norther Cali-

fornia Geological Society, pp. 1-6.

Jousseaume, F. Pp 1884. Etude sur la famille des Cypraeidae.

Bulletin de la Société Zoologique de France 9: 81-100.

Kodama, K. P. and P. D. Ward. 2001. Compaction-corrected

paleomagnetic paleolatitudes for Late Cretaceous rudists

along the Cretaceous Califomia margin: Evidence for less

than 1500 km of post-Cretaceous offset for Baja British

Columbia. Geological Society of America Bulletin 113:

1171-1178, figs. 1-3.

Ludvigsen, R. and G. Beard, G. 1994. West coast fossils: A

guide to the ancient life of Vancouver Island. Whitecap

Books, Vancouver, xii + 194 p., 130 figs.

Ludvigsen, R. and G. Beard. 1997. West coast fossils: A guide

to the ancient life of Vancouver Island, Second edition.

Harbour Publishing, Madeira Park, British Columbia, 216

pp., 157 figs.

Muller, J. E. and J. A. Jeletzky. 1970. Geology of the Upper

Cretaceous Nanaimo Group, Vancouver Island and Gulf

Islands, British Columbia. Geological Survey of Canada

Paper 69-25: 1-77.

Mustard, P., J. Haggart, D. Katnick, K. Treptau and J. Mac-

Eachem. 2003. Sedimentology, paleontology, ichnology

and sequence stratigraphy of the Upper Cretaceous Nee

naimo Group submarine fan deposits, Denman and Horn-

by islands, British Columbia. In: Geological field trips in

southern British Columbia. Geological Association of Can-

ada Cordilleran Section, pp. 103-145, figs, 1-27.

Nilsen, T. H. 1984. Tectonics and sedimentation of the Upper

Cretaceous Hombrook Formation, Oregon and California.

In: Crouch, J. K. and Bachman, S. B. (eds.), Tectonics and

sedimentation along the California margin. Society of

Economic Sedimentologists and Paleontologists, Pacific

Section Guidebook 42: 101-118, figs. 1-7.

Oliveira, P. E. de. 1957. Invertebrados Cretacicos do fosfato

de Pemambuco. Divisao de Geologia e Mineralogia Rio

de Janeiro, Boletim 172: 1-29, pls. 1-2.

Oyama, K. 1959. Review of nomenclature on Japanese shells

(3). Venus 20: 361-362.

Packard, E. L. 1922. New species from the Cretaceous of the

Santa Ana Mountains, Califomia. University of California

Publications, Bulletin of the Department of Geological

Sciences 13: 413-462, pls. 24-38.

Popenoe, W. P. 1937. Upper Cretaceous Mollusca from south-

em Califomia. Journal of Paleontology 11: 379-402, pls.

45-49.

Popenoe, W. P. 1942. Upper Cretaceous formations and faunas

of southern California. Bulletin of the American Associa-

tion of Petroleum Geologists 26: 162-187, figs. 1-4.

Rafinesque, C. S. 1815. Analyse de la nature, ou tableau de

Yunivers et des corps organisés. Palermo, 224 pp.

Russell, J. S., S. L. Baum and R. Watkins. 1986. Late Coniacian

to Early Campanian clastic shelf deposits and molluscan

assemblages of the northeastern Sacramento Valley, Cali-

Page 50

fornia. In: Abbott, P. L. (ed.), Cretaceous stratigraphy,

western North America. Society of Economic Paleontol-

ogists and Mineralogists, Pacific Section Guidebook 46:

179-196, figs. 1-16.

Saul, L. R. 1982. Water depth indications from Late Creta-

ceous mollusks, Santa Ana Mountains, California. In: Bott-

jer, D. J. and others (eds.), Late Cretaceous depositional

environments and paleogeography, Santa Ana Mountains,

southern Califomia. Society of Economic Paleontologists

and Mineralogists, Pacific Section Guidebook 24: 69-75,

figs. 1-3, 4 unnumbered figs.

Saul, L. R. 1996. Three new Turonian muricacean gastropods

from the Santa Ana Mountains, southern California. The

Veliger 39: 125-135.

Schilder, F. A. 1927. Revision der Cypraeacea (Moll., Gastr.).

Archiv fiir Naturgeschichte 91A(10): 1-171.

Schilder, F. A. 1928. Die Cypraeacea des Daniums von Da-

nemark und Schonen. Danmark Geologiske Un-

dersggelse, ser. 4, 2(3): 1-29, figs. 1-16.

Schilder, F. A. 1932. Cypraeacea. In: W. Quenstedt (ed.), Fos-

silium Catalogus, I: Animalia, pt. 55. W. Junk, Berlin, 276

editte M. and F. A. Schilder. 1971. A catalogue of fossil and

living cowries. Institut Royal des Sciences Naturelles de

Belgique, Mémoire 85: 1-246.

Schlotheim, E. F. 1820-1823. Die petrefactenkunde auf ihrem

jetzigen Standpunkte durch die Beschreibung seiner

Sammlung versteinerter und fossilier iiberreste des Thei-

rund Pflanzenreichs der Vorwelt erliutert, Becker’schen

Buchhandlung, Gotha. Iii + 437 pp. Atlas: pls. 15-29

[1820]; pls. 1-21 [1822]; pls. 22-37 [1823].

Squires, R. L. and L. R. Saul. 1997. Late Cretaceous occur-

rences on the Pacific slope of North America of the me-

lanopsid gastropod genus Boggsia Olsson, 1929. The Ve-

liger 40: 193-202.

Squires, R. L. and L. R. Saul. 2001. New Late Cretaceous

gastropods from the Pacific slope of North America. Jour-

nal of Paleontology 75: 46-65.

Trask, J. B. 1856. Description of a new species of ammonite

and baculite from the Tertiary rocks of Chico Creek. Pro-

ceedings of the California Academy of Sciences, Ist ser.

1, 92-93, pl. 2.

Vredenburg, E. W. 1920. Classification of the Recent and fossil

Cypraeidae. Records of the Geological Survey of India 2:

65-152.

Ward, P. D. 1978. Baculitids from the Santonian-Maestrichtian

Nanaimo Group, British Columbia, Canada and Washing-

ton State, USA. Journal of Paleontology 52: 1143-1154.

Wentworth, C. M. 1966. The Upper Cretaceous and lower Ter-

tiary rocks in the Gualala area, northern Coast Ranges,

California [Ph.D. dissertation]. Stanford University, Stan-

ford, 197 pp.

Whiteaves, J. F. 1895. On some fossils from the Nanaimo

Group of the Vancouver Cretaceous. Transactions of the

Royal Society of Canada, 2nd ser., 1: 119-133, pls. 1-3.

APPENDIX 1.

LOCALITIES CITED

CAS 61918 (ex S. G. Clark loc. 251). Near Gualala, sec.

27(?), TIN, R1ISW, MDMB, Gualala quadrangle, Men-

docino County, California. Upper Cretaceous (upper

THE NAUTILUS, Vol. 118, No. 1

Campanian to lower Maastrichtian), Gualala Formation.

Coll.: S. G. Clark.

LACMIP 10441 (ex CIT loc. 1396). Bluffs along south

side of Fossil Bay, south side and east end of Sucia Is-

land, San Juan County, Washington. Upper Cretaceous

(lower Campanian), Cedar District Formation. Coll.: R.

Durbin, H. L. and W. P. Popenoe, 23 Jul, 1935.

LACMIP 17198. Unsorted very indurated conglomerate

lens with pebble and cobble-sized clasts and a sandstone

matrix; at elevation 207 m, on west side of divide on

west side of Bee Canyon, 4496 m south and 3117 m east

of northwest corner of USGS El Toro quadrangle (1949

ed.), Santa Ana Mountains, Orange County, California.

Upper Cretaceous (upper lower Campanian), Williams

Formation, Pleasants Sandstone Member. Coll.: P. Peck

and others, 28 May, 1997. [Locality now inaccessible and

covered by Eastern Transportation Corridor].

LACMIP 17421. In sandstone immediately overlying

basal conglomerate and from spoil piles along south side

of commercial property at 5607 Palmer Way, Carlsbad,

San Luis Rey quadrangle (1975), San Diego County,

California. Upper Cretaceous (uppermost Campanian to

lowermost Maastrichtian), Point Loma Formation. Coll.:

G. L. Kennedy.

LACMIP 17611. Dry Creek, near Pentz, Butte County,

California. Upper Cretaceous (lower Campanian), infor-

mal Pentz Road member, Chico Formation. Coll.: E. S.

Gohre.

LACMIP 24081 (ex UCLA loc. 4081). South of Pentz,

Butte County, California. Upper Creatceous (lower

Campanian), informal Pentz Road member, Chico For-

mation. Coll.: T. Susuki.

SDSNH 3162. Carlsbad area, locality (now covered by

Faraday Avenue) was exposed during development of

Carlsbad Research Center, southwest of El Camino

Real, south of Letterbox Canyon and north of Palomar

Airport, 33°08'02” N, 117°16'41” W, San Luis Rey quad-

rangle, San Diego County, California. Upper Cretaceous

(uppermost Campanian to lowermost Maastrichtian),

Point Loma Formation. Coll.: B. O. Riney, T. A. De-

méré, and M. A. Roeder, Mar—May, 1982.

SDSNH 3162-A. Carlsbad area, at base of stratigraphic

section measured at SDSNH 3162, approximately 6.1 m

below a calcareous marker bed. Upper Cretaceous (up-

permost Campanian to lowermost Maastrichtian), Point

Loma Formation. Coll.: B. O. Riney, T. A. Deméré, and

M. A. Roeder, Mar—May, 1982.

SDSNH 3162-B. Carlsbad area, 2.1—-3.9 m below a cal-

careous marker bed in measured stratigraphic section at

SDSNH 3162. Upper Cretaceous (uppermost Campan-

ian to lowermost Maastrichtian), Point Loma Formation.

Coll.: B. O. Riney, T. A. Deméré, and M. A. Roeder,

Mar—May, 1982.

SDSNH 3162-M. Carlsbad area, near top of stratigraph-

L. T. Groves, 2004

Page 51

ic section measured at SDSNH 3162. Upper Cretaceous

(uppermost Campanian to lowermost Maastrichtian),

Point Loma Formation. Coll.: B. O. Riney, T. A. Deméré

and M. A. Roeder.

SDSNH 3392. Carlsbad area, north of Palomar Airport,

roadcut along west side of College Boulevard, approxi-

mately 424 m south of intersection with E] Camino Real,

33°08'21" N, 117°17'02” W, San Luis Rey quadrangle,

San Diego County, California. Upper Cretaceous (up-

permost Campanian to lowermost Maastrichtian), Point

Loma Formation. Coll.: SDSNH field party May, 1987.

SDSNH 3405. Carlsbad area, north of Palomar Airport,

excavation for College Boulevard, approximately 242—

485 m south of intersection with El Camino Real,

33°08'21" N, 117°17'02” W, San Luis Rey quadrangle,

San Diego County, California. Upper Cretaceous (up-

permost Campanian to lowermost Maastrichtian), Point

Loma Formation. Coll.: B. O. Riney, M. A. Roeder, and

R. Q. Gutzler, Apr-May, 1987.

SDSNH 3454. Carlsbad area, north of Palomar Airport,

excavation for College Boulevard, approximately 153 m

north of College Boulevard and Faraday Avenue inter-

section, 33°08'11” N, 117°17'02" W, San Luis Rey quad-

rangle, San Diego County, California. Upper Cretaceous

(uppermost Campanian to lowermost Maastrichtian),

Point Loma Formation. Coll: B. O. Riney and M. A.

Roeder, Apr-May, 1987.

UCMP A3404. Fossils in float boulders in Lucas Can-

yon, a branch of San Juan Canyon, near San Juan Cap-

istrano, Santiago Peak quadrangle, Santa Ana Moun-

tains, Orange County, California. Upper Cretaceous

(lower Campanian), Ladd Formation. September, 1916.

USGS Mesozoic loc. M8829. Conglomerate at northeast

end of beach on north side of Haven’s Neck in SW%4

SW% SE%, section 12, TI1N, RI6W, Mendocino Coun-

ty, Califomia. Upper Cretaceous (upper Campanian to

lower Maastrichtian), Gualala Formation, informal An-

chor Bay member. Coll.: W. P. Elder, 1992.

THE NAUTILUS 118(1):52-53, 2004

Book Review

Page 52

Checklist of the Land Snails and Slugs of

California

Roth, B. and P. S. Sadeghian. 2003. Checklist of the

Land Snails and Slugs of California. Santa Barbara Mu-

seum of Natural History Contributions in Science, Num-

ber 3, 81 pp., 13 color plates.

Creating lists is one of the most important things that

systematists do. By bringing together in a single place

nomenclatural, bibliogr: aphic, distal bwttomnell aa type

material information, each checklists or catalogs are fun-

damental to revisionary systematics, to biogeogr aphy,

and to biodiversity conservation.

Checklists are primarily works of nomenclature in that

they are compilations of names, with varying amounts of

additional information associated with each name. They

can a lists of all the names in a particular higher taxon

(e.g., the family Partulidae—Richardson, 1990), al-

Tough often they are also geographically constrained

(e.g., the recent catalog of the New World Ampullari-

idae—Cowie and Thiengo, 2003). Some catalogs list all

the names introduced by particular authors (eg., those

of Clench and Turner—Johnson, 2003), perhaps restrict-

ed to a higher taxon of interest (e.g., the Neritidae de-

scribed by Récluz—Kabat and F men 1992). And some

catalogs are compilations of all names in a particular fau-

na, as in the case of the present work on the land gas-

tropod fauna of California. These last are the most valu-

able checklists for conservation because in order to con-

serve a fauna it is necessary to know as well as possible

which species constitute that fauna. Checklists such as

this one provide that essential baseline of knowledge.

In addition to being a compilation of all the names of

land snails and slugs, including non-native species,

known from California, this checklist includes details of

name-bearing type material, and the county-by-county

distributions of the recognized species and subspecies.

Non-native species are clearly identified by asterisks.

The checklist is also in a sense a work of taxonomy. Rath-

er than listing all names separately, with no expressed

opinion as to dnetir synonymy, synonyms (including some

new synonyms) are listed under the species/subspecies

with which they are synonymized. The species are listed

alphabetically under the appropriate genera, and genera

are listed alphabetically under the appropriate higher

taxa. Some new species/genus combinations are intro-

duced.

A bonus, not often seen in such checklists, is the set

of beautiful color plates illustrating, mostly for the first

time, holotypes and lectotypes of 64 land snail taxa de-

scribed by S. Stillman Berry and deposited in the Santa

Barbara Museum of Natural History.

A total of 279 extant species and 112 subspecies are

listed, plus 6 species known only as fossils, for an overall

total of 397 valid taxa. Of these, 20 are as yet unde-

scribed species and subspecies, and 37 are non-native

species in California. Numerous synonyms are also list-

ed. Distributional information is based on extensive re-

search in museum and private collections. As is made

clear, however, even a fauna that might be expected to

be better known than many, in fact probably includes

many species remaining to be discovered, including

cryptic species that may only be discovered using mo-

lecular techniques; and field work will probably extend

the recorded distributions of the known species beyond

those reported in this checklist.

This is a valuable work that gives us just a tantalizing

taste of what is to come in a manual of the land snails

and slugs of California, currently in preparation by Roth.

However, I cannot end without mentioning a very small

number of essentially minor criticisms. First, I would

have liked to have had an index—I went to find one

particular genus and had to thumb through the book to

locate it, but more importantly, the lack of an index

makes it difficult to locate synonyms. Second, it would

have been nice to have had a bibliography of the original

descriptions, but this will no doubt be available in the

forthcoming manual. Finally, I understand the logic be-

hind the authors’ reluctance to use ‘formal [taxonomic]

ranks above the genus’, but in fact they do use the ‘tra-

ditional names of taxa’ above the genus throughout the

checklist, which allows the more traditional among us to

follow it. Among the Helminthoglyptidae, however, they

also use Roth’s (1996) rather obscure Iererenieel cla-

distic nomenclature—difficult to understand in part be-

cause, given the way the book is laid out, it is difficult

to see how far the various clade names are indented,

which is crucial to understanding their relationships. But

these are quibbles. This is a useful and important book,

both for systematists working with the groups repre-

sented in California, and especially as a foundational

documentation of California’s biodiversity that will be an

important tool in its conservation.

LITERATURE CITED

Cowie, R. H. and S. C. Thiengo. 2003. The apple snails of the

Americas (Mollusca: Gastropoda: Ampullariidae: Asolene,

Felipponea, Marisa, Pomacea, Pomella): a nomenclatural

and type catalog. Malacologia 45: 41-100.

Johnson, R. I. 2003. Molluscan taxa and bibliographies of Wil-

liam James Clench and Ruth Dixon Turner. Bulletin of

the Museum of Comparative Zoology 158(1): 16.

Kabat, A. R. and Y. Finet. 1992. Catalogue of the Neritidae

(Mollusca: Gastropoda) described by Constant A. Récluz

including the location of type specimens. Revue Suisse de

Zoologie 99: 223-253.

Book Review, 2004 Page 53

Richardson, C. L. 1990. Partulidae: Catalog of species. Tryonia Robert H. Cowie

15: i + 1-96. Center for Conservation Research and Training

Roth, B. 1996. Homoplastic loss of dart apparatus, phylogeny University of Hawaii

of the genera, and a phylogenetic taxonomy of the Hel- 3050 Maile Way, Gilmore 408

minthoglyptidae (Gastropoda: Pulmonata). The Veliger Honolulu, HI 96822, USA

39: 18-42. [email protected]

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A3Z(4

WA ey yee

Volume 118, Number 2

June 29, 2004

ISSN 0028-1344

A quarterly devoted

to malacology.

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Department of Invertebrates

Field Museum of

Natural History

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Invertébrés Marins et Malacologie

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and Training

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CONTENTS

Nie Wall) -S

Volume 118, Number 2

June 29, 2004

ISSN 0028-1544

John Slapcinsky

Brian Coles

Robert H. Cowie

Néstor J. Cazzaniga

Matthias Glaubrecht

Sven N. Nielsen

Revision of the genus Pilsbryna (Gastropoda: Pulmonata:

Gastrodontidae) and comments on the taxonomic status of

Pilsoryme erdgiens Mlommsom, IGS. 1.0 000caeacccasecoccsceconasoecnceses OD

The South American Mollusca of Johann Baptist Ritter von

Spix and their publication by Johann Andreas Wagner.................... (all

The genus Olivancillaria (Gastropoda: Olividae) in the

Miocene of Chile: rediscovery of a senior synonym and

Glesorijattlomn Git A MEW SOSCHES, ooo ccvecceccosocascocogansovonsnvc0eGnoS 88

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THE NAUTILUS 118(2):55-70, 2004

Page 55

Revision of the genus Pilsbryna (Gastropoda: Pulmonata:

Gastrodontidae) and comments on the taxonomic status of

Pilsbryna tridens Morrison, 1935

Brian Coles

John Slapcinsky

Florida Museum of Natural History

4202 Scottie Smith Drive

University of Florida Jefferson, AR 72079 USA

Gainesville, FL 32611 USA

[email protected]

ABSTRACT MATERIALS AND METHODS

Pilsbryna is revised based on new material collected during

recent surveys of wet leaf-litter microhabitats in the southern

Appalachian Mountains. Five species are recognized including

Pilsbryna aurea Baker, 1929, P. castanea Baker, 1931, P. no-

dopalma new species and P. quadrilamellata new species. All

species are redescribed or described. Pilsbryna vanattai (Walk-

er and Pilsbry, 1902) is transferred to Pilsbryna from Glyphy-

alinia (Glyphyalus) based on genital and juvenile shell anato-

my. Pilsbryna tridens Morrison, 1935, is reexamined based on

newly available material; its placement in the genus Helicod-

iscus sensu lato is supported by new radular evidence. Pilsbry-

na species share unique genital and shell characteristics that

are included in a redescription of the genus. The new generic

definition combined with habitat information for all Pilsbryna

species allows a better understanding of the geographic and

microhabitat distribution of the genus.

INTRODUCTION

Although the terrestrial invertebrate fauna of eastern

North America is among the best known in the world,

new species are discovered frequently. Most are small

species from patchily distributed and poorly sampled mi-

crohabitats. Recent collecting in one such habitat, damp

to wet leaf-litter surrounding mountain seeps, springs

and streams, has uncovered a remarkable radiation of

small terrestrial snails of the genus Pilsbryna. In this

paper, we redescribe the genus, describe two new spe-

cies of Pilsbryna, and redescribe the other three species

in the genus including a species formerly placed in the

genus Glyphyalinia. One other previously described spe-

cies, P. tridens, was reassigned to Helicodiscus by Hu-

bricht (1964) without explanation. This generic place-

ment was used in most subsequent publications listing

the species (Riedel, 1980; Hubricht, 1985; Turgeon et

al., 1988, 1998) although it has remained unconfirmed

until now. Newly collected material of H. tridens allows

for its unequivocal generic assignment.

Specimens were hand-collected from the leaf-litter/soil

interface or sifted from samples of leaf-litter. Live col-

lected animals were narcotized overnight in a suspension

of water containing a 1 cm length of mentholated ciga-

rette and then preserved in 75% ethanol. Gross anatom-

ical dissections were made under 75% ethanol using a

dissecting microscope. Isolated reproductive systems

were stained with Harris haematoxylin and Semichon

aceto-carmine stain and cleared with glycerin. Radulae

were isolated from dissected buccal masses using a sat-

urated KOH solution. Scanning electron micrographs of

shells and radulae were made using a field emission-

SEM. Drawings of the shell and genital anatomy were

made with the assistance of a camera lucida, and mea-

surements were taken using an ocular micrometer. All

line drawings of the internal structures of juvenile shells

are in basal view and use solid lines for shell outlines

and internal structures that make contact with the inner

surface of the translucent base of the shell. Dotted lines

indicate structures that are viewed through the translu-

cent base of the shell but do not make contact with the

base. Shell measures were made as follows. Whorl count

(W) was measured from the suture of the first whorl to

the body whorl (Figure 1) and fractions of a whorl were

determined with the aid of a cardboard circle divided

into 10 equal parts of 36°. Spire diameter (SD) was mea-

sured as the length of a straight line passing from the

apertural edge of the suture through the middle of the

apex to the opposite suture (Figure 1, line a—b). Height

(H) was the greatest distance between the apex and the

base of the aperture measured parallel to the shell axis

(Figure 2, line c-d). Aperture height (AH) was measured

from the suture to the base of the aperture, parallel to

the shell axis (Figure 2, line e—f). Aperture width (AW)

was the greatest distance from the apertural edge of the

umbilicus to the outer edge of the aperture (Figure 2,

line gf). Greater diameter (GD) was the measure of

the greatest width of the shell (Figure 3, line h-i), and

lesser diameter (LD) was the diameter perpendicular to

Page 56

THE NAUTILUS, Vol. 118, No. 2

1 a8 2

3 J

Figures 1-3. Shell measurements: 1. Whorl count (1-4.9), spire width (line A-B). 2. Shell height (line C-D), aperture height

(line E-F), aperture width (line F—G). 3. Greater diameter (line H-I), lesser diameter (line I-]), umbilical width (line K-L).

the greater diameter (Figure 3, line i-j). Umbilical

Width (UW) was measured from the inner edge of the

aperture through the center of the umbilicus to the op-

posite side of the umbilicus (Figure 3, line kl). Maps

were made with the assistance of DeLorme, Topo USA

software. We examined specimens of Pilsbryna in the

collections of the Academy of Natural Sciences, Phila-

delphia (ANSP), Field Museum of Natural History, Chi-

cago (FMNH), North Carolina State Museum, Raleigh

(NCSM), National Museum of Natural History (USNM)

and Florida Museum of Natural History, Gainesville

(UF) and the private collection of one of the authors

(BC).

The following abbreviations are used in figures of gen-

ital anatomy: AG = albumen gland, EP = epiphallus,

HD = hermaphroditic duct, OV = free oviduct, PE =

penis, PG = prostate gland, PP = penial papillae, PR

= penial retractor muscle, SD = spermathecal duct, SP

= spermatheca, UT = uterus, VD = vas deferens. An-

atomical terminology follows Pilsbry (1946); for alternate

usages see Tompa (1984).

SYSTEMATICS

Family Gastrodontidae Tryon, 1866

Genus Pilsbryna Baker, 1929

Type Species: Pilsbryna aurea Baker, 1929 by mono-

SINE

Description: Small snails of the family Gastrodonti-

dae with depressed-helicoid and umbilicate shells of

roughly 5 whorls that are sculptured with closely placed,

shallow axial grooves. Whorl expansion is slow and reg-

ular through the penultimate whorl and more rapid in

the body whorl. Shells of juveniles usually contain spi-

rally arranged lamellae or nodules within the shell.

These lamellae or nodules occur at mid parietal and var-

ious other positions from columellar to sutural, and are

reduced or completely resorbed in adult shells.

The apical %4 to % of the penis bears papillae that are

not located within a well-defined apical chamber. The

base of the penis is thin-walled and simple. The penial

retractor muscle is inserted near the apex of the penis.

The epiphallus is robust, of similar diameter to the pe-

nis, does not bear a caecum and joins the penis laterally

at the apex. The epiphallus is well defined at the junction

with the vas deferens. The spermathecal duct is long and

slender and the spermatheca ovate. There are no well-

defined glandular areas within the free oviduct.

The form of the radula is typical of many small North

American Gastrodontidae. The centrals are symmetric

and tricuspid and of similar height to the first laterals.

There are three asymmetric tricuspid laterals on each

side of the central, and numerous unicuspid marginals.

Remarks: Pilsbryna has been treated as a subgenus of

Paravitrea by Riedel (1980); however, similarities be-

tween the two genera are superficial or shared by several

other groups of North American Gastrodontidae. Al-

though both Pilsbryna and Paravitrea possess internal

barriers in the shell, the form and position of these bar-

riers differs. Juvenile Pilsbryna possess spiral lamellae or

series of nodules, usually at the parietal, palatal and basal

positions. Paravitrea species usually have axial rows of

two to many, evenly spaced, small lamellae grouped 4

to % whorl apart that are not concentrated near the ap-

erture. The shells of Paravitrea are more tightly coiled,

usually with 6 or more whorls while shells of Pilsbryna

attain fewer than 6 whorls and more closely resemble

those of some Glyphyalinia and Nesovitrea. Analysis of

genital characters also does not support the placement

of Pilsbryna within the genus Paravitrea. The genital

anatomy of all known species of Pilsbryna is highly con-

served and the combination of genital characters found

in Pilsbryna are not found in Paravitrea or in any other

group of North American Gastrodontidae. Similarities in

radular anatomy between Pilsbryna and Paravitrea are

not unique; they are shared with most small American

Gastrodontidae, including Glyphyalinia and Nesovitrea.

The unique apertural barriers of the shell, the rows of

papillae at the apex of the penis and the unusual habitat

shared by Pilsbryna species strongly suggest the group

is monophyletic and distinct from other genera of small

American Gastrodontidae. In any case, evidence of a

particularly close relationship between Pilsbryna and

Paravitrea is lacking, and Pilsbryna is here considered a

distinct genus.

J. Slapeinsky and B. Coles, 2004

Pilsbryna aurea Baker, 1929

Common name: ornate bud

(Figures 4-10, Table 1)

Pilsbryna aurea Baker, 1929a: 91-92, pl. 3, figs. 4-8; Baker,

1929b: 260-261, pl. 9, figs. 4-5; Baker, 1931: 112-113, pl.

19, figs. 9-13, pl. 20, fig. 5; Pilsbry, 1946: 389-391, fig.

205(4—5), fig. 206(a—c), fig. 207(9-13), fig. 208(5); Baker,

1962: 3; Burch, 1962: 107, fig. 257; Hubricht, 1973: 14:

Turgeon et al., 1988: 135; Turgeon et al., 1998: 148.

Diagnosis: A medium-sized Pilsbryna with an adult

shell diameter of 2.9-3.6 mm, height of 1.4-1.9 mm and

5.0-5.4 whorls. Shells of less than 3.5 whorls contain two

lamellae, one crescent-shaped at mid-columellar posi-

tion, the second blade-shaped or undulate at mid-pari-

etal position. Both lamellae extend up to % whorl into

the body whorl. The parietal lamella is often thickened

along its distal edge giving it a T-shaped cross section.

Description: Shell depressed-helicoid, umbilicate,

glossy, translucent and densely sculptured with irregu-

larly spaced, indented axial lines (Figures 4-6). Adult

shell (Table 1) about 2.9-3.6 mm (mean = 3.2, n = 10)

in major diameter and 1.4-1.9 mm (mean = 1.7) in

height with 5.0-5.4 (mean = 5.1) slowly expanding

whorls. The shell height/greater diameter ratio is 0.48—

0.56 (mean = 0.52) and the spire width/greater diameter

ratio is 0.53-0.60 (mean = 0.57). The final % of the body

whorl expands slightly more rapidly than previous

whorls; the lesser/greater diameter ratio is 0.82-0.91

(mean = 0.86). The aperture is ovate, widest at or slight-

ly below the middle of the whorl. The umbilicus is 0.4—

0.6 mm (mean = 0.5) in diameter and 0.13—0.19 (mean

= 0.16) of the greater diameter of the shell. The funnel

shaped umbilicus expands regularly until the final third

of the body whorl, where it expands more rapidly. Shells

of immature animals contain two lamellae, a crescent

shaped lamella at mid-columellar position and an un-

dulating lamella at mid-parietal position (Figures 7-8).

Both lamellae extend up to % whorl into the aperture.

The parietal lamella is usually thickest distally and is of

ten T shaped in cross section. Both lamellae are much

reduced and more often completely resorbed in adults.

The basal half of the penis is straight-sided and simple

while the apical half is robust, bearing numerous small

papillae (Figure 9). The penial retractor muscle is sub-

apically inserted. The epiphallus is about the same

length and roughly half the diameter of the penis and

inserts laterally at the apex of the penis. The diameter

of the epiphallus remains roughly constant and widens

only slightly at the junction with the narrow vas defer-

ens. The epiphallus is folded at the mid-point and also

has longitudinal folds internally. The spermatheca is

ovate, and the narrow spermathecal duct expands very

slightly basally. The free oviduct is roughly twice the di-

ameter of the base of the penis, expanding slightly at the

junction with the spermathecal duct. A caecoid outpock-

__eting was not observed on the free oviducts of two dis-

sected adults.

The central tooth of the radula is symmetrically tri-

Page 57

cuspid; the mesocone is slender, especially basally, ex-

panding slightly above the ectocone cusps and then ta-

pers slowly to the apex (Figure 10). The ectocones are

short, roughly % the total height of the tooth, symmet-

rical and diamond-shaped. The three laterals are tall,

slender, and asymmetrically tricuspid. The endocone is

tall about % the height of the mesocone. The ectocone

is % the height of the mesocone and separated from it

by a narrow gap. The marginals are tall, slender and

unicuspid, with concave peripheral edges and convex

proximal edges. We studied two radulae (UF 287063)

using SEM microscopy. Both had twenty marginals on

each side.

Holotype: ANSP 147189a. Baker (1929a) figured and

measured this shell and designated it as the type, there-

by fixing the holotype.

Paratypes: ANSP 147189, type locality, Baker, 1928.

Type Locality: USA, Tennessee, Unicoi County,

Limestone Cove, Big Springs, between Unaka and Stone

Mountains, about 11 km E of Unicoi, 36° 11’ N, 82° 17’

W, elevation 700 m, Baker, 1928.

Other Material Examined: ANSP 158890, USA,

Tennessee, Unicoi County: Limestone Cove; ANSP

152469, type locality, Baker, 1928; ANSP 165583, Baker,

26-30 Aug. 1928; UF 292089, UF 287063, Davis

Springs, 8 km E of Unicoi, 36°10.7’ N, 82°16.4’ W, 687

m elevation, J. Slapcinsky and B. Coles, 31 May 2001;

FMNH 248868, near Davis Springs, 8 km E of Unicoi,

L. Hubricht, 18 May 1961; FMNH 248867, 2.1 km SE

of Limestone Cove, L. Hubricht, 28 May 1974; BC

6761, Washington County, Dry Creek Road, 1 km S of

Jim McNeese Road, 36° 15.6’ N, 82° 21.9’ W, 637 m

elevation, B. Coles, 21 May 2002.

Remarks: Several characters of the genital and radular

anatomy differed from previously published descrip-

tions. A caecoid outpocketing was not observed on the

free oviducts of two adults dissected in this study; in

contrast, Baker (1929b) recorded it in a juvenile. How-

ever, reproductive structures, especially those of the pos-

terior portion of the reproductive system, can vary in

size, shape, color and texture with maturation and re-

productive stage (Emberton, 1985). Our observations of

the structures of the anterior portion of the reproductive

system compare well with those of Baker (1929b). This

study using SEM microscopy found twenty marginals

per side. In contrast, Baker (1929b), using light micros-

copy, recorded only 14 marginals in an immature spec-

imen.

Pilsbryna aurea appears most similar to Pilsbryna

quadrilamellata described below, juvenile shells of these

two share unique crescent-shaped columellar lamellae

and lack spiral rows of nodules at any position. However,

the two species are easily separated using other juvenile

shell characters: P. awrea has a distally expanded parietal

lamella and does not possess the mid-basal and sutural

lamellae of P. quadrilamellata. Anatomically, P. aurea can

Page 58 THE NAUTILUS, Vol. 118, No. 2

Figures 4-10. Pilsbryna aurea. 4-6. UF 287063, diameter 3.2 mm. 7. FMNH 248868, diameter 2.6 mm. 8. UF 287063, diameter

2.1 mm. 9. Camera lucida drawings of genitalia, UF 292089, maximum width 5.9 mm; 10. UF 292089, horizontal field width =

217 wm.

J. Slapcinsky and B. Coles, 2004

Page 59

Table 1. Measurements in mm of undamaged adult shells of five species of Pilsbryna, N = 10. GD = greater diameter, LD =

lesser diameter, H = height, AW = aperture width, AH = aperture height, SW = spire width, UW = umbilicus width, W =

number of whorls.

GD LD AW AH SW UW W

P. aurea Mean= SD 32=02 28+02 17+02 15+=01 13+01 18+£01 05=01 51+01

Range 2.9-3.6 2.5-3.0 1.4-1.9 Lo=I7 1.2-1.4 1.7-2.0 0.4—0.6 5.0-5.4

P. castanea Mem = SID 872= O02 88= 01 2Oz= O01 Lvye Ol 1H = Ol Al=Ol OF 2Ol 58 = Oo

Range 3.5-3.9 3.0-3.4 1.9-2.2 1.6-1.8 1.4-1.6 2.0-2.3 0.6-0.8 5.0-5.4

P. nodopalma Mean= SD 30+£01 26+01 15+01 14+=01 12+01 172=01 05+=01 45201

Range Qisrll 2.42.8 1.4-1.6 eS eS) 1.2-1.3 1.6-1.8 0.40.5 43-4.6

P. quadrilamellata Mean = SD 32+02 28+02 17+01 15+£01 13+00 19+01 05+01 50+02

Range 2.8-3.4 DFS}. 1.5-1.8 1.4-1.6 1.3-1.4 1.8-2.2 0.4-0.6 ATO 2

P. vanattai Mean= SD 40+02 34+02 20+01 20+02 17+01 20+01 05+01 48 +02

Range 3.84.4 SWShT/ 1.9-2.2 1.8-2.3 ESS 1e9) GED 0.5-0.6 4.6-5.1

be distinguished from other species of Pilsbryna by the

unusually inflated apex of its penis and by the clearly

subapical insertion of its penial retractor muscle. The

teeth of the radula are tall and slender. The central tooth

is especially slender, and unlike other Pilsbryna species,

does not bear a strong angle at its widest point. The

peripheral ectocones are relatively poorly defined, sep-

arated from the adjacent mesocone by narrow gaps sim-

ilar to those of P. castanea and P. nodopalma, described

below, and unlike those of P. quadrilamellata and P. van-

attai.

Habitat and Distribution: Previously, Pilsbryna au-

rea was known from sites within 3 km of the town of

Limestone Cove in Unicoi County, Tennessee (Baker,

1929a; Hubricht, 1973). Recent collecting has also un-

covered this species along Dry Creek Road, south of

Johnson City, a range extension of more than 10 km to

the northwest. This species appears to be restricted to

mountain valleys near Unicoi, Tennessee, and has been

found at elevations of 600-700 m. Known populations

occur on rocky wooded hillsides along small streams.

The species is most common in wet leaf-litter along

streams and around seeps but is also found in deep leaf-

litter at the base of limestone and other sedimentary

rocks along stream banks.

Pilsbryna castanea Baker, 1931

Common name: prominent bud

(Figures 11-17, Table 1)

Pilsbryna castanea Baker, 1931: 111-112, pl. 19, figs. 1-8, pl.

20, fig. 4; Pilsbry, 1946: 391-393, fig. 207(1-8), fig. 208(4);

Baker, 1962: 5: Burch, 1962: 94, 98, 106, fig. 219, fig. 2298,

fig. 256; Hubricht, 1973: 14; Turgeon et al., 1988: 135;

Turgeon et al., 1998: 148.

Diagnosis: A medium to large Pilsbryna with an adult

shell diameter of 3.4-3.9 mm, height of 1.9-2.2 mm, and

whorl count of 5.1-5.4 whorls. Shells with fewer than 4

whorls have a sinuous parietal lamella and a spiral series

of basal nodules extending a full whorl into the aperture.

Shells 4.5 whorls contain only a few basal nodules rough-

ly half way into the body whorl and all traces of lamellae

are resorbed in most adults.

Description: The shell is depressed-helicoid and um-

bilicate, with its surface sculptured with dense and ir-

regularly spaced axial indentations (Figures 11-13). Pils-

bryna castanea is relatively high-spired (Table 1), with a

shell height of 1.9-2.2 mm (mean = 2.0, n = 10), width

of 3.4-3.9 mm (mean = 3.7) and height/width ratio of

0.51-0.58 (mean = 0.55). The shell is relatively tightly

coiled with 5.1-5.4 (mean = 5.2) whorls; the final % of

the body whorl expands slightly more rapidly than pre-

vious whorls and the ratio of lesser/greater diameter is

0.82-0.89 (mean = 0.86). The spire is 0.55—0.61 (mean

= 0.58) of the greater diameter. The umbilicus is rela-

tively straight-sided and 0.6-0.8 mm (mean = 0.7) wide;

the ratio of umbilical width/greater diameter is 0.17—

0.21 (mean = 0.19). Juvenile shells with fewer than four

whorls usually have a mid-parietal lamella and a spiral

row of low nodules at basal position (Figures 14—15).

Both often extend a full whorl or more into the aperture.

The position of the basal nodules coincides with the

summits of the undulating parietal lamella. Animals with

shells more than four whorls gradually stop producing

lamellae and begin to resorb previously deposited la-

mellae. The shells of animals approaching maturity often

contain traces of the lamellae a half whorl into the ap-

erture; by the time most animals reach maturity all trac-

es of apertural barriers are resorbed.

The penis is narrow and elongate, slightly constricted

above the base, expanding slowly apically, widest *4 from

the base, then tapering slowly to the apex (Figure 16).

The apical half bears small papillae. The penial retractor

muscle is inserted nearly apically. The epiphallus is lon-

ger than the penis, and joins it laterally at the apex. The

epiphallus is narrowest near the junction with the penis

is folded at mid-point, and slightly inflated towards the

vas deferens. The vas deferens is narrow; the junction

with the much wider epiphallus is well defined. Free

oviduct is slightly wider than the penis, where they meet

and widens at and above the attachment of the sper-

Page 60 THE NAUTILUS, Vol. 118, No. 2

diameter 2.4 mm. 16. Camera lucida drawings of genitalia, UF 297419, maximum width 5.8 mm. 17. UF 297419, horizontal field

width 205 wm.

J. Slapcinsky and B. Coles, 2004

mathecal duct. Spermathecal duct is long and narrow;

the spermatheca is ovate. One animal was dissected.

The central tooth of the radula is tall, slender and

tricuspid (Figure 17). The mesocone is elongate, widest

and angular slightly above the ectocone cusps and ta-

pering sharply towards the apex and narrowing towards

the base. The ectocones are slightly less than % the total

height of the tooth, symmetrical, and diamond-shaped.

The three lateral teeth are tall, slender and asymmetri-

cally tricuspid. The endocone of each is tall and the ec-

tocone short. The seventeen marginal teeth on each side

are tall, slender and unicuspid, with concave peripheral,

and convex proximal, edges. One radula was examined.

Holotype: ANSP 152468a. Baker (1931) figured and

measured this shell and designated it the type, thereby

fixing the holotype.

ANSP 152468, type locality.

Type Locality: USA, Tennessee, Marion County,

Dove, mouth of Cave Cove, west facing hillside south of

big spring, which forms eastern source of Battle Creek,

35°10’ N, 85°47’ W, 244 m elevation, Baker, 1931.

Other Material Examined: ANSP 165582, USA,

Tennessee: Marion County, type locality, Baker, 23-26

Jul. 1928; FMNH 171387, west side of Battle Creal at

junction of Ladd’s Cove Road and Interstate 24, 35°8.7

N, 85°46.7’ W, 193 m elevation, 4 Sep. 1974, G. Good-

friend; UF 297419, UF 297381, 9 Jun. 2002, J. Slapcin-

sky; FMNH 171403, Martin Springs 10 km south of

Monteagle, 4 Sep. 1974, G. Goodfriend; FMNH

248870, 8 km N of Sequatchie, near large spring, 23 Oct.

1962, L. Hubricht; UF 306530, Bledsoe County, Lusk

Loop Road, 0.5 km NW of Cannon Creek, 35°30.1’ N,

85°18.9' W, 300 m elevation, 1 Jun. 2003, J. Slapcinsky;

FMNH 171335, NE side of Rains Gulf, 18 km SW of

Pikeville, 410 m elevation, 7 Sep. 1974, G. Goodfriend;

ANSP 165581, Cannon Creek, W of Pikeville, 1928, H.

B. Baker.

Remarks: Like Pilsbryna nodopalma, described be-

low, and P. vanattai, juvenile P. castanea have spiral se-

ries of nodules. Pilsbryna castanea can be distinguished

from the other two species because it does not possess

a second series of nodules at the palatal position. The

nodules are peg-shaped and not dorsoventrally com-

pressed like those of P nodopalma. The nodules in P.

castanea can extend the entire body whorl, much farther

than those of the other nodulate species. However, this

last character is less useful when comparing older juve-

niles that have discontinued deposition of lamellae and

have begun to resorb previously deposited barriers. The

adult shells of P. castanea are larger than those of P.

nodopalma, have a greater height/width ratio than P.

vanattai, and have more whorls than either species.

Habitat and Distribution: Pilsbryna castanea is

found on wooded hillsides bordering Battle Creek and

the Sequatchie Valley in Marion and Bledsoe Counties,

Tennessee (H.B. Baker, 1931; Pilsbry, 1948; Hubricht,

Paratypes:

Page 61

1973, 1985). All animals were found in deep leaf-litter

at the base of limestone exposures. Live specimens are

most common in wet leaf-litter surrounding seeps.

Pilsbryna nodopalma new species

Common name: oar tooth bud

(Figures 18-24, Table 1)

Pilsbryna (unidentified): Lee, 1990: 7-8, fig. (unnumbered)

Diagnosis: A small and relatively loosely-coiled Pils-

bryna of 2.7-3.2 mm diameter and 1.4-1.6 mm height,

with 4.3-4.6 regularly expanding whorls. The shell

whorls are widest above the middle. The shells of ju-

venile animals contain an undulate parietal lamella and

three or four paired subcolumellar and lower palatal

nodules grouped near the aperture. These nodules are

dorsoventrally compressed.

Description: The shell is depressed-helicoid, umbili-

cate and translucent. The shell is small and loosely coiled

for the genus (Table 1); adults reach 2.7-3.2 mm (mean

= 3.0, n = 10) greater diameter and 1.4—1.6 mm (mean

= 1.5) height with 4.3-4.6 (mean = 4.5) whorls (Figures

18-20). Height is 0.47—0.53 (mean = 0.51) of the great-

er diameter. Whorl expansion is regular and increases

only slightly in the last third of the body whorl; the ratio

of lesser diameter to greater diameter is 0.83-0.90

(mean = 0.86). The shell surface is glossy and sculp-

tured with weak and irregularly spaced indented axial

lines. The apex is relatively flat and the whorls are widest

slightly above mid-point. Spire width is 0.53-0.60 (mean

= 0.58) of greater diameter. The sutures are relatively

deeply impressed for the genus. The umbilicus is nar-

row, 0.4-0.5 mm (mean = 0.45) roughly 0.13-0.17

(mean = 0.15) of shell diameter. Juvenile shells, with

less than four whorls, have an undulate parietal lamella

and two to four, paired, subcolumellar and lower palatal

nodules grouped near the aperture (Figures 21-22).

These nodules are dorsoventrally compressed giving

some of them the appearance of the tips of oar blades.

The parietal lamella is undulate and is tallest where it

passes the paired lower palatal and subcolumellar nod-

ules. The distal edge of the parietal lamella points to-

wards the lower palatal nodules.

The penis is relatively short and moderately robust; it

is apically papillate and basally smooth (Figure 23). The

epiphallus is moderately short, about the same length

and diameter as the penis. The penial retractor muscle

is inserted nearly apically on the penis. The epiphallus

is constricted slightly at the subapical junction with the

penis and has strong internal longitudinal folds. The base

of the spermathecal duct is robust with weak internal

folds, the remainder of the duct is slender; the sper-

matheca is ovate. Interior of the vagina and free oviduct

bear many folds. The free oviduct is roughly % larger

than the base of the penis, expanding greatly at the junc-

tion with the spermathecal duct and narrowing again be-

fore the junction with the uterus. Description is based

on two dissections.

Page 62 THE NAUTILUS, Vol. 118, No. 2

23. Camera lucida drawings of genitalia, UF 294575, maximum width 4.0 mm. 24. UF 294575, horizontal field width 167 jm.

J. Slapcinsky and B. Coles, 2004

Page 63

The central tooth of the radula is tricuspid; the me-

socone is elongate (Figure 24). The ectocones are short,

symmetric and diamond-shaped. The three lateral teeth

are tall, slender and asymmetrically tricuspid. The en-

docones of the laterals are tall and flare away from the

mesocone, while the ectocones are short and not as

strongly differentiated. The peripheral margin of the

mesocone of the laterals is concave above the ectocone.

There are twelve unicuspid marginal teeth on each side,

all with concave peripheral and convex proximal, mar-

gins. Based on the examination of two radulae.

Holotype: UF 304986, J. Slapcinsky and H. G. Lee, 9

Jun. 2001.

Paratypes: UF 286492, type locality, B. A. Brown,

May 1989; UF 294574, UF 294575, type locality, J. Slap-

cinsky and H. G. Lee, 9 Jun. 2001; UF 294573, UF

294574, Betsy's Gap, State Road 209, 0.5 km SW of sum-

mit, 35°41.3’ N, 82°54.3’ W, 1150 meters elevation: UF

294571, UF 294571, J. Slapcinsky, 11 Mar. 2001, 29 May

2001; UF 292056, Harmon Den Road 1.4 km SW of

Max Patch Road, 35°46.5’ N, 82°57.8’ W, 1000 meters

elevation, J. Slapcinsky and H. G. Lee, 9 Jun. 2001; UF

292713, Madison County: State Road 63 at Friezeland

Creek, 35°43.6’ N, 82°50.5’ W, 1000 meters elevation,

J. Slapcinsky and H. G. Lee, 9 Jun. 2001; UF 294577,

0.3 km NE of junction of State Road 1175 and State

Road 1182, 35°44.7’ N, 82°57.0’ W, 1170 meters eleva-

tion, J. Slapcinsky and H. G. Lee, 9 Jun. 2002; BC 1930,

Tennessee, Greene County: Cherokee National Forest,

Paint Creek Use Area, Hurricane Gap Road 3.2 km from

Paint Creek, 35°57.9’ N, 82°50.5’ W, 640 meters eleva-

tion, B. Coles; UF 293022, Paint Creek near Forest

Road 31, 1.3 km SW of fee station, 35°58.3’ N, 82°51.0'

W, 500 meters elevation, J. Slapcinsky, 1 Jun. 01; UF

293059, Paint Creek Road 5.3 km SW of fee station near

junction with Forest Road 31, 35°57.3’ N, 82°53.4’ W,

420 meters elevation, J. Slapcinsky, 1 Jun. 01; BC 6763,

Forest Road 3 by Paint Creek, B. Coles, 21 May 2002,

35°58.6’ N, 82°50.7’ W.

Type locality: USA, North Carolina, Haywood Coun-

ty, Carter Mountain Road at small stream 0.3 km SE of

State Road 209, 35°40.7’ N, 82°54.4’ W, 1030 meters

elevation.

Habitat and Distribution: Pilsbryna nodopalma is

known from sites that extend for 50 km along the moun-

tains on the North Carolina-Tennessee border, north-

west of Asheville. Specimens have been found at 400-

1100 m elevation usually on wooded, rocky hillsides in

leaf-litter. Although, like other species of Pilsbryna, this

species is found in moist leaf-litter, it is also found

among leaves on relatively dry rock outcrops.

Etymology: Named for the dorsoventrally com-

pressed apertural nodes that resemble the ends of oars

(Latin, noda = knot and palma = palm or oar blade).

For the purposes of the American Fisheries Society list

of the common names of mollusks (Turgeon et al., 1988,

1998) and other administrative uses, the common name

“oar tooth bud” is proposed.

Remarks: Juveniles of Pilsbryna nodopalma are most

likely to be confused with P. vanattai, the only other

species with a series of palatal nodules. Pilsbryna no-

dopalma has nodules at the lower palatal position and

the distal edge of the parietal lamella points toward

these. In contrast, in PR. vanattai the nodules are at mid-

palatal position and the distal edge of the parietal lamella

points below these nodules. The nodules of P. nodopal-

ma are dorsoventrally compressed, unlike the simple

peg-shaped to elongate nodules of P. vanattai. Adults of

P. nodopalma differ from all other species of Pilsbryna

in having the whorls widest above mid-point rather than

below. The penis, epiphallus and free oviduct are more

robust than in other species of Pilsbryna except for P.

vanattai. The endocones of the lateral teeth of P. no-

dopalma flare away from the mesocone, more so than

any other species of Pilsbryna. A juvenile Pilsbryna

specimen figured by Lee (1990) is this species.

Pilsbryna quadrilamellata new species

Common name: four blade bud

(Figures 25-31, Table 1)

Diagnosis: A small to medium sized Pilsbryna with a

shell of 2.8—3.2 mm diameter, 1.5-1.8 mm height, with

4.75.2 slowly expanding whorls. Shells of immature an-

imals contain four lamellae, a crescent-shaped umbilical

lamella and blade-shaped parietal, basal and sutural la-

mellae located within 4 whorl of the aperture. Lamellae

are reduced in adult specimens; however traces of la-

mellae, especially the basal lamella are visible in the

shells of many adults.

Description: Shell depressed-helicoid, umbilicate,

glossy, and translucent with a sculpture of dense and

irregularly spaced indented axial lines (Figures 25-27).

Adult shells (Table 1) are about 2.8-3.4 mm (mean =

3.2, n = 10) in major diameter and 1.5-1.8 mm (mean

= 1.7) in height with 4.7-5.2 (mean = 5.0) whorls. Shell

height is 0.47-0.61 (mean = 0.52) of greater diameter.

The whorls expand slowly and regularly; the lesser/great-

er diameter ratio of adult shells is 0.81—0.91 (mean =

0.87) the spire-width/greater diameter ratio 0.55-0.67

(mean = 0.61). The funnel-shaped umbilicus expands

regularly; the umbilical width is 0.4-0.6 mm (mean =

0.5) and the ratio of umbilical width to greater diameter

is 0.13-0.19 (mean = 0.17). The aperture of juvenile

shells is evenly crescentic. In adults the sutural edge of

the lip is flattened and the body whorl is widest basally.

Shells of juveniles contain four lamellae at columellar,

basal, sutural and parietal positions (Figures 28-29). The

columellar lamella is short and crescent-shaped and is

easily seen through the translucent base of the shell,

although it often does not reach near enough to the ap-

erture to be seen in apertural view. The basal lamella is

thick, the distal edge broadly rounded. The sutural la-

mella is narrow, the distal edge evenly rounded and is

Page 64 THE NAUTILUS, Vol. 118, No. 2

Figures 25-31. Pilsbryna quadrilamellata. 25-27. UF 292445, diameter 3.4 mm. 28. UF 292445, diameter 2.5. 29. UF 292445,

diameter 2.1 mm. 30. Camera lucida drawings of genitalia, UF 292442, maximum width 5.8 mm. 31. UF 292442, horizontal field

width 150 wm.

J. Slapcinsky and B. Coles, 2004

most easily seen in an apical view through the translu-

cent shell. The parietal lamella is thin and blade-shaped,

tapering distally to a sharp edge. All four lamellae are

reduced as individuals reach maturity, however some

trace of lamellae, especially the basal lamella, remains in

many adults.

The penis is relatively long and slender, apically pa-

pillate and basally smooth (Figure 30). The epiphallus is

moderately long and slender, roughly the same diameter

and length as the penis. The penial retractor muscle is

inserted nearly apically on the penis. The epiphallus

joins the penis subapically. The interior of the epiphallus

bears strong internal folds. The spermathecal duct is

long and slender and expands slightly at the junction

with the free oviduct. The interior of the vagina, free

oviduct and base of the spermathecal duct have weak

folds. The free oviduct is about 4% larger than the base

of the penis, and does not expand significantly at the

junction with the base of the spermathecal duct. Two

animals were dissected.

The central tooth of the radula is tricuspid; the me-

socone is very slender and elongate (Figure 31). The

ectocones are relatively short, a little more than % the

total tooth height, symmetric and not significantly con-

stricted basally. The three lateral teeth are tall, slender

and asymmetrically tricuspid. The endocones of the lat-

erals are tall, while the ectocones short, less than % the

height of the entire tooth. Both ectocones and endoco-

nes flare strongly away from the mesocone. Each side of

the radula has eighteen, tall, slender and unicuspid mar-

ginal teeth, with concave peripheral and convex proximal

margins. Description is based on examination of two rad-

ulae.

Holotype: UF 304987, J. Slapcinsky and B. Coles, 31

May 2001.

Paratypes: All from type locality: UF 292442, UF

292445, J. Slapcinsky and B. Coles, 31 May 2001; BC

6760, B. Coles, 20 May 2002; UF 288172, UF 297415,

J. Slapcinsky, 10 Jun. 2002; UF 299538, J. Slapcinsky, 10

Sep. 2002.

Type Locality: USA, Tennessee, Unicoi County, Una-

ka Springs, cold air slope along Unaka Springs Road, 3.2

km S of Banner Hill, 36°05.9’ N, 82°26.7' W, 520 meters

elevation.

Habitat and Distribution: All Pilsbryna quadrila-

mellata specimens were collected from leaf-litter within

approximately 20 m of the base of a talus slope on a NE

facing slope of the Nolichucky River. The riverbank sup-

ports hemlock forest with cove hardwoods on thin, rich

soil overlaying sandstone talus, with pockets of deep leaf-

litter. A steady stream of cold air emanates from the base

of the talus slope throughout the spring, summer, and

fall. Local residents visit the cold air slope during the

summer months and part of the base of the slope has

been cleared to provide a seating area.

Etymology: Named for the unique arrangement of

Page 65

four blade-like apertural barriers (Latin, quattuor = four

and lamella = blade). For the purposes of the American

Fisheries Society list of the common names of mollusks

(Turgeon et al., 1988, 1998) and other administrative

uses, the common name “four blade bud” is proposed.

Remarks: The juvenile shell of P. quadrilamellata is

unique in having a sutural lamella and a basal lamella

that is long, thick and evenly rounded distally. It is most

similar to P. aurea, both species having a crescent-

shaped columellar lamella. The body whorl of P. quad-

rilamellata does not expand rapidly at maturity like most

other Pilsbryna species. Adult P. quadrilamellata are un-

usual in having the sutural margin of the final third of

the body whorl flattened. The penis and epiphallus of P.

quadrilamellata are unusually delicate and elongate. The

penis and epiphallus are uniform in width throughout,

similar only to P. nodopalma and P. vanattai. The en-

docones of the lateral teeth of the radula flare strongly,

similar only to those of P. nodopalma and the ectocones

are relatively shorter than in any other species of Pils-

bryna.

Pilsbryna vanattai (Walker and Pilsbry, 1902) new

combination

Common name: honey glyph

(Figures 32-38, Table 1)

Vitrea vanattai Walker and Pilsbry, 1902: 432, pl. 23, figs. 4-6

Retinella (Glyphyalus) vanattai (Walker and Pilsbry, 1902).

Baker, 1930: 205, pl. 10, figs. 9, 10; Pilsbry, 1946: 273-

274, Fig 135; Burch 1962: 98, fig. 229.

Glyphyalinia vanattai (Walker and Pilsbry, 1902). Baker, 1962:

20; Hubricht, 1970: 13; Hubricht, 1985: 23, Map 212; Tur-

geon et al., 1988: 134; Turgeon et al., 1998: 147.

Diagnosis: A medium to large-sized, relatively de-

pressed and loosely coiled Pilsbryna with a shell of 3.8—

4.4 mm diameter and 1.9-2.2 mm height with 4.6-5.1

whorls. The body whorl of adults expands rapidly; the

ratio of greater/lesser diameter of adult shells is 0.80-

0.89. Juveniles have a sinuous parietal lamella and two

spirally arranged series of one to three widely spaced

paired basal and palatal nodules. The palatal lamella oc-

cupies a position lateral to and slightly above the parietal

nodules.

Description: Shell depressed-helicoid, umbilicate,

and fragile, sculptured with irregular, impressed, axial

lines (Figures 32-34). The sutures are shallow. Adult

shells are relatively large (Table 1), 3.84.4 mm (mean

= 4.0, n = 10) diameter, and 1.9-2.2 mm (mean = 2.0)

height with 4.6-5.1 (mean = 4.8) whorls. The whorls

expand slowly and regularly up to four whorls and then

very rapidly, the ratio of lesser/greater diameter of adult

shells is 0.80-0.89 (mean = 0.84). The rapid expansion

of the body whorl results in low ratios for shell height/

greater diameter of 0.48—-0.56 (mean = 0.51) spire

width/greater diameter 0.48-0.56 (mean 0.52) and um-

bilical width/greater diameter of 0.12-0.16 (mean =

0.14). The umbilicus is narrow until the final third of the

Page 66 THE NAUTILUS, Vol. 118, No. 2

Figures 32-38. Pilsbryna vanattai. 32-34. UF 279916, diameter 4.0 mm. 35-36. UF 279916, diameter 2.0 mm. 37. Camera

lucida drawings of genitalia, UF 279917, maximum width 5.7 mm. 38. UF 279917, horizontal field width 160 ym.

J. Slapcinsky and B. Coles, 2004

Page 67

body whorl and then expands rapidly to 0.5-0.6 mm

(mean = 0.5). Immature shells with fewer than 3.5

whorls have three lamellae: a sinuous parietal lamella

and a series of paired basal and palatal nodules. When

viewed from the aperture, the palatal lamella is situated

lateral to and slightly above the parietal lamella. Two to

three pairs of palatal and basal nodules can usually be

seen through the base of the juvenile shell (Figures 35—

36). These barriers are widely spaced, up to % whorl

apart, and are usually completely resorbed in shells over

4 whorls. The undulating parietal lamella extends up to

%; of a whorl and is more prominent where it passes the

paired basal and palatal lamellae.

The penis is papillate on its apical third and smooth

basally (Figure 37). The penial retractor muscle inserts

apically on the penis. The epiphallus joins the penis lat-

erally at the apex, is short, only % the length of the penis,

and robust, slightly narrower than the penis. The epi-

phallus is folded near mid-point, its interior has several

strong, longitudinal folds and it is slightly swollen at the

junction with the vas deferens. The interior of the va-

gina, free oviduct, and base of the spermathecal duct,

bear many longitudinal folds. Three animals were dis-

sected.

The central tooth of the radula is tall, slender and

tricuspid (Figure 38). The mesocone is elongate, widest

near mid-point and tapers apically and narrows basally.

The ectocones are short, symmetrical and diamond-

shaped. The three lateral teeth are tall, slender and

asymmetrically tricuspid. The endocones of the laterals

are tall and the ectocones short. The ectocones and en-

docones are well defined, but do not flare strongly away

from the mesocone. The peripheral edges of the first

two lateral teeth are concave. The third lateral tooth is

notched above the ectocone. The twenty-two marginal

teeth on each side of the radula are tall, slender and

unicuspid, with concave peripheral edges and convex

proximal edges. Two radulae were examined.

Lectotype: ANSP 8326la.

Paralectotype: ANSP 410030, from type locality, J.H.

Ferriss, 1901.

Type Locality: USA, North Carolina, Yancey County,

Mount Mitchell, J. H. Ferriss, 1901.

Other Material Examined: USA, North Carolina:

Avery County, Cranberry, Baker (ANSP 158889). Bun-

combe County: Pisgah National Forest: State Road 197

ca. 9 km E of Barnardsville, 35°47.9’ N, 82°22.1’ W, 985

m elevation, J. Slapcinsky, 29 Apr 2000 (UF 279987);

Walker Cove, Forest Road 74, ca. 7 km E of Dillingham,

35°45.7' N, 82°21.6’ W, 1140 m elevation, J. Slapcinsky

and R. Caldwell, 24 Apr 1998 (UF 279967), J. Slapcinsky

and S. Florence, 27 Apr 2000 (UF 279916, UF 279917),

J. Slapcinsky, 13 Mar 2001 (UF 287012), J. Slapcinsky,

1 Jun 2001 (UF 287010, UF 287011); Forest Road 74

at Perkins Road Trail, 35°44.9’ N, 82°21.4’ W, 1200 m

elevation, J. Slapcinsky and S. Florence, 27 Apr 2000

(UF 279983, UF 279984): Forest Road 74, 0.5 km N of

Laurel Gap Trail, 35°44.5’ N, 82°21.9’ W, 1200 m ele-

vation, J. Slapcinsky and S. Florence, 27 Apr 2000 (UF

279986, NCSM P-4731); Douglas Falls Trail, 0.5 km S

of Forest Road 74, 35°43.2’ N, 82°22.4’ W, 1350 m el-

evation, J. Slapcinsky and S. Florence, 27 Apr 2000 (UF

279988); Forest Road 63 ca. 8 km S of Dillingham,

35°42.6’ N, 82°23.7' W, 1190 m elevation, J. Slapcinsky,

2 Jun 2001 (UF 287013); Bent Creek Experimental For-

est, Forest Road 479, ca 1 km N of Blue Ridge Parkway,

35°27.7' N, 82°39.7' W, 860 m elevation, J. Slapcinsky,

2 Jun 2001 (UF 288621, UF 288622). Mitchell County:

1.9 km E of Spruce Pine, L. Hubricht, 4 Jun 1964

(FMNH 240499); Magnetic City, A. G. Wetherby, 1893

(ANSP 64609). Yancey County: near South Toe River,

7.2 km E of Mount Mitchell, 975 m elevation, L. Hu-

bricht, 26 May 1962 (FMNH 240500); Black Mountains,

Cat Tail Cove, J. H. Ferriss, 1901 (ANSP 84066). Ten-

nessee: Carter County: Iron Mountain, Forest Road

4331 at Fall Branch, 36°9.1’ N, 82°10.7’ W, 1180 m el-

evation, J. Slapcinsky and B. Coles, 30 May 2001 (UF

992523): Roan Mountain, State Road 143 at Dave Miller

Hollow Road, 36°10.3' N, 82°6.1' W, 840 m elevation,

J. Slapeinsky and B. Coles, 30 May 2001 (UF 292505);

Roan Mountain, behind picnic area W of Dave Miller

Hollow Road, 36°10.3’ N, 82°5.9’ W, 850 m elevation,

J. Slapcinsky and B. Coles, 30 May 2001 (UF 292470);

4.0 km S of Roan Mountain, L. Hubricht, 21 Sep 1967

(FMNH 240502): N outliers of Roan Mountain, Baker

(ANSP 158887). Unicoi County, State Road 107, 0.5 km

E of Red Fork Road, 36°9.2’ N, 82°14.9' W, 880 m el-

evation, J. Slapcinsky and B. Coles, 31 May 2001 (UF

293083).

Habitat and Distribution: Pilsbryna vanattai has

been found at sites between 800 and 1400 m elevation,

ranging 120 km along the Blue Ridge of North Carolina

and W into extreme eastern Tennessee (Pilsbry, 1946;

Hubricht 1970, 1985). Specimens located in this survey

were found at the soil leaf-litter interface on rich wet

soils within a few meters of seeps, springs and small

streams, often among stinging nettles, Laportea cana-

densis (Linnaeus, 1753), but also, less commonly, in

deep leaf-litter at the base of rock outcrops.

Remarks: The juvenile shell of Pilsbryna vanattai dif-

fers from that of P. nodopalma, the only other species

with palatal nodules, in having nodules that are not dor-

soventrally compressed and that are located at mid-pal-

atal rather than sub-palatal position. The body whorl of

P. vanattai expands more rapidly than any other species

of Pilsbryna. P. vannattai has the lowest ratios of lesser

diameter/greater diameter, height/greater diameter,

spire width/greater diameter, and umbilical width/great-

er diameter of any Pilsbryna species. The penises of P.

vanattai and P. nodopalma are more robust than those

of other species of Pilsbryna. The epiphallus of P. van-

attai is relatively shorter than any other species, only

about % the length of the penis. The ectocones and en-

docones of the lateral teeth of PR vanattai are well dif-

ferentiated, unlike P awrea and P. castanea, but do not

Page 65

THE NAUTILUS, Vol. 118, No.

Figures 39-43. Helicodiscus tridens. 39-41. UF 286491, diameter 1.6 mm. 42. UF 286491, diameter 1.4 mm. 43. UF 286491,

horizontal field width 42 m.

flare away from the mesocone, unlike P nodopalma and

P. quadrilamellata.

Walker and Pilsbry (1902) did not designate a holo-

type for P. vanattai. A single set of approximate mea-

surements was given, but these could refer to either of

the two adults of the three specimens mentioned in the

description. Baker (1962) selected the specimen figured

by Walker and Pilsbry (1902, figs. 4-6) to be the ‘type’,

thereby designating the lectotype, and segregated it as

8326 1a. The rernentndler of the lot, the two “paralecto-

types, remained ANSP 83261. Later, the juvenile para-

lectotype was lost and the remaining paralectotype was

recataloged (ANSP 410030).

Family Helicodiscidae

Genus Helicodiscus Morse, 1864

Helicodiscus tridens (Morrison, 1935)

Common name: crosstimbers coil

(Figures 39-43)

Pilsbryna tridens Morrison, 1935: 546, figs. 8-10; Pilsbry, 1946:

393, fig. 209; Burch, 1962: 106 fig. 25; Cheatum and Ful-

lington, 1971: 9, fig. 12.

Halkicadhisene ti idens (Morrison, 1935). Hubricht, 1964: 28: Rie-

del, 1980: 52; Hubricht 1985: 20, fig. 182; Turgeon et al.,

1988: 131; Turgeon et al., 1998: 148.

Holotype: USNM 359722, P. V. Roundy.

Type Locality: USA, Texas, Palo Pinto County, near

Strawn.

Other Material Examined: USA, Oklahoma: Mus-

kogee County, South Canadian River at Highway 2

35°15.8’ N, 95°14.4’ W, 150 m elevation, B. Galles. 28

Nov 1998 (UF 286491); Haskell County, South Canadian

River, Whitefield, L. Hubricht, 1935 (FMNH 239114):

Texas, Colorado County, Colorado River at Columbus,

H. F. Wickham, 8 Jan 1953 (ANSP 189563).

Habitat and Distribution: All previously known

specimens of Helicodiscus tridens were collected from

river drift or Pleistocene deposits from central

Oklahoma to central Texas (Hubricht, 1985). Recent

specimens of H. tridens including one with a dried an-

imal were sifted from leaf-litter under willows, Salix sp.

Remarks: Comparisons of the species described in the

genus Pilsbryna have highlighted the need to resolve the

generic placement of Teahaadiaons tridens (Morrison,

1935). Helicodiscus tridens was described as a Pilsbryna

species based on similarities in the apertural barriers.

Hubricht (1964) moved the species to the genus Heli-

codiscus. However, the basis for this change was not giv-

en, and the generic placement of the species has re-

J. Slapcinsky and B. Coles, 2004

Page 69

Figure 44. Distribution of Pilsbryna: a = Pilsbryna aurea, c = P. castanea, n = P. nodopalma, q =

vanattai. Map scale: 1 em = 18 kn.

mained unresolved. Characters other than those of the

shell previously have not been available for study. Newly

collected material, including a single dried animal, allows

comparisons of both shell and radular morphology

among H. tridens and species of Pilsbryna and Helicod-

iscus. Characters of the shell (Figures 39-42), including

the nearly circular aperture, the small number of whorls

(about 4 in adults), the flat apex and base, the impressed

sutures, and the regularly expanding umbilicus, are not

similar to species of Pilsbryna. However, they are con-

sistent with Helicodiscus, sensu lato. The radula of H.

tridens differs from Pilsbryna species. The row of tiny

central teeth (Figure 43, far right), the much larger sym-

metric tricuspid lateral teeth and the short, broad, mul-

ticuspid marginal teeth are characteristic (Solem, 1975)

P. quadrilamellata, v = P.

of the helicodiscidae but are not found in Pilsbryna,

thereby supporting Hubricht’s (1964) placement of H.

tridens in Helicodiscus.

DISCUSSION

When species of Paravitrea and Helicodiscus are sepa-

rated from Pilsbryna, the geographic and microhabitat

distribution of Pilsbryna is clarified. All known popula-

tions of Pilsbryna occur near springs, seeps and moun-

tain streams in the southern Appalachian Mountains and

nearby Cumberland Plateau (Figure 44). Within these

areas, Pilsbryna species occupy moist microhabitats in

damp, often deep, leaf-litter. It is likely that Pilsbryna

species would be intolerant of habitat changes affecting

Page 70

THE NAUTILUS, Vol. 118, No. 2

soil hydrology and leaf-litter cover. The narrow distri-

bution and habitat specificity of species of Pilsbryna

should make them species of special concern to land

managers.

The terrestrial molluscan fauna of the southern Ap-

palachian Mountains is by far the most diverse in eastern

North America (Hubricht, 1985). Diversity notwith-

standing, the region is still not well sampled, especially

for small species with narrow habitat requirements. Re-

cent collecting efforts in wet leaf-litter microhabitats un-

covered not only the species reported here but also sev-

eral others that require additional material to adequately

describe. These results suggest there is a largely unre-

ported radiation of Pilsbryna at springs and seeps

throughout the southern Appalachian Mountains and

Cumberland Plateau. Additional collecting in this region

is necessary to determine the scope of this radiation.

ACKNOWLEDGMENTS

Staff of the Cherokee National Forest and Pisgah-Nan-

tahala National Forest particularly Joe McGuiness and

Sandy Florence generously shared their knowledge of

the lands in their care. Peter Wyatt of the Tennessee

Wildlife Resources Agency showed us the unusual cold-

air talus slope at Unaka Springs. Harry G. Lee (Jackson-

ville, Florida) drew our attention to North Carolina pop-

ulations of Pilsbryna nodopalma, donated specimens,

and helped collect additional specimens. Ron Caldwell

and Richard and Wanda Ott graciously hosted visits. We

are particularly grateful to private land owners, Billie

and John Brown, Walter McClain and the Davis family

who allowed access to their lands. Jochen Gerber

(FMNH) and Gary Rosenberg (ANSP) lent specimens

and/or facilitated visits to collections in their care. Fred

Thompson and Gustav Paulay (UF) commented on ear-

lier drafts of this manuscript. Fieldwork was conducted

with financial support from the Thomas L. McGinty En-

dowment Fund, University of Florida Foundation.

LITERATURE CITED

Baker, H. B. 1929a. New southem Appalachian land snails. The

Nautilus 42: 86-93.

Baker, H. B. 1929b. Pseudohyaline American land snails. Pro-

ceedings of the Academy of Natural Sciences of Philadel-

phia 81: 251-266.

Baker, H. B. 1930. The North American retinellae. Proceed-

ings of the Academy of Natural Sciences of Philadelphia

82: 193-219.

Baker, H. B. 1931. Nearctic vitreine land snails. Proceedings

of the Academy of Natural Sciences of Philadelphia 83:

85-117.

Baker, H. B. 1962. Type land snails in the Academy of Natural

Sciences of Philadelphia I. North America, North of Mex-

ico. Proceedings of the Academy of Natural Sciences of

Philadelphia 114: 1-21.

Burch, J. B. 1962. How to know the eastern land snails. Wm.

C. Brown, Dubuque. 214 pp.

Cheatum, E. P. and R. W. Fullington. 1971. Keys to the fam-

ilies of the recent land and fresh-water snails of Texas.

Dallas Museum of Natural History, Bulletin 1(supple-

ment): 1-18.

Emberton, K. C. 1985. Seasonal changes in the reproductive

gross anatomy of the land snail Triodopsis tridentata tri-

dentata (Pulmonata: Polygyridae). Malacologia 26: 225—

239.

Hubricht, L. 1964. Helicodiscus tridens and Helicodiscus ald-

richiana. The Nautilus 78: 28.

Hubricht, L. 1970. The land snails of North Carolina. Sterkiana

39: 11-15.

Hubricht, L. 1973. The land snails of Tennessee. Sterkiana 49:

UL=N7/,

Hubricht, L. 1985. The distribution of the native land mollusks

of the eastern United States. Fieldiana, Zoology, New Se-

ries 24: ]-19].

Lee, H. G. 1990. Toward an improved strategy for landsnail

collecting. The Shell-O-Gram 31(1): 3, 6-8.

Morrison, J. P. E. 1935. Three new land shells from the south-

em United States. Journal of the Washington Academy of

Sciences 25(12): 545-547.

Pilsbry, H. A. 1946. Land Mollusca of North America (north

of Mexico), The Academy of Natural Sciences of Phila-

delphia, Monograph 3, 2(1): 1-520.

Pilsbry, H. A. 1948. Land Mollusca of North America (north

of Mexico), The Academy of Natural Sciences of Phila-

ere Monograph 3, 2(2). 521-1113.

Riedel, 1980. Genera zonitidarum diagnosen supraspezi-

es taxa der familie Zonitidae (Gastropoda, Stylom-

matophora). Dr. W. Backhuys, Rotterdam. 197 pp

Solem, A. 1975. Polygyriscus virginianus (Burch, 1947) a hel-

icodiscid land snail (Pulmonata: Helicodiscidae). The

Nautilus 89(3): 80-86.

Tompa, A. S. 1984. Land snails (Stylommatophora). In A. S.

Tompa, N. H. Verdonk and J. A. M van den Biggelaar

(eds.), The Mollusca, Volume 7, Reproduction. Academic

Press: New York, pp. 47-140.

Turgeon, D. D., A. E. Bogan, E. V. Coan, W. K. Emerson, W.

G. Lyons, W. L. Pratt, C. F. E. Roper, A. Scheltema, F.

G. Thompson and J. D. Williams. 1988. Common and

scientific names of aquatic invertebrates from the United

States and Canada: mollusks. American Fisheries Society

Special Publication 16, 277 pp.

Turgeon, D. D., J. F. Quinn, Jr., A. E, Bogan, E. V. Coan, F.

G. Hochburg, W. G. Lyons, P. M. Mikkelsen, R. J. Neves,

C. F. E. Roper, G. Rosenberg, B. Roth, A. Scheltema, F.

G. Thompson, M. VWeaditione and J. D. Williams. 1998.

Common and scientific names of aquatic invertebrates

from the United States and Canada: mollusks, second edi-

tion. American Fisheries Society Special Publication 26,

526 pp.

Walker, B. and H. A. Pilsbry: 1902. The Mollusca of the Mt.

Mitchell region, North Carolina. The Proceedings of the

Academy of Natural Sciences of Philadelphia 54: 413-442.

THE NAUTILUS 118(2):71-87, 2004

Page 71

The South American Mollusca of Johann Baptist Ritter von Spix

and their publication by Johann Andreas Wagner

Robert H. Cowie

Center for Conservation Research and

Training

University of Hawaii

3050 Maile Way, Gilmore 408

Honolulu, HI 96822 USA

cowie @hawaii.edu

y Farmacia

Juan 670

ARGENTINA

[email protected]

Néstor J. Cazzaniga

Departamento de Biologia, Bioquimica

Universidad Nacional del Sur. San

(8000) Bahia Blanca

Matthias Glaubrecht

Museum fiir Naturkunde

Humboldt-Universitit zu Berlin

Institut fiir Systematische Zoologie

D-10115 Berlin

GERMANY

[email protected]

ABSTRACT

Dr. Johann Baptist Ritter von Spix collected Mollusca in South

America from 1817 to 1820. After his return to Europe he

completed the plates, including their legends, and brief diag-

noses for a monograph on the taxa he had collected, but died

in 1826 before the main text was written. Dr. Johann Andreas

Wagner was enlisted to complete the monograph, which he

did, and it was published in 1827. In total, 64 gastropod and

20 bivalve taxa were illustrated. In developing the monograph

for publication, Wagner altered Spix’s concepts of many of the

taxa, in some Cases using his own name, not Spix’s, as the au-

thor of the name. We discuss Wagner's rationale for making

these changes. We discuss the appropriate citation of the au-

thorship of the work, concluding that it should be cited as of

Wagner alone. We also discuss the appropriate authorship of

the species, concluding that in all cases in which Spix had pro-

vided a name on the plates authorship should be given as “Spix

in Wagner”. Wagner also created some new names, three of

which are replacements for Spix’s names, which are preoccu-

pied, but the remaining ones are unnecessary replacement

names for Spix’s names (of which they are therefore junior

objective synonyms) . Spix’s type material is in the Zoologische

Staatssammlung in Munich, although some of it is missing as

a result of damage sustained during World War II.

INTRODUCTION

Johann Baptist Ritter von Spix was born in Héchstadt

an der Aisch, near Bamberg, Bavaria, on February 9,

1781, as the son of a surgeon. [For additional biographic

information see Fittkau (1983) and Huber and Huber

(1993)]. He gained a doctorate in theology at the Uni-

versity of Bamberg but then changed his career to med-

icine, gaining a medical degree in 1806 from the Uni-

versity of Wirzburg. He then practiced medicine for a

short period before traveling to Paris in 1808 to meet

Cuvier, Lamarck, and other well-known naturalists. On

October 31, 1810, the Bavarian king, Maximilian I Jo-

seph, made him adjunct (scientific assistant) to the cu-

rator of the zoological-zootomical collections of the Ba-

varian Academy of Sciences in Munich and, less than six

months later, on April 24, 1811, following orders from

the king, the Academy made him curator, and thus, di-

rector of these collections. During the period 1808-1811

Spix traveled not only in France but also in Italy and

Switzerland. In 1811, he published his major work, Ges-

chichte und Beurtheilung aller Systeme in der Zoologie

[= History and evaluation of all systems in zoology] (xiv

+ 710 p., Achrag’sche Buchhandlung, Niirnberg). Spix

greatly improved the old-fashioned arrangement of the

natural history cabinet of the Bavarian Academy of Sci-

ences and is regarded as the founder of the modern

Bavarian zoological collections, now the Zoologische

Staatssammlung Miinchen (ZSM).

In 1817, in the company of botanist Carl Friedrich

Philipp von Martius (1794-1862) and a number of other

naturalists, Spix embarked on a major expedition of ex-

ploration and natural history collecting to Brazil. They

returned to Europe on December 10, 1820, with an

enormous quantity of material that they had collected,

which was deposited in the Bavarian Academy of Sci-

ences where Spix was curator.

Based on that extensive material, Spix published nu-

merous works, on monkeys and bats, turtles and frogs,

lizards, and birds. Martius published on the plants. Sad-

ly, only six years after his return from South America,

Spix died, in Munich on May 15, 1826, apparently as a

result of lingering illness contracted during the expedi-

tion. As a consequence, Spix’s intended publications on

fishes, insects, and mollusks remained unfinished and

had to be completed by others.

Under the auspices of the editors (Franz von Paula

von Schrank and Martius), the mollusk volume (Testacea

fluviatilia ...) was completed by Dr. Johann Andreas

Wagner (March 21, 1797—-December 17, 1861) and pub-

lished in 1827. The primary aim of this paper is to clarify

the authorship of the species described therein, follow-

ing, when pertinent, the International Code of Zoologi-

cal Nomenclature (ICZN, 1999), hereafter the Code, as

Page 72

well as correct attribution of the work itself. In order to

do this, it is necessary to discuss in detail the history of

the production of the volume.

HISTORY OF THE TESTACEA FLUVIATILIA

The title page of the original issue of the mollusk work

printed in Munich is shown in Figure 1. The title trans-

lates as follows:

Freshwater mollusks that, while traveling through

Brazil during the years 1817-1820 commanded by and

under the auspices of Maximilian Joseph I, Most

August King of Bavaria, were collected and taken care

to be painted by Dr. J. B. von Spix, former Civilian

Knight of the Royal Order of the Bavarian Crown,

Ordinary Member of the Bavarian Academy of

Sciences, curator of the collections of the Royal

zoological, zootomical and ethnographical Museum.

Arranged, described and illustrated with observations

by Dr. J. A. Wagner

Edited by Dr. F. von Paula von Schrank and

Dr. C. F. P. von Martius

Munich

Publisher C. Wolf

1827

A later issue was printed in Leipzig, also in 1827. It

differs slightly in a number of ways, as discussed below.

In total, 64 gastropod and 20 bivalve species were il-

lustrated. The title (Testacea fluviatilia), however, does

not reflect the contents correctly because, although the

preface reads that the mollusks studied only live in fresh

waters (“quae nonnisi aquas dulces incolunt”), 43 out of

64 gastropod species are land snails (e.g., genera Buli-

mus, Strophocheilus, Helix, Achatina). This suggests that

perhaps at least the editors (who were not malacologists)

were not fully aware of the contents of the book. If Spix

himself had had the opportunity of publishing his work,

then he may have titled it “Testacea fluviatilia et terres-

tria” or “Testacea Brasiliensia’; the latter perhaps being

his intention, as suggested by the heading on page 1| of

the descriptive text, above the title of the Ampullaria

section. Wagner, of course, knew that many species were

terrestrial, which suggests that he was not involved in

deciding the title of the work nor in the writing of the

preface.

The preface (“Praefatio”) (Figure 2) to the work ex-

plains some of its history. It is difficult to translate it

because several terms and structures are not classical,

with many embedded sentences and quotations. For ex-

ample, the preface begins with “Reliquerat” |= had left

behind], which has its object “icones” [= illustrations]

21 words later. This object, “icones”, has two modifying

sentences connected by “ac” [= and]; the first runs from

“ad exemplaria” to “illustratas” (14 words), while the

second runs from “reliquis” to “servituras” (12 words).

Within this second sentence, “animalium collectorum’” is

a genitive construction with a seven-word adverbial

phrase in between. We have tried to make the following

translation as close to the original as possible; however

THE NAUTILUS, Vol. 118, No. 2

this has led to the use of somewhat awkward English in

places. Also, we have placed some nouns in brackets

since in Latin it was common to omit them and leave

the reader to recover the meaning from the adjective.

In addition, some explanatory material and some of the

original Latin wording is also placed in brackets for clar-

ity. The following is our translation of the “Praefatio”.

PREFACE.

Dr. Johann Baptist von Spix, formerly our colleague

in the Royal Academy of Sciences, whose death, pre-

mature and calamitous for letters [i.e., science], we

grieve, had left behind him illustrations of the animals

with shells [“Testaceorum’| and the fishes, from the ex-

amples deposited by him in the Brazilian collections of

the Academy [“in Museo academico Brasiliano” |, drawn

on stone [i.e., lithographs] and illustrated in colors, and

had intended to use the [illustrations] to serve [creation

of] the missing descriptions of the animals collected

while traveling through the wide provinces of Brazil.

These [illustrations] were handed to us by a brother of

the now deceased man to be shared with the supporters

of Spix’s works. And there were not-weak arguments that

impelled us to put hands to work; mainly the vividness

and fidelity of the illustrations themselves, and the value

of the depicted shells, most of which are now made

known for the first time to those curious about nature;

the rest [of the illustrations], even if [the species] have

been described by other authors, will nonetheless be

worth viewing because, as they were collected by Spix,

these authors will themselves have future testimonial [to

their work] by matching their [shells] with the illustra-

tions of the indefatigable traveler.

Doubtless, descriptions, observations and other

[notes], which could be useful to explain the illustrations,

were lacking; but nature offers itself to the eyes. More-

over, being constrained by other issues and dedicated to

other studies, we would have hardly had the necessary

free time to accomplish the work: it seemed to us that

this was a task for a man that measured up to him [i.e.,

Spix]. Conveniently, we happened to find out that Dr.

J. A. Wagner had been engaged for several years in a

large and important study of conchology, and was striv-

ing to publish a continuation of the works of Chemnitz,

under the name of the Museum Conchyliologicum; the

learned man did not refuse the commission that we re-

quested of him, and got to work in order to complete

[the present book].

What he achieved is evident from the work itself. He

worked very hard in order that these mollusks, which

only live in freshwater, should be included within the

genera established by Lamarck and other recent authors,

in the conviction that he had to do so in the interest of

science because those [genera] that were given by Spix

did not rely on a solid foundation, but rather seemed to

be based on weak characters. Similarly, for the species,

he reduced many to already known species, and carefully

added all their synonyms. Even though by doing this the

number of species that we consider new decreases, it is

still the case that a large number of new [species] sur-

vives, which cannot be unpleasing to those interested in

nature.

We hereby offer this posthumous work, trusting that

it will enhance the memory of this man of natural sci-

R. H. Cowie et al., 2004

Page 73

TESTACGCEA FLUVIATILIA

QUAE

IN ITINERE PER BRASILIAM

ANNIS, MDCOCKYII — MDCCCKX

JUSSU ET AUSPICIIS

MAXIMILIANI JOSEPHIIL

BAVARIAE REGIS AUGUSTISSIMI

SUSCEPTO

COLLEGIT ET PINGENDA CURAVIT

Dre od COB den SPTX

Quondam Ordinis Regu Coronee Bavarice Ciwilis Eques, Academie scientiarum Bavarice

Socius Ordinarius, Muset Regi zoologici, zootomici et ethnographict Conservator rel.

DIGESSIT, DESCRIPSIT ET OBSERVATIONIBUS ILLUSTRAVIT

Dr. J. A WAGNER

EDIDERUNT

Dr. F. ad PAULA de SCHRANE et Dr. C. F. P. de MARTIUS.

MONACHII,

NeoPa (S) (Chn WV Oak

1 is-2 @

BNL

Figure 1. The title page of the Munich edition of the “Testacea fluviatilia ...”.

Page 74

THE NAUTILUS, Vol. 118, No. 2

PRAEFATIO.

Raises Dr. Joannes Baptista de Spix, Collega quondam nosterin Aca-

demia Scientiarum Regia, cujuspraematuram ac litteris calamitosam dolemus

mortem, Testaceorum Pisciumque icones, ad exemplaria in Museo acade-

mico Brasiliano a se deposita in lapide delineatas coloribusque illustra-

tas, ae reliquis animalium in itinere per amplas Brasiliae provincias in-

stituto collectorum descriptionibus seryituras. Has beati nunc viri fra-

ter nobis tradidit, cum fautoribus operum Spixianorum communi-

candas. Et erant argumenta non levia, quae nos impellebant, operi ma-

num ut adhiberemus; nitor praecipue et fides iconum ipsarum, delineato-

rum conchyliorum pretium, quorum pleraque nunc primum naturae cu-

riosis innotescunt, alia yero, quanquam jam ab aliis autoribus descripta,

digna tamen videbantur, quae, quod, a Spixio collecta, illis ipsis autori-

bus testimonio futura essent, inter icones ab indefesso peregrinatore para-

tas collocarentur.

Deerant quidem descriptiones, obseryationes, aliaque, quae ad tabu-

larum explicationem usui esse possent; sed natura ipsa ob oculos versa-

tur. Ceterum, quum nos et aliis negotiis impediti et alienis studiis dedi-

ti vix otii necessarii satis ad perficiendum opus habituri nobis videremur,

de viro laborandum erat, qui illi par esset. Commodum accidit, uti re-

sciremus, D. J. 4. Wagner ab annis aliquot magno et egregio studio

Conchyliologiae operam dare, atque continuationem ejus operis quod

Chemnitius, Musei Conchyliologici nomine, edidit moliri; nec abnuit,

quam obtulimus, provinciam, vir eruditus operique perficiendo manus

admovit.

hy aes

Quae praestiterit, ex ipso constat opere. Plurimum laborayit, ut

haee conchylia, quae nonnisi aquas dulces incolunt, ad ea genera referret,

guae Lamarckius aliique recentiores constituerunt, ratus, id se scientiac

commodo debere, propterea quod quae a Spirio statuta sunt, genera non

satis firmo fundamento, nec nisi fluxis characteribus niti yidebantur. Idem in

speciebus obseryayit, quarum plures ad species notas reyocayit, synonyma-

que diligenter omnibus addidit. Qua re quanquam factum est, ut novya-

rum, quas dicimus, specierum numerus minucretur, magna tamen yere

noyarum copia superest, quae naturae curiosis non potest non esse per-

grata.

His nos opus hoc posthumum offerimus, spe freti, fore, uti et memo-

ria viri de scientia naturali optime meriti accessione augeatur noya, et

ipsi scientiae non contemnendum accedat incrementum.

Dabamus Monachii Tdib. Augusti MDCCCXXFTL

Dr. Franc. a Paula de Schrank.

Dr. Car. Frid. Phil. de Martius.

Figure 2. The “Praefatio” of the Munich edition of the “Testacea fluviatilia ...”.

ence of excellent merit through a new addition, and that

it will be a non-negligible contribution to science itself.

Written in Munich, on August 13, 1827.

Dr. Franz von Paula von Schrank.

Dr. Carl Friedrich Philipp von Martius.

Thus, it is clear that Spix produced lithographs and

that his brother passed them to the editors (Schrank and

Martius), who in turn gave them to Wagner in order that

he could produce the finished work, based upon Spix’s

illustrations. It is also clear that Spix provided names for

his species.

Wagner was a systematist of the Blumenbach and Cu-

vier traditions (Martius, 1862). He is known to have

avoided the creation of new genera and to have placed

new species as far as possible in genera already estab-

lished. His first published scientific work was in fact the

description of the mollusks collected by Spix in Brazil.

Gotthilf Heinrich von Schubert (1780-1860) and Wag-

ner published, in 1829, the 12" volume of Neues Sys-

tematisches Conchylien- Cabinet, the famous German

conchological series started by Friedrich Heinrich Wil-

helm Meera (volumes 1-3, 1769-1774) and Johann Hi-

eronymus Chemnitz (volumes 4-11, 1780-1788), as is

alluded to in the preface (above), where reference is

made to the “Museum Conchyliologicum” (which trans-

lates into English as “Conchological Museum” but was

probably intended as the Latin translation of the Ger-

man “Conchylien-Cabinet”). In 1831, Wagner published

a handbook of the natural history of the animal King-

dom, focusing in particular on mammals.

Wagner obtained his Ph.D. in 1826 from the Univer-

sity of Erlangen, having previously spent two years

(1814-1816) at the University of Wiirzburg. He then be-

came a Privatdozent—a privately paid lecturer—in Ee

langen, following a journey to Paris. On October 22,

1832, the Bavarian King awarded Wagner the position

of adjunct (scientific assistant) to Schubert, his friend

and Spix’s successor as curator of the Munich zoological

collection. It was apparently Schubert, following Spix’s

death in 1826, who arranged that Wagner be asked to

publish Spix’s material. The year 1832 is the earliest date

we know for sure that Wagner y was actually working in

Munich. He became a anenibeE of the Royal Bavarian

Academy of Science in 1835, and in 1849 was installed

as the third curator of the zoological-zootomical cabinet.

[For additional biographic information see Martius

(1862) ].

Wagner received, as the basic material for his study,

the 29 plates that Spix had created, with Spix’s names,

and probably some notes and Latin diagnoses (see be-

low). This corpus was wholly included in the book and

Wagner used Spix’s names, either confirming them or

R. H. Cowie et al., 2004

Page 75

TABULAE EXPLIGATAE.

Tab. 1. ‘Fig. 2. Ampallaria Gigs Spit. 5 0. we es Page

Tob. I. Fig. 1. Ampulllonin zonata Spix , aye eee ee se

Fieve) Ampillarinalsossiag Yar. (cf. Archimedes svi he

Tab. WM. Fig. 2- — Ampullaria olivacea Spix, ea

ig- 2. Ampulloriae olivaceae Var. (f. quercina Shi): 2.

Tab, TY. Fig. 1. 2. Ampullaria popyracea Spix 3.

Fig. 3. Ampullaria ? rosea Spix . 3.

Fir 4) “Acpulliviallinealalsntaor’ ((4/¢figulina Spt) 3.

Fig. 5. Ampnillaria cyclostoma Spix. . ‘ 4

TV. ~ Fig. 1. Ampullaria orasea\Swainss |(Ziclée\fasclotata Spt)» he

Fig. 2. Ampullaria lineata Wagn. (Helix lineata Spix) . 3

Fig. 3 Ampullarin crassa Swains. (Helix crassa Spix) . 4

Fig: A Ampulllaria erassa Swains. (Elelicina exumbilicata Spiz) . 4

‘Tab. VI. Fig. 1. —_‘Bulimus corrugatus Wi ‘sn. (B. terrestris Sra = vehi ae Bs

ig. 2. ‘Bulimus durus Spix.

Fig. 3. Bolimus hyelinus Wagn. “a. aa Soe A 3 6.

Fig. 4. _-Bulimus angiostomus Wagon. (B. virgalus Spix). . . 14.

Tab, YI. Fig. 1. Bulimus Magus Wogn. (B. aye Spx) UT ‘ 6.

Bulimus perlocidus) Spx « z.

Bulimus lituratus Spix kjjeous 7

Fig Bolimus vittatus Spix 7

Fig. Bolimus zebra Spix. Tmyar Bal. 8.

ig- Bulimus lincatus Spix Lig hes 8

Tab, YUL. Fig. Achatina melanostoma Swains. (4. perversa Spix) 16.

Tis penlaeestentaiconig lena era wltreu Spiz) , Zz

Bulimus decapitatus Spix, /ntlsfut, fe

Melania tuberculata Wagn. (dylacostoma Ripert Spi)

Melanin scalaris Wagn, (dylacostoma labrum) « ere

Balimus Gallina -Sultana ‘Lam. (dchatina pavonina Spix). . ,

Bulimus undatus Brug. (Achatina pulchella Spix) .

4 Bulimus floccosus Wagn. (Achalina floccosa Spix)

Dulimas caleareus Brug. (Colunna maritima Spix) .

Bulimus caleareus B. (Columna 8-gyrata Spix) - ‘

Bulimus Spixii Wago. (Columna bulimea Spix). . . - - . al.

15.

Bulimns sylyaticus Wagan. (Columna sylvatica Spix). . =.» ah

- 4.

ig. 3.

~ 4

igs 1

Age

+ Ae

aE5

Lymnacus papyraceus Spix Bea re Tc,

Tob, XI. Fig. ai oiitete ratte go (Sica HuclavtiarrcoriadlomuntSpesyi om ae eles

Fig. 2. 3. Bulimus virgincus Brug. (Strophocheilus dimeida Spx). » . «az

Tab. XIL Fig. 1. 2. Bulimus melanostomus Swains. (Auris melastoma Spiz) - + +» + 13,

Tig. Auricula signata. (Auris signata Spix). . + © + + = + Ih

Fig. Auriculne signatac Var. (Auris vittata Spix) . . +» - « + 1%

Tab. XU. Fig. 1. 2. Auricula Leporis Lam. (Stenostoma auritum Spix) . - - . = “U58>

Fig. Bulimus angulatus Wogn. (Stenostoma Puru Spix) - Bai na, Nits

Fig. Bulimus angiostomus Wagn. (Slenostoma Capucira Spiz) e eo Seay

Figure 3.

reducing them to synonymy. Wagner also included two

pages of “Tabulae explicatae” (Figure 3), essentially an

index that listed all the names in the sequence in which

they appeared on Spix’s plates, with reference to the text

page on which Wagner gave his own opinions regarding

what he had interpreted about each species.

AUTHORSHIP OF THE WORK

Prior to Spix’ss death, he had produced plates, with

names, illustrating the species. Following his death, the

editors enlisted Wagner to provide descriptions of the

species, which he did. The completed work was pub-

lished in 1827. In the past, some authors (e.g., Ihering,

1890; Haas, 1969) have considered Spix as sole author

of the work, perhaps because he provided the plates,

legends, and probably a short diagnosis for each species

(see below) that are the foundation of the work, which

Wagner simply prepared for publication, that is, as a sci-

entific editor of Spix’s work. Others, however, most likely

because Spix did not provide the text of the descriptions

but only the names and illustrations (and probably the

short diagnoses), have attributed authorship of the work

either to Wagner alone (because he was the author of

the descriptions) or to Spix and Wagner (as suggested

for example by Glaubrecht (1996: 488)), following the

convention that a name and illustration alone (Spix’s con-

tribution) did not satisfy the criteria of nomenclatural

Tob, XIV. Fig. 1. Pupa exesa Wogn. (Clausiliaexcsa Spix) . apy Pog. 19:

Fig. 2. Popa striata Wagn. (Clausilia striata Spix). . of a es 19-

7 gn. (Clausilia 6- dentata Shia) «We teny oka hacaas

Pupa sexdentai

Popa inflata (Clausilia pupoides Spix). . #1 BAe er eager ke

Tab, XY. Papa clatior Spix . «Pigs OSS Hid noun aaa

Helix Navicula Wagn, (Wovicuta Garcia oi) a0 Wht Pin: ce een

Helix clsusa Wagn. (Tomigerus clausus Spix) . : . . . . gh

Tab, XVI. Helix Pyramidella Wagn. (Helicina Pyramidetla Spix) - . . . Qh

Helicina yariab ‘gn. (Helicina fasciata Spiz) . . - * « oh

Helicinae yari Var. (Helicina flava Spix) . . Pes |,

Tab. XVU. Helix Pellis serpentis Chemnitz, (Solarium Serpens Spx) oe GSS tage

Helix punctata Wagn. (Solarium Serpens Spix), . . + = . ah

Helix perspectiva Ww ‘agn. (Solarium candidum Spix), . . > (he

Helix'perspectiva Wagn. Solarium vitreum Spiz) . . . . . oh

Holix yitrina Wago, (Solarium imperforatum Spix) - oe

Helix nana Wogn. (Solarium pygmacum Spix) . o7 (React reepe

Tab. XV. Fi Planorhis olivaceus Wagn. (Planorbis Merrie Spix) ate Sonam

Planorbis olivaceus Waga. ob Spi” = Visca ereeroe

Planorbis lugubris Wagn. (Planorbis witha Sey ce a7

Plavorbia lugubris VW, (Planorbis albefcavs pix) io Noa vente tne

Planorbis lugubris W. (Planorbis viridis Spix) . 9. . . . . ar

Tab. XIX. F 2. Anodon giganteus Spix . . RAR Cece ee TT Bt oe eh

‘Tab. XX. AnodopttrapexsunSpix 25-0) Sipe elie h «Ae SeR? > acy 498%

» Anodon rotundos Spix . 4 _ " a - . . . +) age

Tab. XXI Anodon anserinus Spix. © 5 ee ew

Tab. XX Anodon longus Spix, - 2 yee ee

Mnodim triponusiSpix 2c.) 7a ine pes Abeer Oriani Sees

Anodon obtusus Spix, Separates

(Sr(odontia/abtase spectaten juntue mgr (arcane taste Spi) 2 Abst = 508

Anodon radiatus Spic =. wy se - gts

Anodon siliquosus Spix soeeenego

» Auodontis iilquosi apdcimen juaius WVaen. (Andon pygmaeus Sri) + 50

Anodon ensiformis Spix. ‘ a esas

. Anoion ‘rotundas?! (Aptodon inermis Spiz). ashley

. Unio pectinatus Wagn. (Tetraplodon pectinatus Spi) eta Oe eee

Gyclas bahicnsis Wagn. . eS Aly nearer

(atUniolellipiteus Wazno(DiplodonelliprcilsySpis)) 5...) aan

Unio rolundus Wagn. (Diplodon rotundus Spiz). - . + + « 3h

Tab, XXI.

Tab. XXIV.

‘Tab. XXY.

Tab, XXYVI.

Tab. XVI. Fe . Unio raudatus Wagn. (Diplodon furcatus Spiz). . » ~ +» + 3

MSV Fige i.e. Unio rhombeus Wagn. (Diplodon rhombeus Spix) - . + = = 3h

TabXxex Fis 1. 2. Unio rugosus Wagn. (Triplodon rugosus Spix) . 9. + + + + 3

The “Tabulae explicatae” of the Munich edition of the “Testacea fluviatilia . ..”.

availability. However, the Code (Article 12.2.7) states

that a name associated with an illustration, published

before 1931, is sufficient to establish availability. Thus,

Spix’s contribution to the work is sufficient to validate

his authorship of the names associated with the illustra-

tions on the plates, even in the absence of Wagner's de-

scriptions, which also, however, standing alone, would be

sufficient to validate the names. Additional reasons for

attributing most of the names to Spix are discussed be-

low.

Given Spix’s contribution, that it is explicitly acknowl-

edged in the preface, that his name is clearly part of the

title page (in fact appearing first), and that there is no

evidence that the plates were published separately from

the text, authorship of the work should arguably be at-

tributed to both Spix and Wagner, and in that order be-

cause that is the order in which they appear on the title

page. Based on this argument, that is, that the work is

neither Wagner's nor Spix’s work alone, but their mixed

contributions, authorship of the work as of Spix and

Wagner could be justified, as was accepted by Fechter

(1983b).

A similar argument was used by Kottelat (1988) in

deciding that authorship of the work “Selecta genera et

species piscium quos in itinere per Brasiliam . . . collegit

et pingendos curavit Dr. J.B. de Spix” on the fishes col-

lected by Spix but published (in two fascicles: June 1829,

January 1831) by Louis Agassiz (1807-1873) should be

Page 76

attributed to Spix and Agassiz. This volume, the first

monograph on Brazilian fishes, also contains descriptions

of numerous new species and genera. While the plates

of this fish volume were prepared at least in part under

Spix’s supervision, the text was written by Agassiz. How-

ever, Kottelat (1988: 73) argued, based on article 50(a)

of the Code (3" edition, 1985) and the fact that Spix was

not author of both the names and the conditions making

them available, that for nomenclatural purposes Agassiz

is the sole author of the names. It appears that Agassiz

wrote the whole of the text, and whatever Spix wrote,

can only have been rough notes, although he provided

the species names for the plates. According to Kottelat

(1988: 73), most plates of the first fascicle were engraved

and colored before Spix’s death and bear the names Spix

intended to give them. As we have seen above, however,

a name associated with an illustration, published before

1931, is indeed sufficient to establish availability (and

this was the case also in the 3“ edition of the Code).

Thus, in this regard, Kottelat’s interpretation of the Code

was incorrect. As Wagner did in the case of the mollusks,

in several cases Agassiz did not follow Spix’s names on

the plates of the first fascicle (regarding the names as

inappropriate or in a “barbarian language” or the fishes

as misidentified) and therefore introduced other names

in the text. In contrast, the plates of the second fascicle

bear names consistent with the text, undoubtedly given

by Agassiz. Nevertheless, Kottelat decided, following

Recommendation 51B of the Code (3 edition), that the

species names should be cited as “Agassiz in Spix and

Agassiz”, arguing that this seemed dlesizable to him “for

bibliographic purposes”.

Notwithstanding this entire argument, authorship of

the work is not regulated by the Code and the most

appropriate attribution of the work remains open for dis-

cussion. Spix’s name forms part of the title, since the

relative pronoun “quae” [= “that’] on the second line

of the title page (Figure 1) opens a subordinate sentence

with a subject (“Dr. J. B. de Spix”) that requires one or

more verbs (“collegit et pingenda curavit”) to give mean-

ing to the sentence. But conversely, Wagner's and the

editors’ contributions are not part of the title—Wagner

“arranged, described and illustrated with observations”

(see above) the work of Spix. The book does not contain

only Spix’s concepts, nor did Wagner communicate with

him to clarify the reasons why Spix considered some of

the shells as belonging to new species. Indeed there is

no evidence that they ever met. In fact, Spix’s views were

contradicted several times in the text. So, the book is

not a unity (i.e., a Spix-and-Wagner production) but an

earlier work by Spix, critiqued and modified by Wagner.

Therefore we consider that Wagner, being responsible

for presenting the material following his own criteria,

and deciding the fate of the work after Spix’s death, is

to be credited with the final product, as the only author

of a book into which Spix’s contribution has been incor-

porated.

We therefore consider it most appropriate to treat

THE NAUTILUS, Vol. 118, No. 2

Wagner as the sole author of the book, which should

then be cited as

WAGNER, J.A. 1827. Testacea fluviatilia quae in itinere

per Brasiliam annis MDCCCXVII-MDCCCXxX jussu et

auspiciis Maximiliani Josephi I. Bavariae regis augustis-

simi suscepto collegit et pingenda curavit Dr. J. B. de

Spix, quondam ordinis regii corone Bavarice civilis

eques, academiz scientarum Bavarice socius ordinarius,

musei regii zoologici, zootomici et ethnographici conser-

vator rel. C. Wolf, Monachii [= Munich]. iv + [ii] + 36

pp. 29 pls.

Another issue of the work was also published in Leip-

zig, but we consider this to have been published sub-

sequent to the Munich edition, as discussed below.

AUTHORSHIP OF THE SPECIES

The following discussion relates to the Munich issue, as

the Leipzig issue, published after the Munich issue (see

below), has no bearing on nomenclature.

Spix consistently provided binomina in the figure leg-

ends for the new species he illustrated in the plates. He

thereby accomplished the minimum pre-1931 conditions

of availability for those names (Code, Articles 11 and 12),

except for publication. It seems that he also provided

two to three line diagnoses (see below). Arguably, be-

cause Wagner provided the longer descriptions (for most

species), he also could be construed as an author, since

the plates, diagnoses, and longer descriptions were pub-

lished simultaneously. So, arguably, authorship could un-

doubtedly be considered as “Spix and Wagner” for all

those species originally named by Spix.

Of course, Spix failed to publish his new species in-

dependently, but the act of being published by another

person (Wagner) does not necessarily deprive Spix of

nomenclatural authorship because, “if it is clear from the

contents that some person other than the author of the

work is alone responsible both for the name or act and

for satisfying the criteria of availability other than actual

publication [our italics], then that other person is the

author of the name or act” (Code, Article 50.1.1). Wag-

ner was careful to keep authorships recognizable by

identifying them in the text and in the index [the “Ta-

bulae explicatae”]. So, the illustrations, created by the

deceased Spix, with associated names that were attri-

buted to Spix, were published as a corpus together with

an index in which each plate was listed in Spix’ se-

quence. Thus, the names attributed to Spix in the work

should indeed be attributed to Spix alone and not to Spix

and Wagner. This has been the conclusion reached by

earlier authors (e.g., Ihering, 1890; Morrison, 1954;

Haas, 1969: Fechter, 1983a, b).

Furthermore, regarding precedence of Spix and Wag-

ners names and nomenclatural acts, the Code (Recom-

mendation 24B) states that “Zoologists acting as First

Revisers to determine the precedence of identical names

published in the same or different works, and on the

same day, are advised to follow attributions by authors

concerned if these are known”. And the Code (Article

R. H. Cowie et al., 2004

Page 77

50.6) states that “When two or more identical names for

the same taxonomic taxon are published on the same

date, by different authors in the same or different works,

their precedence (and hence the authorship of the

name) is determined by the application of Article 24”.

Therefore, because Wagner attributed the names to

Spix, the latter's names take precedence, even though

published simultaneously with the former’s names. The

following example is illustrative. Wagner considered Am-

pullaria archimedes Spix a synonym of A. zonata “Wag-

ner on the basis of Spix’s figure alone; ie. “... quae

mihi tantum ex hac figura cognita est” [= “... which I

only know from this figure”]. Therefore, A. archimedes

cannot be co-authored by Wagner, because he did not

consider it a valid species. The species has to be credited

to Spix; thus, Wagner's act was to create a subjective

synonym.

Both authors worked independently and did not share

their taxonomic concepts, a further reason for author-

ship of the taxa not being Spix and Wagner, but Spix (or

Wagner in a few cases) alone. Table 1 lists all the names

of both authors, with their correct authorship and status.

Wagner's rationale for attributing authorship does not

follow current rules of nomenclature. He accepted as of

Spix only those species that he considered valid and cor-

rectly combined with a generic name (25 of the 84 spe-

cies illustrated by Spix). In modem terms, citation of

these species should be either as of Spix alone (Code,

Article 50.1.1) or as of Spix in Wagner (Code, Recom-

mendation 51E). On the basis of the following discus-

sion, we consider that the names should be cited as of

Spix in Wagner, with a small number of exceptions.

In the 13 cases in which Wagner transferred one of

Spix’s species to another genus he presented himself as

the author (Ampullaria lineata Wagner for Helix lineata

Spix, Pupa exesa Wagner for Clausilia exesa Spix, Helix

clausa Wagner for Tomigerus clausus Spix, etc.). This

was common practice for several decades in the late

1700s and early 1800s. He did not question the validity

of such species, but simply re-assigned them to genera

established by Lamarck and other authors (as explained

in the Preface, above). This action is now treated as es-

tablishing a new combination, so a reference such as

“Bulimus sylvaticus Wagn. (Columna sylvatica Spix)’ [in

the “Tabulae explicatae”| is now treated as Bulimus syl-

vaticus (Spix) or Bulimus sylvaticus (Spix) Wagner

(Code, Recommendation 51G), although the latter prac-

tice has rarely, if ever, been used in mollusks.

Wagner changed three names because those given by

Spix were preoccupied (i.e., Bulimus hyalinus Wagner

for B. fragilis Spix, non Lamarck; Bulimus magus Wag-

ner for B. inflatus Spix, non Lamarck; Helicina variabilis

Wagner for H. fasciata Spix, non Lamarck). In these cas-

es, Spixs names are junior primary homonyms and

therefore invalid.

Wagner also changed other names, but for no explicit

reason (e.g., Melania scalaris Wagner for Aylacostoma

glabrum Spix; Unio caudatus Wagner for Diplodon fur-

catus Spix). In these cases, Spix’s names are valid, while

Wagner's are unnecessary replacement names and thus

junior objective synonyms (Code, Article 72.7).

Finally, in some cases Wagner added his name follow-

ing what appear to be replacement names for some of

Spix’s binomina; however, the new names were not bi-

nominal and therefore not available. For example, An-

odon lituratus Spix became “Anodontis obtusi specimen

junius Wagn.” [= “a young specimen of Anodon obtusus

Wagner’, although he attributed Anodon obtusus to

Spix]. Similarly, and although Wagner did not add his

name in this case, Ampullaria figulina Spix became “Am-

pullaria lineata minor” [= a smaller Ampullaria lineata].

In this instance “minor” is not to be considered a sub-

specific name forming part of a valid trinomen, but as a

purely descriptive term. We conclude from these in-

stances that Wagner aimed not to claim authorship of

the species but to establish subjective synonymy.

Again, on the basis of all this evidence, the names are

to be cited as of “Spix in Wagner’, except in the case of

the three invalid, preoccupied names of Spix, the re-

placements for which should be cited as of Wagner

alone. The remainder of Wagner's names are unneces-

sary replacement names.

RELATIONSHIP OF THE TEXT TO THE PLATES

Most species are described twice. The first text, in a

larger font, is a two to three line diagnosis mostly based

on shape and colors, as are visible in Spix’s figures. The

second text, in a smaller font, is a more detailed descrip-

tion that also includes some measurements and infor-

mation that can only be accurately assessed on actual

shells (e.g., whorl number). An example is given in Fig-

ure 4. While it is possible that the first two-line text for

each species is a description by Wagner of Spix’s figure,

and that the second text was written after comparing the

plate with additional materials, we have no compelling

evidence that this is the case. Rather, we believe that

the short first text is attributable to Spix and only the

longer second text to Wagner, for the following reasons.

The customary practice among conchologists at the

time when describing new species was either to write all

the text in Latin, ie., a single description sometimes fol-

lowed by comments (e.g., many papers in the Zeitschrift

fiir Malakozoologie by Philippi, Pfeiffer, etc., Pfeiffer’s

Monographia Heliceorum Viventium), or to give a short

Latin diagnosis, followed by additional descriptions and

comments in a vernacular language (e.g., the Voyage of

dOrbigny, contributions to the Systematisches Conchy-

lien-Cabinet von Martini und Chemnitz by Philippi,

Kiister, etc., Reeve’s Conchologia Iconica). Neither of

these patterns is followed. When Wagner shared Spix’s

concept and name for the species, then the two texts are

arranged directly one after the other (e.g., Ampullaria

gigas, A. papyracea, A. rosea). When, for some reason,

Wagner changed the original name, usually because of a

genus change (e.g., Melania tuberculata Wagner for Ay-

lacostoma tuberculatum Spix), or to give a replacement

name (e.g., Pupa inflata Wagner for Clausilia pupoides

Page 78 THE NAUTILUS, Vol. 118, No. 2

Table 1. The names of Spix and Wagner listed in the order of Spix’s plates and as they appear in the “Tabulae explicatae”, with

explanations of their treatment by Wagner if different from their treatment by Spix, and additional comments, as appropriate. In

the “Tabulae explicatae” Wagner's treatment of the names appears in plain Roman type, with Spix’s names in parentheses and italic

type on the same line, if Wagner's treatment of them differed. All their new names are nomenclaturally available. Note that there

are 29 printed plates, but that the Tabulae explicatae only number 27; the final two lines of the Tabulae explicatae lack the plate

numbers, though the plates themselves have the correct legends: “Tab. XXVIII” and “Tab. XXIX”.

Plate and

Names as listed in the figure Names as they appear Treatment by Wagner if different from that by Spix, with

“Tabulae explicatae” numbers on the plates additional comments in square brackets

GASTROPODA

Ampullaria Gigas Spix PI. I, figs. 1,2 © AMPULLARIA Gigas.

Ampullaria zonata Spix Pl. I, fig. 1 AMPULLARIA 1. zona- [Listed as Ampullaria zonata Spix in the Tabulae expli-

ta. 2. Archimedes. catae but Ampullaria zonata Wagner in the text.]

Ampullaria Archimedes PI. Il, fig. 2 [see Ampullaria zonata] Smaller variety of A. zonata Spix.

Spix

HA OUR olivacea Pl. III, fig. 1 AMPULLARIA 1. oliva- A. guyanensis Lamarck, 1822, given as a synonym but

Spix cea. 2. quercina. Spix’s name retained as valid. Compared to A. rugo-

sa Lamarck, 1801, A. globosa Swainson, 1823, al

A. leucostoma Seetinacin. 1823. [Junion primary

homonym of Ampullaria olivacea Lamarck, 1816.]

Ampullaria quercina Pl. Il, fig. 2 [see Ampullaria olivacea] Smaller variety of . olivacea Spix.

Spix

paola papyracea __ PI. IV, figs. 1, 2. AMPULLARIA 1. 2. pa-

Spix pyracea. 3. rosea. 4. fi-

gulina. 5. Cyclostoma.

Ampullaria ? rosea Spix PI. IV, fig. 3 [see Ampullaria papyra- Valid species, although its identity as an Ampullaria

cea] doubted; considered a terrestrial snail in the genus

Bulimus.

Ampullaria figulina Pl. IV, fig. 4 [see Ampullaria papyra- Smaller, yellowish variety of Ampullaria lineata (Spix).

Spix cea]

Rene? cyclostoma PI. IV, fig. 5 [see Ampullaria papyra- Extreme affinity to Ampullaria effusa (Miiller, 1774)

Spix cea] Swainson, 1823 [= Nerita effusa Miiller, 1774] not-

ed.

Helix fasciolata Spix Pl. V, fig. 1 HELIX 1. fasciolata. 2 Variety of Ampullaria crassa Swainson, 1823, with a

lineata. 3. crass. 4. narrow umbilicus.

HELICINA exumbilica-

ta.

Helix lineata Spix Pl. V, fig. 2 [see Helix fasciolata] Placed in Ampullaria as A. lineata “Wagn.” |= (Spix)].

A. fasciata Swainson, 1822, given as a synonym but

Spix’s name retained as valid. |A. fasciata Swainson,

1822, is a misidentification of A. fasciata Lamarck,

1816 (Cowie and Thiengo, 2003).] Compared to A.

reflexa Swainson, 1823. [Junior primary homonym of

Helix lineata Renier, 1804, and Helix lineata Say,

1817.]

Helix crassa Spix PI. V, fig. 3 [see Helix fasciolata] Not a new name; = Ampullaria crassa Swainson,

1823.

Helicina exumbilicata Pl. V, fig. 4 [see Helix fasciolata] Variety of Ampullaria crassa Swainson, 1823, lacking

Spix an umbilicus.

Bulimus terrestris Spix Pl. VI, fig. 1 BULIMUS 1. terrestris. [see Bulimus corrugatus, below]

2. durus. 3. fragilis. 4.

virgatus.

Bulimus corrugatus Pl. VI, fig. 1 [see Bulimus terrestris] Bulimus terrestris Spix synonymized with B. corruga-

Wagner tus “Wagner” and considered to be but a juvenile of

Bulimus ovatus (Miiller, 1774) [= Helix ovata Miill-

er, 1774], although B. corrugatus “Wagner” retained

as the valid name. [Not a new name; = Bulimus

corrugatus Bruguiére, 1792. ]

Bulimus durus Spix Pl. VI, fig. 2 [see Bulimus terrestris |

Bulimus fragilis Spix Pl. VI, fig. 3 [see Bulimus terrestris] Junior secondary homonym of Bulimus fragilis (Mon-

tagu, 1803) Lamarck, 1822 [= Helix fragilis Monta-

gu, 1803].

Bulimus hyalinus Wag- Pl. VI, fig. 3 [see Bulimus terrestris] New replacement name for Bulimus fragilis Spix, non

ner

Bulimus fragilis (Montagu, 1803) Lamarck, 1822.

R. H. Cowie et al., 2004

Table 1. Continued.

Names as listed in the

“Tabulae explicatae”

Plate and

figure

numbers

Page 79

Names as they appear

on the plates

Treatment by Wagner if different from that by Spix, with

additional comments in square brackets

Bulimus virgatus Spix

Bulimus angiostomus

Wagner

Bulimus inflatus Spix

Bulimus Magus Wagner

Bulimus perlucidus

Spix

Bulimus lituratus Spix

Bulimus vittatus Spix

Bulimus zebra Spix

Bulimus lineatus Spix

Achatina perversa Spix

Bulimus vitreus Spix

Bulimus decapitatus

Spix

Aylacostoma tubercula-

tum Spix

Aylacostoma glabrum

Spix

Melania scalaris Wag-

ner

Achatina pavonina Spix

Pl. VI, fig. 4

Pl.

Al.

Pl.

Al.

Pl.

All.

Pl.

VII, fig.

VII, fig,

VII, fig.

VIL, fig.

VIII, fig.

VI, fig. 4

VII, fig. 1

VII, fig. 3

VII, fig. 6

[see Bulimus terrestris |

BULIMUS 1. inflatus. 2.

perlucidus. 3. lituratus.

4. vittatus. 5. Zebra. 6.

lineatus.

see Bulimus inflatus

[see Bulimus inflatus

see Bulimus inflatus

see Bulimus inflatus]

[see Bulimus inflatus|

1. ACHATINA perversa.

2. BULIMUS vitreus.

3. BULIMUS decapi-

tatus. 4. AYLACOS-

TOMA tuberculatum.

5. glabrum.

[see Achatina perversa

ACHATINA 1. pavonina.

2. pulchella. 3. 4. floc-

cosa.

Incomplete shell of Bulimus angiostomus Wagner |=

Stenostoma capueira Spix].

[see additional listing, below]

Junior primary homonym of Bulimus inflatus Olivier,

1801, and Bulimus inflatus Lamarck, 1822.

New replacement name for Bulimus inflatus Spix, non

Olivier, non Lamarck.

Helix lita |“Freycinet, Voyage autour du monde”; =

Helix lita Férussac in Quoy and Gaimard] given as a

synonym but Spix’s name retained as valid.

Treated as a valid species. Also mentioned a larger va-

riety with a basal, dark color band.

[Primary junior homonym of Bulimus zebra Olivier,

1801, and Bulimus zebra Perry, 1810. Perhaps not

intended by Spix as a new name, but a misidentifi-

cation of Buccinum zebra Miiller, 1774].

Exreme similarity to “Bulimus radiatus” noted.

[Junior primary homonym of Bulimus lineatus Drapar-

naud, 1801. ? = Bulimus radiatus de Blainville,

1825—apparently the only Bulimus radiatus de-

scribed. |

Synonym of Achatina melanostoma Swainson. Dextral

and sinistral shells of this species mentioned as de-

posited in the Munich Museum.

[Not a new name; = Achatina perversa Swainson,

1821 (originally spelled “Achatinia” by Swainson).

The original name of “Achatina melanostoma Swain-

son” is melastoma. Even if melanostoma were pref-

erable from a scholarly perspective, a poor latiniza-

tion is not to be corrected (Code, Article 32.5.1).

Because Wagner cited the original and the changed

name, and used the latter as valid, the change is

considered demonstrably intentional (Code, Article

33.2.1). Wagner’s nomenclatural act being an unjus-

tified emendation, Achatina melanostoma Wagner,

1827, is a junior objective synonym of Achatina me-

lastoma Swainson, 1823.]

Synonym (an incomplete shell) of Bulimus perlucidus

Spix.

Extreme similarity to Bulimus decollatus (Linnaeus,

1758) Bruguiére, 1789 [= Helix decollata Linnaeus,

1758] noted.

Placed in Melania as Melania tuberculata “Wagn.” [=

(Spix)]; compared to Melania truncata Lamarck,

1822.

Replaced by Melania scalaris Wagner.

[Unnecessary replacement name for Aylacostoma gla-

brum Spix; junior objective synonym of A. glabrum

Spix.]

Not mentioned in the text; synonym of Bulimus galli-

nasultana Lamarck, 1822, in the “Tabulae explica-

>»

tae .

Page 80

Table 1. Continued.

THE NAUTILUS, Vol. 118, No. 2

Names as listed in the

“Tabulae explicatae”

Achatina pulchella Spix

Achatina floccosa Spix

Columna maritima Spix

Columna 8-gyrata Spix

Columna bulimea Spix

Bulimus Spixit Wagner

Columna sylvatica Spix

Lymnaeus papyraceus

Spix

Strophocheilus

Haemastromus Spix

Strophocheilus Almeida

Spix

Auris melastoma Spix

Auris signata Spix

Auris vittata Spix

Stenostoma auritum

Spix

Stenostoma Purw Spix

Bulimus angulatus

Wagner

Stenostoma Capueira

Spix

Bulimus angiostomus

Wagner ;

Plate and

figure

numbers

Names as they appear

on the plates

Treatment by Wagner if different from that by Spix, with

additional comments in square brackets

Pl. IX, fig. 2

Pl. IX, figs. 3, 4

Pl. X, fig. 1

Pl. X, fig.

Pl. X, fig. 3

Pl. X, fig. 4

Pl. X, fig. 5

Pl. XI, fig. 1

Pl. XI, figs. 2, 3

Pl. XII, figs. 1,

Pl. XII, fig. 3

Pl. XI, fig. 4

Pl. XIII, figs. 1,

Pl. XIII, fig. 3

Pl. XIII, fig, 4

Pl. XIII, fig. 4

[see Achatina pavonina]

COLUMNA 1. maritima.

2. 8-gyrata. 3. bulimea.

4, sylvatica. 5. LYM-

NAEUS papyraceus.

see Columna maritima

STROPHOCHEILUS 1.

Haemastomus. 2. 3. Al-

meida.

[see Srophocheilus Hae-

mastomus |

AURIS 1. 2. Melastoma.

3. signata. 4. vittata.

see Awris melastoma]

see Auris melastoma]

STENOSTOMA 1. 2. au-

ritum. 3. Puri. 4. Ca-

pueira.

see Stenostoma auritum|

[see Stenostoma auritum|

Synonym of Bulimus undatus Bruguiére, 1789. Buccin-

um zebra Miiller, 1774, among other names, listed

in synonymy, but Bruguiére’s name retained as valid.

Placed in Bulimus as Bulimus floccosus “Wagn.” [=

(Spix)].

Synonym of “Bulimus calcareus Bruguiére” [= Helix

calcareus Born, 1778].

Synonym of “Bulimus calcareus Bruguiére” [= Helix

calcareus Born, 1778].

Replaced by Bulimus spixii Wagner.

[Unnecessary replacement name for Columna bulimea

Spix; junior objective synonym of C. bulimea Spix.]

Placed in Bulimus-as Bulimus sylvaticus “Wagner” [=

(Spix)].

Spix’s original spelling, “Lymnaeus papyraceus” (leg-

end of plate X) changed to “Limnaeus papyraceus

Spix’.

ee Pfeiffer, 1821 (emendation of Limneus Dra-

pamaud, 1801, Limnus Montfort, 1810, and Lym-

naeus Brand, 1810) is a junior synonym of Lymnaea

Lamarck, 1799.]

Synonym of “Bulimus ovatus Bruguiére” [= Helix ova-

ta Miiller, 1774].

Synonym of “Bulimus virgineus Bruguiére, 1789” [=

Helix pudica Miiller, 1774].

Not a new name but a reference to Bulimus melasto-

mus Swainson, 1820; emended to “Bulimus melanos-

tomus Swainson’, citing the original name together

with the emended name, the latter used as valid.

[Wagner's emendation was unjustified, so Bulimus

mealnostomus Wagner is an available, junior objec-

tive synonym of Bulimus melastomus Swainson,

1820 (Code, Article 33.2.3).]

Placed in Auwricula as Auricula signata “Wagner” [=

(Spix)]; compared with Auricula sileni Férussac,

1807.

Faded specimen of Awricula signata Spix.

Synonym of Auricula leporis “Lamarck” [= “Brugui-

ére”, in Férussac, 1807].

Replaced by Bulimus angulatus Wagner. The name

Purti was printed with a written accent in the plate

legend but without an accent in the Tabulae explica-

tae (Puru).

[Unnecessary replacement name for Stenostoma puru

Spix; junior objective synonym of Stenostoma puru

Spix. Spix’s name is the name of the Purti River, in

apposition; perhaps Wagner considered such a name

unacceptable, since he replaced both such names of

Spix (see B. angiostomus, below).]

Replaced by Bulimus angiostomus Wagner.

[Unnecessary replacement name for Stenostoma capueira

Spix; junior objective synonym of Stenostoma capueira

Spix. Spixs name is the name of the Capueira River,

in apposition. See B. angulatus, above.]

R. H. Cowie et al., 2004

Table 1. Continued.

Page 81

Names as listed in the

“Tabulae explicatae”

Clausilia exesa Spix

Clausilia striata Spix

Clausilia 6-dentata Spix

Clausilia pupoides Spix

Pupa inflata Wagner

Pupa elatior Spix

Navicula fasciata Spix

Helix Navicula Wagner

Tomigerus clausus Spix

Helicina Pyramidella

Spix

Helicina fasciata Spix

Helicina variabilis

Wagner

Helicina flava Spix

Solarium Serpens Spix

Helix Pellis serpentis

Chemnitz

Helix punctata Wagner

Solarium candidum

Spix

Solarium vitreum Spix

Helix perspectiva Wag-

ner

Solarium imperforatum

Spix

Plate and

figure

numbers

Pl. XIV, fig.

ran

Pl. XIV, fig.

Pl. XIV, fig,

me & OO bo

Pl. XV, fig. 1

Pl. XV, figs.

Pl. XV, figs. 4,

PL XVI figs. 1,

Pl. XVI, figs. 3,

Pl. XVI, fig. 5

Pl. XVII, figs.

iL, 2

Pl. XVII, fig. 1

Pl. XVII, fig. 2

Pl. XVII, figs.

3,4

Pl. XVII, fig. 5

Pl. XVII, figs.

345

Pl. XVII, fig. 6

Names as they appear

on the plates

Treatment by Wagner if different from that by Spix, with

additional comments in square brackets

CLAUSILIA 1. exesa. 2

striata. 3. 6-dentata. 4.

pupoides.

[see Clausilia exesa]

1. PUPA elatior. 2. 3.

NAVICULA fasciata.

4.5. TOMIGERUS

clausus.

see Pupa elatior]

see Pupa elatior|

HELICINA 1. 2. Pyram-

idella. 3. 4. fasciata. 5.

flava.

see Helicina Pyramidel-

la]

see Helicina Pyramidel-

la]

see Helicina Pyramidel-

la]

SOLARIUM 1. 2. Ser-

pens. 3. 4. candidum.

5. vitreum. 6. imper-

foratu mM, Lo pygm aeum.

see Solarium Serpens

see Solarium Serpens|

see Solarium Serpens

[see Solarium Serpens|

Placed in Pupa as Pupa exesa “Wagner” [= (Spix)].

Placed in Pupa as Pupa striata “Wagner” [=

Placed in Pupa as Pupa sexdentata “Wagner” |=

Replaced by Pupa inflata Wagner.

Compared to Clausilia sexdentata Spix. [Unnecessary

replacement name for Clausilia pupoides Spix; ju-

nior objective synonym of Clausilia pupoides Spix.]

(Spix)].

Replaced by Helix navicula Wagner. [The genus-group

name Navicula Spix is a junior primary homonym of

Navicula Blainville, 1825.]

[Unnecessary replacement name for Navicula fasciata

Spix; junior objective synonym of Navicula fasciata

Spix.]

Placed in Helix as Helix clausa “Wagner” [= (Spix)].

Placed in Helix as Helix Pyramidella “Wagner” [=

(Spix)].

Junior primary homonym of Helicina fasciata Lamarck,

1822; replaced by Helicina variabilis Wagner.

New replacement name for Helicina fasciata Spix.

Variety of Helicina variabilis Wagner.

Spix’s apical (pl. XVII, fig. 1) and basal (pl. XVII, fig.

2) views of this species identified as two distinct but

very similar species (see below, and see the discus-

sion of this species in the text).

The apical view (pl. XVII, fig. 1) of Solarium serpens

Spix identified by Wagner as Helix pellisserpentis

“Chemnitz” [= Gmelin, 1794, since Chemnitz is un-

available].

New name for Solarium serpens Spix of pl. XVII, fig.

2 (non Spix of pl. XVII, fig. 1).

[Junior primary homonym of Helix punctata Miiller,

1774]

[see Helix perspectiva, below]

Solarium candidum Spix and S. vitreum Spix listed as

adult (“testa completa adulta”) and younger (“testa

junior”) specimens, respectively of Helix perspectiva

Wagner.

[Junior primary homonym of Helix perspectiva Megerle,

1816, and Helix perspectiva Say, 1817. As first revisers,

we treat Helix perspectiva Wi agner as an unnecessary

new name for Solarium ceundbichoica Spix, not for S. vi-

treum Spix. Helix perspectiva Wagner is therefore a ju-

nior objective synonym of S. wagons Spix, and So-

larium vitreum Spix is a junior subjective synonym of

S. candidum Spix, according to Wagner's opinion.

Replaced by Helix vitrina Wagner.

Dyevarm (2X9)

Page 82

Table 1. Continued.

THE NAUTILUS, Vol. 118, No. 2

Treatment by Wagner if different from that by Spix, with

additional comments in square brackets

[Unnecessary replacement name for Solariwm imper-

foratum Spix; junior objective synonym of Solarium

Plate and

Names as listed in the figure Names as they appear

“Tabulae explicatae” numbers on the plates

Helix vitrina Wagner Pl. XVIL, fig. 6 [see Solarium Serpens|

imperforatum Spix.|

Solarium pygmaeum Pl. XVIL, fig. 7 [see Solarium Serpens]

Spix

Helix nana Wagner

Planorbis ferrugineus

Spix

Planorbis olivaceus

Wagner and Spix

Planorbis nigricans

Spix

Planorbis albescens

Spix

Planorbis viridis Spix

Planorbis lugubris

Wagner

BIVALVIA (see footnote)

Anodon giganteus Spix

Anodon trapezeus Spix

Anodon rotundus Spix

Anodon anserinus Spix

Anodon longinus Spix

Anodon trigonus Spix

Anodon obtusus Spix

Anodon lituratus Spix

Pl. XVII, fig. 7

Pl. XVI, fig. 1

Pl. XVIII, fig. 2

Pl. XVIII, figs.

3, 4

Pl. XVIII, fig. 5

Pl. XVIII, fig. 6

Pl. XVIII, figs.

3-6

Pl. XIX, figs.

Pl. XX, fig. 1

Pl. XX, figs. 2-4

Pl. XXI, figs.

Pl. XXII, fig,

Pl. XXII, fig.

PI. XXII, fig.

Pl. XXII, fig,

Hw CO

[see Solarium Serpens|

PLANOBBIS 1. ferrugi-

neus. 2. olivaceus. 3. 4.

nigricans. 5. albescens.

6. viridis.

[see Planorbis ferrugi-

neus |

[see Planorbis ferrugi-

neus |

[see Planorbis ferrugi-

neus |

[see Planorbis ferrugi-

neus |

[see Planorbis ferrugi-

neus |

ANODON giganteum. 1.

juv. 2. adult.

ANODON 1. trapezeum.

2. 4, rotundum.

[see Anodon trapezeus|

ANODON anserinum.

ANODON 1. longinum.

2. trigonum. 3. obtu-

sum. 4. lituratum.

[see Anodon longinus]

[see Anodon longinus|

Replaced by Helix nana Wagner.

[Junior primary homonym of Helix nana Martens, 1824,

and Helix nana Megerle, 1824. Unnecessary replace-

ment name for Solarium pygmaeum Spix; junior ob-

jective synonym of Solarium pygmaeum Spix.]

Synonym of Planorbis olivaceus Spix.

Compared to “Planorbis corneus” [= Helix cornea

Linnaeus, 1758].

[The only claim of joint authorship is in the “Tabulae

explicatae”. However, no author is mentioned for

this species in the text (p. 26); if there was a reason

for this departure, it is not evident from the work

itself, and species authorship is therefore assignable

to Spix alone.]

[see Planorbis lugubris, below]

Planorbis nigricans Spix, P. albescens Spix, and P. viri-

dis Spix considered as juvenile specimens of P. lu-

gubris Wagner. Compared to “Planorbis corneus” [=

Helix cornea Linnaeus, 1758).

[Unnecessary replacement name. Planorbis nigricans

Spix is the first of the three names listed as Planor-

bis lugubris Wagner in the “Tabulae explicatae”. We,

as first revisers, take P. lugubris Wagner to be the

replacement name for P. nigricans; the former is

thus a junior objective synonym of the latter. Pla-

norbis albescens Spix and P. viridis Spix are then

subjective synonyms according to Wagner's opinion. ]

Anodon crassus Swainson, 1823, considered a variety

of this species, even though Swainson’s name has

priority.

Anodon membranaceus |= Mytilus memranaceus Ma-

ton, 1811] given as a synonym, but Spix’s name re-

tained as valid. Compared to Anodon trapezeus Spix.

Compard to Anodon giganteus Spix. [Though the fig-

ures have itary. ih (outer view) and 2 (inner view

of both valves), the legend does not mention any

numbers].

Compared to “Anodon sulcatus Lamarck” [= Anodon-

ta sulcatus Lamarck, 1819].

Synonym (young specimen) of Anodon obtusus Spix.

R. H. Cowie et al., 2004

Table 1. Continued.

Page 83

Names as listed in the

“Tabulae explicatae”

Plate and

figure

numbers

Names as they appear

on the plates

Treatment by Wagner if different from that by Spix, with

additional comments in square brackets

Anodon radiatus Spix

Anodon siliquosus Spix

Anodon pygmaeus Spix

Anodon ensiformis Spix

Aplodon inermis Spix

Tetraplodon pectinatus

Spix

Cyclas bahiensis Wag-

ner

Diplodon ellipticus Spix

Diplodon rotundus Spix

Diplodon furcatus Spix

Unio caudatus Wagner

Diplodon rhombeus

Spix

Triplodon rugosus Spix

Pl. XXIII, fig. 1

Pl. XXIII, fig. 2

Pl. XXIII, figs.

3,4

Pl. XXIV, figs.

1,3

Pl. XX, figs. 1,

Pl. XXVI, figs.

i, 2

Pl. XXVI, figs.

3, 4

Pl. XXVII, figs.

i, 2

Pl. XXVII, figs.

i, 2

Pl. XXVIII, figs.

1,2

Pl. XXIX, figs.

1,2

ANODON 1. radiatum.

2. siliquosum. 3. 4.

pygmaeum.

[see Anodon radiatus]

ANODON ensiforme.

1. 2. APLODON inerme.

3. 4. TETRAPLO-

DON pectinatum. 5.

6. CYCLAS bahiensis.

[see Anodon inermis]|

[see Anodon inermis]

DIPLODON 1. 2. ellipti-

cum. 3. 4. rotundum.

[see Diplodon ellipticus]

DIPLODON furcatum.

[see Diplodon furcatus]

DIPLODON rhombeum.

TRIPLODON rugosum.

Compared to “Anodon glaucus Humboldt” [= Ano-

donta glaucus Valenciennes, 1827].

Compared to Anodon longinus Spix.

Synonym (young specimen) of Anodon siliquosus Spix.

[Though the figures have numbers 1 (outer view) and

2 (inner view of both valves), the legend does not

mention any numbers].

Possible synonym of Anodon rotundus Spix.

Placed in Unio as Unio pectinatus “Wagner” [=

(Spix)]. Castalia ambigua Lamarck, 1819, given as a

synonym, but Spix’s name retained as valid.

Compared to Cyclas fontinalis Draparnaud, 1801.

[Authorship should be Spix not Wagner; probably a

lapsus calami, because the legend on plate XXV

reads “Cyclas bahiensis” (Spix’s original spelling) and

the text subtitle on p. 32 reads “Cyclas bahiensis

Spix’.]

Placed in Unio as Unio ellipticus “Wagner” [= (Spix)].

Compared to Unio pictorum (Linnaeus, 1758).

Placed in Unio as Unio rotundus “Wagner” [= (Spix)].

Mya variabilis Maton, 1811, given as a synonym but

Spix’s name retained as valid.

Replaced by Unio caudatus Wagner. [No numbers

printed either with the figures (outer view and inner

view of both valves) or in the legend].

Hyria avicularis Lamarck, 1819, given as a synonym

but Spix’s name retained as valid.

[Unnecessary replacement name for Diplodon furcatus

Spix; junior objective synonym of Diplodon furcatus

Spix.]

Placed in Unio as Unio rhombeus “Wagner” [=

(Spix)]. Compared to Unio peruvianus Lamarck,

1819. [Though the figures have numbers 1 (outer

view) and 2 (inner view of both valves), the legend

does not mention any numbers].

Placed in Unio as Unio rugosus “Wagner” [= (Spix)].

Compared to Hyria corrugata Lamarck, 1819.

[Though the figures have numbers | (outer view)

and 2 (inner view of both valves), the legend does

not mention any numbers].

Note—Most Greek nouns ending in -on are neuter. However, the word -odon (= tooth) and its derived genera ending in -odon are

masculine. Spix apparently did not realize the difference and gave neuter endings (-wm or -e) to all species in Anodon, Diplodon,

Triplodon, and Tetraplodon. All these names are, therefore, incorrect original spellings that were rightly corrected by Wagner in

the “Tabulae explicatae” and in the text (Code, Aritcles 31.2, 32.5, 34.2).

Spix), he provided Spix’s species name after the short

diagnosis, before describing the species in the longer

description. When he combined two of Spix’s species as

varieties under a single Wagner species (e.g., Helix li-

neata Spix and Ampullaria figulina Spix under Ampul-

laria lineata Wagner), or synonymized two of Spix’s spe-

cies

this

(Ampullaria zonata Spix and A. archimedes Spix un-

der A. zonata Wagner), he provided another, still shorter

diagnosis on the same line as each of Spix’s species

names, following the two-line diagnosis, before describ-

ing the species in the longer description. We suggest that

reflects the likelihood that Spix left not only the

Page 84 THE NAUTILUS, Vol. 118, No. 2

AMPULUARIA.

<> |

4. AMPULLARIA papynacea spix. Tb. 1V. Fig. x. o.

A. testa ovato- globosa, tenuissima, longitudinaliter subtilissime striata, nigro-fus-

ca: umbilico angusto: apertura nigra,

7o-ovala slobosa, te issime Arelig Rf 2 : . / :

Testa oblongo-ovata, globosa, tenuissima, fragilis; striis longituidinalibus crebris, tenuibus,

Anfractus quingue Ul Sex convexi, ultimus

maximus, Spira brevis, striis longitudinalibus profundioribus. Apertura ovato-oblonga; labrum

oOo”

acutum, lenue, margine sinistro subreilexo, Umbilicus angustus , longituiinalis,

strias Iransversas, remotas, Obsoletes decussantibus.

Color testae

nigro — fuscus; ultimus anfractus interdum fascia olivaceo brannea cinctus, Spirafusco-rubra; Opers

tura nigra.

Longitude 2 poll. 2 lin; lat. poll. g lin.

Tlubitat in flactis et stagnis Provinciarum Bahiensis, Pernambucanae et Piauhiensis.

5. AMPULLARIA nosea spix. tab. tv. Fig. 5.

A. testa ovata, yentricosa, tenui, pellucida, longitudinaliter striata, perforata,

albido-rubella; apertura oblongo-ovata, inferne ampla.

Testa oblongo-ovata, ventricosa, tenuis, pellucida, Jongitadinaliter striata. Anfractus quinque

conyexi; ultimus maximus, usque ad medium striis Jongitudinolibus eleganter ornatus , infra me-

dium Isevis, Spira exsertiuscula, obtusa; striis longitudinalibus crebris, Apertura oblongo-ovats ,

superne anfractu penultimo yalde anguslata, inferne ample. Labrum acutum, tenue, margine sini-

stro subreflexum. Umbilicus angustissimus. Color testae rubello-albidus, margo aperturae sinister

Toseus.

Longitude 9 '/; lin; lau. 7 14 lin.

Habitat in aguis Brasiliae auvstralioris.

Obsercatio. Waec species, mihi tantum ex unico specimine cognita, dubic Ampallariis adseri-

benda est, forsan in Bulimi genus amandanda.

6. AMPULLARIA xineara wacn. Tab. V. Fig. et Tab. IV. Fig. &

A. testa ovato-globosa, olivaceo-virente aut lutescente, fasciis obscure purpurets

ornata; spira elevata; umbilico mediocri; apertura alba, transversim fasciata.

5) VAR. testa majore, adulta, olivaceo-virente, fasciis purpursscentibus cincta: hee

neata Spix, Tab. F. Fig. 2.

Swainson, Zoological /ilustrations. No. 21. Tab. 103.: Ampullaria Sasciata.

. aise : : ; f

h) VAR. testa minore latescente, fasciis fuscis cincta: -dimpallaria figulina, Spiz Tab IV.

Fig. 4.

Figure 4. An example of species descriptions, from p. 3 of the Munich edition of the “Testacea fluviatilia ...”.

R. H. Cowie et al., 2004

Page 85

plates but also the two to three line diagnoses. This con-

curs with Fechter (1983b) who concluded that Spix pro-

duced the lithographs with hand coloration, named the

species, and attributed a brief, two-line diagnosis to

them, and that Wagner elaborated and completed the

descriptions. We are not aware of the particular evidence

Fechter based these statements on. However, this same

conclusion is elaborated on in a paper by Ihering (1890)

on Spix’s unionoid type material (see below).

We also consider that Spix provided the legends to

the plates, thereby permitting his illustrations to be iden-

tified. Comparing the Munich issue (publisher [“Typis”]

C. Wolf) and the Leipzig issue (publisher T. O. Weigel)

is instructive. The title page of the Leipzig issue has the

same wording but slightly different type settings com-

pared to the Munich issue, but it does not have a pub-

lication date. Nevertheless, we consider it logical to de-

duce that the Munich issue was published first, with no

changes to Spix’s legends, and that Wagner made chang-

es subsequently for the Leipzig issue. Our reasoning is

as follows.

The Munich issue has the species names with no at-

tributions of authorship, but the Leipzig issue has attri-

butions included in the legends, but only if Wagner had

changed the concept of Spix’s species. It also lacks the

plate numbers. For instance, in the Munich issue, the

legend of plate I reads “AMPULLARIA Gigas” [on line

1] and “Tab. I.” [line 2]; and, because Wagner made no

change to this species, the legend in the Leipzig issue

simply reads “AMPULLARIA Gigas”, in the same type,

but lacks “Tab. I.”. In contrast, the Munich issue legend

of plate 2 reads “AMPULLARIA 1. zonata. 2. Archi-

medes” [line 1] and “Tab. I.” [line 2], but because Wag-

ner synonymized two of Spix’s species, the Leipzig issue

legend reads “AMPULLARIA zonata. Wer. [= Wagner]”

on line 1 and “A. 1. zonata. 2. Archimedes. Sp. [= Spix]”

on line 2, with the plate number lacking. Similar alter-

ations can be seen on other plates.

Additional instances also suggest that the Munich is-

sue retained Spix’s original legends. For example, in the

Munich issue, the legend on plate XIX reads “Anodon

giganteum”, but “Anodon giganteus Spix” in the “Tabu-

lae explicatae” and on text page 27, the latter name with

the masculine ending being correct (see footnote to Ta-

ble 1). When Wagner referred to one of Spix’s plates, he

gave the incorrect original name, although he used the

correct masculine names in the subtitle for each species.

For instance, on page 30, Wagner treated Anodon ob-

tusus and Anodon siliquosus, but he mentioned as vari-

eties or forms the names Anodon obtusum Spix, Anodon

lituratum Spix, Anodon siliquosum Spix, and Anodon

pygmaeum Spix. This indicates that Spix engraved the

(incorrect) names on the original plates. Wagner's action

in correcting the species endings was mandatory, and

does not affect Spixs authorship (Code, Article 34.2).

Although Spix’s name does not appear on the plates,

all these examples strongly suggest that Spix provided

both the original plates and legends. For the Munich

issue, Wagner, for whatever reason, did not change the

legends to reflect his text, whereas, apparently later, for

the Leipzig issue, he changed the legends so that they

did now reflect his text.

Did Wagner work entirely from Spix’s lithographs,

with no reference to the shells, or did he see at least

some of the shells? Ihering (1890) has argued that Wag-

ner only had access to the lithographs and did not work

with the material. He based this deduction on a com-

parison of the original type material in ZSM in 1888-

1889 (i.e., before the partial destruction of Spix’s type

material during World War II) with Wagner's descrip-

tions in the 1827 volume. Ihering noted not only that

measurements were sometimes inaccurate but also that

Wagner had made what appeared to Ihering (1890: 119)

to be an unbelievable error in terms of the orientation

of the bivalve shells. In fact some of the printed litho-

graphs are reversed (e.g., Aplodon inerme Spix on pl.

XXV figs. 1-2, Anodon trigonus Spix on pl. XXII fig. 2,

and Diplodon ellipticus Spix on pl. XXVI figs. 1-2, as

discussed by Ihering 1890: 126-127, 162, and 163-165,

respectively). Lithographs are mirror images of the ac-

tual plates, and perhaps the engraver assumed that the

bivalve halves were alike and engraved some of the fig-

ures without taking the trouble to reverse them (the gas-

tropods are illustrated correctly). Wagner, apparently not

realizing this, erroneously gave left for right and vice

versa, and, accordingly, anterior/posterior and the wrong

dentition on the valves of the bivalve shells. Further-

more, Ihering noted that a comparison of Spix’s Latin

diagnoses (based on the actual type material) with Wag-

ners more elaborate descriptions shows that the latter

made mistakes that were derived from errors introduced

in the illustrations, that is, features in the illustrations

that are not actually seen on the shells. Ihering therefore

concluded that Wagner had not seen the shells.

At the time, Wagner was working in Erlangen and

there is no definitive evidence that he actually worked

in the Munich collection of Spix until 1832 (see above).

However, Wagners comments on some species strongly

suggest that he had access to the shells. For example,

Wagner occasionally mentioned that he knew a species

of Spix only from Spix’s figure (e.g., Ampullaria archi-

medes). On p. 19, Wagner stated that “Ceterum Pupam

exesam, cum in Museo Monacensi specimen nullum adsit,

accuratis describere non possum’ (Because there is no

specimen in the Munich Museum, I cannot accurately

describe Pupa exesa). The description of Bulimus galli-

nasultana (p. 9) ends with a reference to Lamarck,

which is justified by Wagner, who says: “Speciminibus,

nescio quo casu, in Museo Monacensi omnino deficien-

tibus, hunc Bulimum rarissimum, pretiotissimum non

describere possum” (Because all specimens in the Mu-

nich Museum—I do not know why—are totally defec-

tive, I cannot describe this very rare and most beautiful

Bulimus). This means that, when the Munich Museum

did not have adequate material, Wagner was not able to

elaborate a description, and so he included only the two-

line diagnosis, probably provided by Spix (Pupa exesa),

or copied Lamarck’s description (Bulimus gallinasul-

Page 86

tana). Thus, for those species for which Wagner gave his

own full description, or for which he did not explicitly

state that he had not seen specimens, he had arguably

seen the actual material.

Regarding Bulimus floccosus [= Achatina floccosa

Spix], Wagner stated that “Musewm Monacense possidet

specimen unicum, cujus apex abruptus est; itaque nu-

merum anfractorum et longitudinem totius testae non in-

dicare possum” (The Munich Museum has only one

specimen, the apex of which is broken; so I cannot in-

dicate either the number of whorls or the total length

of the shell). However, Spix’s figures 3 and 4 on plate IX

do not show a broken apex; perhaps the shell was dam-

aged during its shipping or handling, before Wagner had

ie opportunity of measuring it. This also implies that

shell measurements given by \ Wagner were probably not

taken from the lithographs but from the actual shells.

In some instances Wagner made unambiguous state-

ments that could probably not have been made had he

not studied the collection. For example, on p. 31, he

said that “Anodon siliquosum et pygmaeum cl. Spixii ae-

tate sola inter se diversa esse plurimus speciminibus in-

termediis, quae in Museo Monachensi asservatur, facile

probatur’ (Anodon siliquosum and pygmaeum of the

eminent Spix only differ from each other by their age,

as can be easily demonstrated by the many intermediate

specimens conserved in the Munich Museum). On the

same page, Wagner made the following comment on An-

odon radiatus Spix: “Museum Monachense permulta spe-

cimina hujus speciei asservat, quae omnia ab Anodonte

glauco cl. Humboldtii diversa sunt” (The Munich Mu-

seum conserves a lot of specimens of this species, all of

which differ from Anodon glaucus of the eminent Hum-

boldt).

The instance of Solarium serpens Spix is also intrigu-

ing. Wagner divided the genus Helix into three sections:

a) dentatae, non carinatae (with apertural teeth, not car-

inated); b) carinatae, and c) planorbes, neque carinatae,

neque dentatae (planorbid, neither carinated, nor

toothed). He placed Helix punctata Wagner |= Solarium

serpens Spix of pl. XVII fig. 2] in section b (carinatae),

while Helix pellisserpentis “Chemnitz” [= Solarium ser-

pens Spix of pl. XVII fig. 1] was placed in section c.

However, Spix’s illustrations are of apical (Spix’s fig. 1)

and basal (Spix’s fig. 2) views, from which it is not pos-

sible to determine whether the shell was carinate or not,

suggesting that if he was not simply guessing Wagner

saw the shell(s).

So, Wagner introduced some severe errors in the de-

scription of the bivalves, which induced Ihering (1890)

to conclude that he had not seen the specimens. How-

ever, the above discussion leads us to conclude that he

did indeed work with at least the greater part of Spix’s

type material, though perhaps not all it.

TYPES

The natural history material collected by Spix and Mar-

tius in Brazil formed a major part of what was intended

THE NAUTILUS, Vol. 118, No. 2

as the “Museum Brasiliense” (Fittkau, 1983). The zoo-

logical material and the types were later integrated into

the collection of the Zoologische Staatssammlung in Mu-

nich, where most of it is still held. Fechter (1983a, b)

surveyed the molluscan types of Spix, listing 64 gastro-

pod species and 19 species of Unionoidea, collected, il-

lustrated and named by Spix, as evidenced by the labels

and the 1827 publication. However, some of the type

material is now missing, as a result of damage the mu-

seum suffered during World War II. Also, Spix’s original

labels are not all extant, and according to Fechter

(1983a) it was only possible in four cases, by comparing

the labels to letters written by Wagner, to attribute some

of the existing older labels to Wagner, apparently stem-

ming from his working in the collection years after Spix’s

death (see above).

ACKNOWLEDGMENTS

We thank the following for comments and discussion:

Gene Coan, Kevin Cummings, Neal Evenhuis, Riccardo

Giannuzzi-Savelli, Otto Kraus, Fabio Moretzsohn, Mi-

chael Ohl, Andrew Rindsberg, Barry Roth, and Michael

Schrédl. They may not all agree with our interpretations.

We especially thank Dick Petit for a detailed review of

the manuscript. We also thank Michael Schrédl and En-

rico Schwabe (ZSM), Ingeborg Kilias (Museum fiir Na-

turkunde, Humboldt-Universitit zu Berlin), and Peter

Mordan (The Natural History Museum, London) for

help with obtaining literature. For help with Latin trans-

lation of the “Praefatio”, M. G. thanks Bernhard Richer

(Ahrensburg) and N. J. C. thanks Lea Cazzaniga. N. J.

C. is a staff researcher of the Comisién de Investiga-

ciones Cientificas de la Provincia de Buenos Aires (Ar

gentina). R. H. C. thanks the USDA T-STAR program

for partial support on his work on Ampullariidae.

LITERATURE CITED

Cowie, R. H. and S. C. Thiengo. 2003. The apple snails of the

Americas (Mollusca: Gastropoda: Ampullariidae: Asolene,

Felipponea, Marisa, Pomacea, Pomella): a nomenclatural

and type catalog. Malacologia 45: 41-100.

Fechter, R. 1983a. Das Typenmaterial der von J.B. v. Spix in

Brasilien gesammelten Unionacea. Spixiana Supplement

9: 295-255.

Fechter, R. 1983b. Liste des Typenmaterials der von J.B. v.

Spix in Brasilien gesammelten Gastropoda. Spixiana Sup-

plement 9: 221-223.

Fittkau, E. J. 1983. Johann Baptist Ritter von Spix. Sein Leben

und sein wissenschaftliches Werk. Spixiana Supplement 9:

11-18.

Glaubrecht, M. 1996. Evolutionsékologie und Systematik am

Beispiel von Siib- und Bnclanassasdanedlean (Mollusca:

Caenogastropoda: Cerithioidea): Ontogenese-Stratigien,

paliontologische Befunde und historische Zoogeographie.

Backhuys Publishers, Leiden, xvi + 544 pp.

Haas, F. 1969. Superfamilie Unionacea. In: Das Tierreich. Lie-

ferung 88 (I-X). Berlin. 663 pp.

Huber, B. and W. Huber. 1993. Dr. Johann Baptist Ritter von

R. H. Cowie et al., 2004

Page 87

Spix—eine “beriihmte Miinchner persénlichkeit”. Spixi-

ana 16: 97-104.

ICZN [Intemational Commission on Zoological Nomencla-

ture]. 1999. International Code of Zoological Nomencla-

ture. Fourth edition. Intemational Trust for Zoological

Nomenclature, London, xxix + 306 pp.

Ihering, H. von. 1890. Revision der von Spix in Brasilien ges-

ammelten Najaden. Archiv fiir Naturgeschichte 56(1):

117-170.

Kottelat, M. 1988. Authorship, dates of publication, status and

types of Spix and Agassiz’s Brazilian fishes. Spixiana 11(1):

69-93.

Martius, C. F. P. von. 1862. Denkrede auf Joh. Andreas

Wagner. Miinchen. Verlag der Kéniglichen Akademie.

16 pp.

Morrison, J. P. E. 1954. The relationships of Old and New

World melanians. Proceedings of the United States Na-

tional Museum 103: 357-394, pl. 11.

THE NAUTILUS 118(2):88-92, 2004

Page 88

The genus Olivancillaria (Gastropoda: Olividae) in the Miocene

of Chile: rediscovery of a senior synonym and description of a

new species

Sven N. Nielsen

Geologisch-Palaéontologisches Institut

und Museum

Universitit Hamburg

Bundesstrasse 55

20146 Hamburg

GERMANY

[email protected]

ABSTRACT

Revision of historical collections from the Tertiary of Chile

housed in Paris and in new collections in Santiago de Chile

revealed the presence of an undescribed species of the olivid

genus Olivancillaria dOrbigny, 1840. Also, the used name of

a common Miocene species, O. tumorifera (Hupé, 1854), is

preceded by an almost completely ignored senior synonym.

The older and therefore valid name for the known species is

reintroduced as O. claneophila (Duclos, 1835), status and re-

pository of type material is given, types are redescribed and

figured and the new species O. matanzana is described. Oli-

vancillaria matanzana differs from O. claneophila in having a

higher spire, a less thick callus, and by lacking a node on the

callus.

INTRODUCTION

Only one species of the genus Olivancillaria dOrbigny,

1840, has been described from the Chilean Miocene;

that species is generally known as Oliva tumorifera

Hupé, 1854. It is a common species of moderate local

biostratigraphic potential because it occurs in almost all

Miocene localities of Chile and is easily recognized.

Thus it might help field geologists to date sediments on

a provisional basis. Subsequent workers (e.g. Philippi,

1887; Méricke, 1896) used the name O. tumorifera, and

only later Klappenbach (1966) realized that an earlier

name existed for the species. This, however, remained

unnoticed by subsequent workers (Fleming in Watters

and Fleming, 1972; Tavera, 1979). Discovery of a new

species of Olivancillaria in the Matanzas collection of V.

Covacevich and D. Frassinetti in the Museo Nacional de

Historia Natural, Santiago de Chile, initiated a deeper

interest in this genus and lead to the rediscovery of the

older name for the known species: Oliva claneophila

Duclos, 1835.

The genus Olivancillaria @Orbigny is usually referred

either to the year 1839 (e.g. Rios, 1994; Pastorino, 1995)

or 1841 (e.g. Kantor, 1991). However, Burch and Burch

(1964) showed the correct date to be 1840. Species of

Olivancillaria today live predominantly along the Atlan-

tic coast of South America but one species occurs also

in India. Fleming (1972) discussed Olivancillaria as an

Atlantic element in the Miocene fauna of Chile, but a

wide distribution in the Pacific during the Miocene is

indicated by the presence of O. altenai Beets, 1986 in

the late Miocene of East Borneo (Beets, 1986). This for-

merly wide distribution explains the disjunctive Recent

distribution in South America and India. Olivancillaria

must have migrated from Chile across or around South

America during the Miocene, but it is unknown where

it survived during the Pliocene because there is no re-

cord of post-Miocene species from Chile and, on the

other hand, there is no Neogene fossil record in Argen-

tina. The genus is also absent in the Caribbean faunas

from Neogene to Recent.

Olivancillaria claneophila has been found at almost all

Miocene localities from the Navidad Region (Duclos in

Chenu, 1846; Philippi, 1887; my own data) to Chiloé

Island (Fleming, 1972; my own data). The new species

has been found at a locality described by Frassinetti and

Covacevich (1993) who, in an earlier paper (Covacevich

and Frassinetti, 1980), also provided a preliminary list of

the mollusk species encountered there. The fossiliferous

level is situated on the tidal platform about one kilo-

meter north of Matanzas (33°57'27”" S, W 71°52'15” W)

and is normally covered by beach-sand. The sediments

range from mudstone to sandstone and yield a fauna

somewhat unusual for the region, since a number of taxa

are only known from this locality. The reason for this,

however, is still unresolved. Macrofauna and sediments

seem to indicate deposition in a near-coast environment,

while benthic foraminifera and ostracods show that dis-

placement into great depths (>1500 m) occurred (Fin-

ger et al., 2003).

S. N. Nielsen, 2004

Page 89

Estero Maitenlahue

coast north of

Rio Rapel

38°5 Arauco

Figure 1. Type locality of Olivancillaria matanzana new spe-

cies and localities of O. claneophila mentioned in the text.

The age of the Navidad Formation is still in debate.

While some authors suggested a lower Miocene (Dremel

in Herm, 1969: Tavera, 1979: Frassinetti and Covacev-

ich, 1993) others gave an upper Miocene age (Tsuchi et

al., 1990; Ibaraki, 1992) for the same locality. Both es-

timates where made with Foraminifera, the former also

on the basis of mollusks. Dating of own foraminifer ma-

terial (Finger et al., 2003) indicates a Late Miocene age

for the localities containing O. claneophila.

MATERIALS AND METHODS

Specimens described or mentioned in this study are de-

posited in the collections of the following institutions:

Museo Nacional de Historia Natural, Departamento de

Paleontologia de Invertebrados, Santiago de Chile

(SGO.PI); Muséum national d’ stoi naturelle, Labor-

atoire de Biologie des Invertébrés marins et Malacologie

(MNHN-BIMM); Laboratoire de Géologie (MNHN-

LG), Paris, France and Senckenberg Museum Frankfurt

am Main (SMF), Germany.

SYSTEMATICS

Family Olividae Latreille, 1825

Genus Olivancillaria @Orbigny, 1840

Type Species: Oliva brasiliensis Chemnitz, 1788 (=

Porphyria urceus Réding, 1798); Recent, Brazil to Ar-

gentina.

Olivancillaria claneophila (Duclos, 1835)

(Figures 2-7, 14-18)

Oliva claneophila Duclos, 1835: pl. 29, figs. 8, 9; Duclos in

Chenu, 1846: 31, pl. 31, figs. 8, 9.

Oliva tumorifera Hupé, 1854: 217-218, Conch. pl. 3, fig. 8;

Philippi, 1887: 72, pl. 8, fig. 9.

Oliva pyriformis Philippi, 1887: 73, pl. 8, fig. 11.

Oliva lebuensis Philippi, 1887: 73, pl. 8, fig.

Oliva otaeguii fa 1887: 74, pl. 8, fig. 2

Ancillaria tumorifera (Hupé, 1854) Méricke, eee 572.

Olivancillaria tumorifera (Philippi, 1887) Ihering, 1907: 514;

Tavera, 1979: 90, pl. 16, fig. 43.

Olivancillaria claneophila (Duclos, 1835) Klappenbach, 1966:

Olivancillaria (Lintricula) tumorifera (Hupé, 1854) Fleming,

1972: 398, figs. 6t, 6w.

Diagnosis: Shell elongate- oval with convex to angu-

lated sides, solid; spire low, covered by thick callus. Col-

umella with two prominent folds, upper fold splitting

into four finer folds in juveniles. Heavy parietal callus

with low, well-defined node. Suture channeled appar-

ently only on three quarters of last whorl. Aperture sub-

rectangular.

Description: The solid, elongate-oval shell has convex

to angulated sides and is characterized by its low spire

with very heavily developed callus. The callus covers the

spire whorls completely and about one whorl of callus is

visible. The columella is covered by callus, the lower part

bearing two moderately prominent folds, of which the

upper feathers into four finer ones in juveniles. The pa-

rietal-field produced by callus, bears a low, well-defined

node. The suture is channeled but due to the heavy cal-

lus only visible on the last three quarters of the body

whorl. The aperture is sub-rectangular.

Type Material: The specimen figured by Duclos is

deposited in MNHN-BIMM (Figure 5, unnumbered,

Navidad, height 35.4 mm). The specimen it is here re-

garded as holotype because there is no indication that

he had seen other material. Holotype of O. tumorifera

MNHN-LG Gg2002/75 (Figures 2-4, Topocalma, height

31.4 mm). Holotype of O. otaegui SGO.PI.541 (Figures

14-15, Curauma, height 21 mm). Holotype of O. le-

buensis SGO.PI.528 (Figure 16, Lebu, height 25 mm).

Holotype of O. pyriformis SGO.PI.532 (Figures 17-18,

locality unknown, height 33 mm).

Other Material Examined: Three specimens from

the coast north of Rio Rapel, Navidad Formation; 23

specimens from Punta Perro, Navidad Formation; five

specimens from Las Brisas, Navidad Formation; four

specimens from Matanzas, Navidad Formation; one

specimen from southern coast of Peninsula Chocoi near

Carelmapu; 29 specimens from Cucao, Chiloé Island (all

own collections). SMF 236001 (1 specimen, Punta Per-

ro), SMF 326002 (20 specimens, north of Rio Rapel),

SGO.PI.5495 (6 specimens, Punta Perro), SGO.PI.5498

(13 specimens, Estero Maitenlahue), SGO.PI.5528 (1

specimen, Rapel Norte), SGO.PI.5533 (2 specimens,

Rapel Norte), SGO.PI.5550 (1 specimen, Rapel Norte),

SGO.PI.5568 (1 specimen, Rapel Norte), SGO.PI.5577

(7 specimens, Rapel Norte).

Type Locality: The coast near Navidad. “Fossile du

Page 90

1 cm

(2 cm for 6-7)

14 15

THE NAUTILUS, Vol. 118, No. 2

Figures 2-18. Miocene Olivancillaria from Chile. 2-7. Olivancillaria claneophila. 2-4. Holotype of Oliva tumorifera Hupé,

1854, MNHN-LG Gg2002/75. 5. Holotype of Oliva claneophila Duclos, 1835, MNHN-BIMM unnumbered. 6-7. Juvenile specimen

from Rapel, SMF 326001. 8-13. Olivancillaria matanzana new species. 8-10. Holotype, SGO.PI.6008. 11. Paratype 1,

SGO.PI.6009. 12-13. Paratype 2, SGO.PI.6009. 14-18. Olivancillaria claneophila. 14-15. Holotype of Oliva otaegui Philippi,

1887, SGO.PI.541. 16. Holotype of Oliva lebuensis Philippi, 1887, SGO.PI.528. 17-18. Holotype of Oliva pyriformis Philippi,

1887, SGO.PI.532. Note that specimen in Figure 5 is uncoated while all others are coated with magnesium oxide.

Chili (. . .) il appartient au terrain tertiaire recouvrant le

granit de la Trinidad, canton de la Navidad, et ne se

rencontre que sur les escarpements des bords de la mer”

(Duclos in Chenu, 1846, p. 31).

Occurrence: Southern Peru (DeVries and Frassinetti,

2003) to Chiloé, southern Chile (Fleming, 1972; own

data); lowermost to upper Miocene.

Discussion: Of the South American species, Olivan-

cillaria claneophila most resembles O. deshayesiana

(Ducros de Saint Germain, 1857) in having convex

whorls and a low spire. It differs from that species

through the presence of a callus node on the parietal

area. Olivancillaria claneophila also resembles O. car-

cellesi Klappenbach, 1965, as figured by Pastorino

(1995). From this it differs in its broader, heavier shell,

a concave columella and a well-defined node on the pa-

rietal callus. Olivancillaria vesica (Gmelin, 1791) differs

from O. claneophila in having more convex whorls, es-

pecially in adult specimens. It also has a node on the

parietal callus, but a bigger and less defined one. Oli-

vancillaria vesica has a narrower spire, an upward-pro-

jecting inner-lip callus and the columella is angulated at

about half height, all these features cannot be observed

in O. claneophila. Olivancillaria urceus, type species of

the genus, has a conical shell with an almost flat spire

and slightly convex sides, while O. claneophila has a

more angulated shell with a low, heavily calloused spire.

The type specimens of O. lebuensis and O. pyriformis

are not well preserved but comparison with the available

material, containing specimens of a wide range of pres-

ervation quality, confirms the synonymy. The prominent

groove on both specimens resembles a pseudolivid

groove (see Vermeij, 1998; Nielsen and Frassinetti,

2003); that groove, however, seems to represent a pres-

ervation artifact.

Olivancillaria matanzana new species

(Figures 8-13)

Diagnosis: Shell elliptic with convex to angulated

sides; spire moderately high, covered by callus. Colu-

mella with prominent fold, above it a second one, split-

ting into four finer folds in juveniles. Strong parietal cal-

S. N. Nielsen, 2004

Page 91

lus. Suture about three quarters of the last whorl chan-

neled.

Description: The elliptic shell has convex sides and is

characterized by its moderately high spire with well-de-

veloped callus covering the spire whorls. The columella

is covered by callus, the lower part bears two moderately

prominent folds, of which the upper splits into four finer

ones in juveniles. The suture is channeled but due to

the heavy callus only visible on the last three quarters

of the body whorl.

Type Material: Holotype SGO.PI.6008 (Figures 8-

10, Matanzas, height 40.9 mm), four paratypes

SGO.PI.6009 (Figures 11-13, Matanzas).

Other Material Examined: Further ca. 20 specimens

(unnumbered) present in the Matanzas collection of

Frassinetti and Covacevich (SGO.PI).

Type Locality: Intertidal platform about 1 km north

of Matanzas, Navidad Formation, Central Chile.

Occurrence: Only known from type locality, upper

Miocene.

Etymology: After the type locality near the village of

Matanzas.

Discussion: Olivancillaria matanzana has a higher

spire and a thinner callus on it than O. claneophila. It

also lacks a prominent node on the parietal callus. No

intermediate forms have been found. Both species occur

together at the Matanzas locality, but O. matanzana is

known from nowhere else. Olivancillaria matanzana re-

sembles O. carcellesi even more than O. claneophila, but

O. carcellesi has a high aperture, almost as high as the

shell, while O. claneophila and O. matanzana have lower

apertures. Olivancillaria vesica has a narrower and lower

spire than O. matanzana.

ACKNOWLEDGMENTS

Thanks are due to Alan Beu (Lower Hutt, New Zealand)

for helpful discussions, Dick Kilburn (Pietermaritzburg,

South Africa) for taking the photograph of O. claneo-

phila, and Eva Vinx (Hamburg, Germany) for taking

some of the other photographs. I am grateful to Agnés

Rage (MNHN, Laboratoire de Paléontologie) for pro-

viding facilities and to Virginie Héros (MNHN-BIMM),

Emmanuelle Vennin (MNHN-LG), and Daniel Frassi-

netti (SGO.PI) for loan of or access to material. Reviews

by two anonymous reviewers improved this work. This

work has been made possible by financial support of the

Deutsche Forschungsgemeinschaft (DFG) grant Ba 675/

25, the University of Hamburg, and a COLPARSYST

grant to review the collections of the MNHN.

LITERATURE CITED

Beets, C. 1986. Molluscan fauna of the Lower Gelingseh Beds

s. str., Sangkulirang area, Kalimantan timur (East Borneo).

Scripta Geologica 82: 1-82.

Burch, J. Q. and R. L. Burch. 1964. The genus Agaronia J. E.

Gray, 1839. The Nautilus 77: 110-112.

Covacevich, V. and D. Frassinetti. 1980. El género Ficus en el

Mioceno de Chile Central con descripcién de F. gayana

sp. nov. Gastropoda: Ficidae. Boletin del Museo Nacional

de Historia Natural de Chile 37: 281-294.

DeVries, T. J. and D. Frassinetti. 2003. Range extensions and

biogeographic implications of Chilean Neogene mollusks

found in Peru. Boletin del Museo Nacional de Historia

Natural de Chile 52: 119-135.

Duclos, P. L. 1835. Histoire naturelle génerale et particuliére

de tous les genres de coquilles univalves marines a l’etat

vivant et fossile, publiée par monographies; ou description

et clasification méthodique de toutes les espéces connues

jusque a ce jour, representées en couleur avec la figure et

lanatomie d'un afses grand nombre de mollusques nou-

vellement decouverts. Firmin Didot Fréres, Paris. Genre

Oliva.

Duclos, P. L. 1846. G. Oliva. Oliva Lamarck, In: J.-C. Chenu,

Illustrations Conchyliologiques. III. Univalves Marins

2(1), 31 pp., 36 pls.

Finger, K., A. Encinas, S. Nielsen and D. Peterson. 2003. Mi-

crofaunal indications of late Miocene deep-water basins

off the central coast of Chile. 10° Congreso Geoldgico

Chileno. Concepcién, Chile. Abstract Volume CD-ROM,

8 pp.

Fleming, C.A. 1972. Part II. Pliocene marine invertebrates

from Chepu district, Chiloe. In: W. A. Watters and C. A.

Fleming. Contributions to the geology and palaeontology

of Chiloe Island, southem Chile. Philosophical Transac-

tions of the Royal Society of London B 263: 380-408.

Frassinetti, D. and V. Covacevich. 1993. Bivalvos del Mioceno

de Matanzas (Formacién Navidad, Chile Central). Boletin

del Museo Nacional de Historia Natural de Chile 44: 73-

97.

Herm, D. 1969. Marines Plioziin und Pleistoziin in Nord- und

Mittel-Chile unter besonderer Beriicksichtigung der En-

twicklung der Mollusken-Faunen. Zitteliana 2, 159 pp.

Hupé, H. 1854. Malacologia y conquiliologia. In Gay, C. (ed.),

Historia fisica y politica de Chile. Vol. 8 and Atlas (Zool-

6gica). Maulde et Renou, Paris, 449 pp.

Ibaraki, M. 1992. Planktonic foraminifera from the Navidad

Formation, Chile: their geologic age and paleoceano-

graphic implications. In: Ishizaki, K. and T. Saito (eds.),

Centenary of Japanese Micropaleontology. Terra Scientific

Publishing Company, Tokyo, pp. 91-95.

Ihering, H. von 1907. Les mollusques fossiles du Tertiaire et

du Crétacé Supérieur de Argentine. Anales del Museo

Nacional de Buenos Aires 3/7: 1-611.

Kantor, Y. I. 1991. On the morphology and relationships of

some oliviform gastropods. Ruthenica 1: 17-52.

Klappenbach, M. A. 1966. Olivancillaria vesica (Gmelin, 1791)

has priority over Olivancillaria auricularia (Lamarck,

1810). Archiv fiir Molluskenkunde 95: 75-77.

Moricke, W. 1896. Versteinerungen der Tertiirformationen von

Chile. Neues Jahrbuch fiir Mineralogie, Geologie und Pa-

laontologie Beilage Band 10: 548-612.

Nielsen, S. N. and D. Frassinetti. 2003. New and little known

species of Pseudolividae (Gastropoda) from the Tertiary

of Chile. The Nautilus 117: 91-96.

Pastorino, G. 1995 (1994). Moluscos costeros recientes de

Puerto Pirémide, Chubut, Argentina. Academia Nacional

de Ciencias de Cérdoba, Miscelénea 93, 30 pp.

Page 92

THE NAUTILUS, Vol. 118, No. 2

Philippi, R. A. 1887. Die tertiiren und quartiren Versteine-

rungen Chiles. F.A. Brockhaus, Leipzig, 266 pp.

Rios, E. C. 1994. Seashells of Brazil. 2nd edition. Fundagao

Universidade do Rio Grande, Rio Grande, 492 pp.

Tavera, J. 1979. Estratigrafia y paleontologia de la Formacién

Navidad, Provincia de Colchagua, Chile (Lat. 30°50’—

34°S). Boletin del Museo Nacional de Historia Natural de

Chile 36, 176 pp.

Tsuchi, R., T. Shuto, T. Takayama, I. Koizumi, A. Fujiyoshi, R.

Nomura, M. Ibaraki, H. Duque-C., R. Tirado-S., M. Al

dana-A., E. Villavicencio-R. and R. Martinez-P. 1990.

Trans-Pacific correlation of Neogene geologic events. Re-

ports of Andean Studies, Shizuhoka University, Special

Vol. 3: 1-7.

Vermeij, G. J. 1998. Generic revision of the neogastropod fam-

ily Pseudolividae. The Nautilus 111: 53-84,

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Volume 118, Number 3

October 4, 2004

ISSN 0028-1344

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CONTENTS

INN gC) 8 i ales.

Volume 118, Number 3

October 04, 2004

ISSN 0028-1344

M. G. Harasewych

M. Gonzalez-Wangiiemert

A. Pérez-Ruzafa

M. J. Rosique

A. Ortiz

Diego G. Zelaya

Carlos S. Gallardo

Cristian Manque

Marcela Filtin

Elizabeth C. Davis

Kathryn E. Perez

Daniel J. Bennett

Gongalo Calado

New Columbariinae (Gastropoda: Turbinellidae) from the

Inrange Ocean tee lra tye ae eee tae reece pre res rea pe MENU ie ccc br OA ee 93

Genetic differentiation in two cryptic species of Ostreidae,

Ostrea edulis (Linnaeus, 1758) and Ostreola stentina

(Payraudeau, 1826) in Mar Menor Lagoon, southwestern

IMeditername amis Gabry toke Le latent yee eR ene ie lastest. cit cet h sneer apes 103

The genus Margarella Thiele, 1893 (Gastropoda:

Trochidae) in the southwestern Atlantic Ocean...................0.005. 112

Comparative resistance to starvation among early juveniles

Of GONE MMMM maMACGONCOaM SMEG ...5c0ccccc0cccccsscconcnovosoveves 121

Euglandina rosea (Férussac, 1821) is found on the ground

AIT MI TIe ELE S SOIT EG] OL GL alesse ee Reema ea ee tes nine ones eet Ay ae IQ7

Rediscovery of the syntypes of Doriopsilla pelseneeri

ID) <@liveimawealtS Oey eek ee Ms cece ere sm Aa oases cea cwr ee ese hacker ye cae 129

THE NAUTILUS 118(3):93-102, 2004

Page 93

New Columbariinae (Gastropoda: Turbinellidae) from the

Indian Ocean

M. G. Harasewych

Department of Zoology, MRC 163

National Museum of Natural History

Smithsonian Institution

PO Box 37012

Washington, DC 20013-7012 USA

[email protected]

ABSTRACT

Four new species of Columbariinae are described from the

Indian Ocean, based on museum material. Coluzea kallistropha

inhabits the lower continental slope off Mozambique and the

KwaZulu-Natal coast of South Africa, and may easily be distin-

guished from its congeners by its lower spire and broad, flange-

like peripheral keel. Coluzea madagascarensis occurs on the

upper continental slope along the southwestern coast of Mad-

agascar. Diagnostic features include long, radial spines along

the shell periphery and strong spiral sculpture. Columbarium

quadrativaricosum is presently known only from the upper

continental slope off Transkei, South Africa. Its distinctive,

thick, squarish varices serve to distinguish it from closely re-

lated species. Coluzea naxa is known only from its type locality

along the uppermost continental slope off the Northwest Shelf

of Weston Australia. It is readily discerned from its geograph-

ically proximal congeners on the basis of its smaller, thinner,

shell, prominent rounded axial sculpture, and posteriorly di-

rected spines.

INTRODUCTION

The subfamily Columbariinae represents an early [Maes-

trichtian (Darragh, 1969)] and diverse radiation of ver-

mivorous Turbinellidae that has, since the Neogene,

been restricted to habitats ranging from outer continen-

tal shelf to abyssal depths along continental margins. The

subfamily is currently known from 52 Recent species

and subspecies, with greatest diversity occurring along

the western margins of ocean basins [western Atlantic

Ocean = 11 species; western Indian Ocean = 14 species

(3 new herein); western Pacific = 17 species and sub-

species]. Six species (1 new herein) are presently known

from the eastern Indian Ocean, and single species have

been described from the eastern Pacific, the Antarctic,

and the Bering Sea. The taxa described herein are based

on specimens collected by the research vessels ANTON

Bruun, MAscAREIGNES III, Merrinc Naup#, and Lapy

BasTEN and housed in the collections of the National

Museum of Natural History, Smithsonian Institution

(USNM), the Museum national d'Histoire naturelle,

Paris (MNHN), the Natal Museum, Pietermaritzburg

(NM), and the Western Australian Museum (WAM).

While the increased sampling of soft bottom deep-sea

habitats is likely to continue to bring additional colum-

bariine taxa to light, the known Ewnoa is beginning to

provide insights “a patterns of allopatric speciation and

bathymetric zonation, as well as to common ecopheno-

typic adaptations in different oceans.

SYSTEMATICS

Family Turbinellidae Swainson, 1840

Subfamily Columbariinae Tomlin, 1928

Genus Coluzea Allan, 1926

Coluzea kallistropha new species

(Figures 1-7)

Diagnosis: A moderate sized species with a narrow,

fusiform, white shell with a broad, laterally oriented,

flange-like keel along the shell periphery with an un-

dulating edge and spiral threads along both surfaces. A

weak but distinct anterior carina is present. Shoulder

sloping. Spiral sculpture consists of 1-2 strong, rectan-

gular, widely spaced spiral cords between the suture and

the periphery, 1-3 cords between periphery and siphon-

al canal, and 5-8 along the proximal portion of the long,

axial siphonal canal. Inner lip formed of a smooth glaze

deposited after the outer shell layer of the previous

whorl is resorbed.

Description: Shell (Figures 1, 3, 6) of moderate size

(to 52 mm), thin to moderately heavy, fusiform; spire

angle 35—-43°. Protoconch (Figure 4) eroded, estimated

to consist of 1% inflated whorl, first whorl deflected from

coiling axis of shell by 75-85°. Transition to teleoconch

indistinct, eroded. Teleoconch extrapolated to consist of

8 strongly convex nearly triangular whorls. Suture abut-

ted to previous whorl just anterior to strong cord form-

ing anterior carina (Figure 1, ac). Earliest discernible

sculpture consists of strong axial ribs (10 per whorl on

Page 94 THE NAUTILUS, Vol. 118, No. 3

Figures 1-6. Coluzea kallistropha new species. 1-2. Holotype [USNM 718517], 60 mi NE Ponta Sao Sebastiaéo, Mozambique,

21°18’ S, 36°18’ E, in 1510-1600 m. 1. Apertural, right lateral, dorsal, and apical views of shell. 2. Outer and inner surfaces of

operculum. 3-5. Paratype 1 [USNM 717921] 100 mi SE Lourenco Marques, Mozambique, 27°09’ S, 34°09’ E, in 1335 m. 3.

Apertural, right lateral, and dorsal views of shell. 4. Lateral view of protoconch and early whorls. 5. Periostracum between suture

and shell periphery. 6. Paratype 2 [NM A76] Durban Bay, from dredgings dumped at head of bay. Seale bar (1 cm) applies to all

entire shells. Abbreviations: ac, anterior carina, c, spiral cord.

M. G. Harasewych, 2004

20°E 30°E 40°E 50°E

40°S

Figure 7. Geographic distributions of new species of Colu-

zea and Columbarium from the western Indian Ocean. Coluzea

kallistropha new species, open circle = type locality, solid cir-

cles = additional localities. Coluzea madagascarensis new spe-

cies, open diamond = type locality, solid diamonds = addi-

tional localities. Columbarium quadrativaricosum new species,

open triangle = type locality, solid triangles = additional lo-

calities.

second teleoconch whorl) crossed by strong spiral cords

(3 above and 3 below periphery). By third teleoconch

whorl, short, broad, open, laterally directed spines form

along periphery. By fourth teleoconch whorl, axial ribs

diminish in prominence; spines fuse to form continuous

peripheral keel with 3 weaker spiral cords above, and

two below the keel. By fifth teleoconch whorl, axial

sculpture becomes indistinct except for fine growth

lines, keel increases in size. Spiral sculpture becomes

more pronounced, with 1-2 strong hollow cords be-

tween suture and periphery, 1—3 between periphery and

siphonal canal, 5-8 along proximal portion of siphonal

canal, decreasing in prominence distally. Finer spiral

threads (1-3) between adjacent cords, 7-12 undulating

threads along adapical and abapical surfaces of periph-

eral keel. Aperture broadly ovate, roughly triangular,

sharply tapering anteriorly, deflected from shell axis by

23-26°. Outer lip glazed, furrows beneath spiral cords,

deepest beneath peripheral keel. Inner lip smooth, outer

shell layer comprising surface sculpture resorbed along

parietal region, columella and siphonal canal prior to de-

position of thin porcellaneous glaze. Siphonal canal long,

axial, stout, straight. Shell color uniformly white. Perios-

tracum (Figure 5) thin, straw-colored, lamellose and

finely hirsute, forming low, broad tufts along spiral cords

and threads (Figure 5, c). Operculum (Figure 2) thin,

claw-shaped, broadly rounded posteriorly, sharply taper-

ing anteriorly, leading to a terminal nucleus. Inner sur-

face with thin glaze covering anterior region and nucle-

us, defining rounded attachment region. Soft tissues,

radula unknown.

Type Locality: 60 mi NE Ponta Sao Sebastiaio, Mo-

Page 95

zambique, 21°18’ S, 36°18’ E, in 1510-1600 m, Gulf of

Mexico Shrimp Trawl, Anton Bruun Cruise 8, sta.

399C, 2 October 1964.

Type Material: Holotype, USNM 718517, 51.35 mm,

from the type locality; Paratype 1, USNM 717921, 26.60

mm, 100 mi SE Lourenco Marques, Mozambique,

27°09' S, 34°09’ E, in 1335 m, ANTON BRUUN Cruise 7,

sta. 374C, 23 August 1964; Paratype 2, NM A76, 42.22

mm, Durban Bay, from dredgings dumped at head of

bay, 2 December 1972.

Distribution (Figure 7): The presently known range

of this species extends from southern Mozambique

southward to KwaZulu-Natal, South Africa, at depths of

1335 to 1600 m.

Etymology: Kallistropha = kallimos—Greek, beauti-

ful + strophe—Greek, a turning, twist.

Remarks: Coluzea kallistropha is most closely related

to Coluzea eastwoodae Kilburn, 1971, and Coluzea juliae

Harasewych, 1989, but may easily be distinguished from

both by its broad, flange-like peripheral keel, and by the

sculpture between the suture and peripheral keel

(smooth in C. eastwoodae; 2-3 broad, squarish hollow

cords in C. juliae; 1-2 strong, narrow cords in C. kallis-

tropha). These three species inhabit the same geograph-

ic area off the southeastern coast of Africa, but have

different bathymetric ranges. Coluzea eastwoodae, the

most widely sampled species, has been reported from

depths ranging from 150 to 740 m, with the deepest live

taken specimen presently known being from 570 m.

Based on the few specimens of C. juliae to be sampled

with precise depth data, the confirmed bathymetric

range for this species is 600 to 700 m. The two live

collected types of C. kallistropha were collected along

the lower continental slope, at depths of 1335 to 1600

m, among the deepest for any species of Columbariinae.

While most living Columbariinae occur on sand and mud

bottoms along the outer continental shelf and upper

continental slope, two species of Fulgurofusus, F. ben-

thocallis (Melvill and Standen, 1907) and F. aequilonius

Sysoev, 2000, are known to occur on the abyssal plain in

polar regions, the former off the South Orkney Islands,

the latter in the Bering Sea.

It is interesting to note that a phenotype with a broad,

laterally directed, flange-like keel occurs in most regions

with a high diversity of columbariines (e.g., Fulgurofusus

brayi Clench, 1959—western Atlantic; Coluzea kallistro-

pha—western Indian Ocean; Coluzea altocanallis (Dell,

1956)—New Zealand; Columbarium pagodoides (Wat-

son, 1882)—eastern Australia), suggesting that the mor-

phology may be an ecophenotypic response to a partic-

ular substrate or habitat.

Coluzea madagascarensis new species

(Figures 7-17)

Diagnosis: A large species, with heavy, narrow, fusi-

form shell, prominent and persistent strongly shouldered

THE NAUTILUS, Vol. 118, No. :

Figures 8-16. Coluzea madagascarensis new species. 8-9. Holotype [MNHN], off SW Madagascz 8’ S, 43°05’ E, in 450-

500 m. 8. Apertural, right lateral, and dorsal views of shell. 9. Outer and inner surfaces of operculum. 10-11. Par atype 1 [USNM

1018: , off SW Madagascar, 22°17’ S, 43°04’ E, in 425-450 m. 10. Ape aoe and dorsal views of shell. 11. Periostracum. 12.

Paratype 2 [MNHN], off SW Madagascar, 22°16’ S, 43°06’ E, in 360-4 . 13. Par: ees 9. 14. atype 10, both [

SW Madagascar, 23°36.4’ S, 43°31.1’ E, in ‘450—46( m. 15-16. Views of the eed of Paratype 6. [MNHN], off SW Madagascar,

22°14.8' S, 43°04.7' E, in 450 m. Scale bar (2 cm) applies to all entire shells.

M. G. Harasewych, 2004

Figure 17. Coluzea madagascarensis new species. Left lat-

eral and dorsal views of radular teeth at mid-length of radular

ribbon.

axial ribs [9-10 on body whorl], each with a long, axially

oriented, open spine along the periphery. Anterior carina

weakly developed. Shoulder sloping, with 5-8 distinct

cords between suture and periphery, 2-3 between pe-

riphery and weak anterior carina, 5-6 between carina

and base of siphonal canal, 25-30 along proximal % of

long, axial siphonal canal. Inner lip of smooth glaze de-

posited after outer shell layer of previous whorl re-

sorbed.

Description: Shell (Figures 8, 10, 12-14) large (to 78

mm), moderately heavy, strongly fusiform; spire angle

28—34°. Protoconch (Figures 15-16) of about 14—1%

smooth, glassy, inflated whorls, first whorl smaller than

second, deflected from shell axis by about 80°. Transition

to teleoconch indistinct, marked by formation of a pe-

ripheral keel, followed within % whorl by axial ribs, then

by spiral cords above and below periphery. Teleoconch

of up to 10% evenly convex whorls with strong periph-

eral carina bearing tubercles on early whorls that change

to long, open, radially oriented spines by sixth postnu-

clear whorl. Suture abutted to previous whorl just an-

terior to dominant spiral cord that forms weak anterior

carina (Figure 8, ac). Axial sculpture begins within %

whorl of transition to teleoconch, consists of 8—10

strongly shouldered ribs that extend from suture to su-

ture on first teleoconch whorl. Axial ribs overlaid by

strong, evenly rounded spiral cords, 3-4 above, 2—3 be-

low periphery. By sixth teleoconch whorl, short, open,

spines perpendicular to coiling axis appear along periph-

ery, supporting axial ribs become broader, more promi-

nent near periphery, less so near sutures. As shoulder

spines become longer, more pronounced in subsequent

whorls [9-10 on body whorl], axial ribs become less dis-

cernible. Spiral cords increase in number and promi-

nence with increasing whorl number (body whorl: 5-8

between suture and periphery, 2-3 between periphery

and weak anterior carina, 5-6 between carina and base

of siphonal canal, 25-30 along proximal % of siphonal

canal, becoming weaker, less distinct distally). Aperture

broadly ovate, rounded abaxially, tapering anteriorly, de-

flected from coiling axis by 26-29°. Outer lip thin. Fur-

row beneath periphery deepest and widest beneath

Page 97

spines. Second, weaker furrow beneath anterior carina.

Inner lip smooth, sculptural elements on outer shell lay-

er resorbed prior to deposition of thin porcellaneous

glaze. Siphonal canal long (~0.46 shell length), axial,

stout, weakly twisted along distal half. Shell color uni-

formly white. Periostracum (Figure 11) moderately

thick, yellowish brown, of closely spaced lamellae, weak-

ly hirsute along spiral cords (evident in early whorls).

Operculum (Figure 9) moderately thin, rounded, taper-

ing to terminal nucleus. Inner surface with rounded

glaze along nucleus, surrounding attachment area.

Holotype a mature, poorly preserved, female, with

mantle cavity spanning ~%4 whorl, kidney, upper whorls

not recovered. Animal light tan colored. Foot, small,

rectangular. Retracted tentacles, short, conical; eyes ab-

sent. Disposition of mantle cavity and cephalic haemo-

coel organs similar to Coluzea aapta (see Harasewych,

1986:161). Proboscis, long, convoluted within proboscis

sheath. Radula (Figure 17) short (3.3 mm), narrow (153

wm), with 116 rows of teeth. Rachidian teeth with 3

short, stout cusps spanning anteriorly indented middle

half of basal plate that broadens laterally.

Type Locality: SW Madagascar, 22°18’ S, 43°05’ E,

in 450-500 m, Chalutier “MascargIcngs III”, sta. 33,

20 January 1986.

Type Material: Holotype, MNHN, 77.50 mm, from

the type locality; Paratype 1, USNM 1018395, 65.53

mm, SW Madagascar, 22°17’ S, 43°04’ E, in 425-450 m,

Chalutier “MascaREIGNES III”, sta. 6, 21 December

1985; Paratype 2, MNHN, 67.50 mm, SW Madagascar,

92°16' S, 43°06’ E, in 360-415 m, Chalutier “Mascar-

EIGNES III”, sta. 41, 22 January 1986; Paratype 3,

MNHN, 78.32 mm, SW Madagascar, 22°14.7’ S,

43°04.5' E, in 470-475 m, Chalutage 114, 2 December

1973; Paratype 4, MNHN, 72.72 mm, SW Madagascar,

22°18’ S, 43°05’ E, in 425 m, Chalutier “MascaREIGNES

IIT’, sta. 13, 23 December 1985; Paratype 5, MNHN,

54.87 mm, SW Madagascar, 22°21.6’ S, 43°04.3’ E, in

450 m, Chalutage 95, 27 November 1973; Paratype 6,

MNHN, 69.62 mm, SW Madagascar, 22°14.8’ S,

43°04.7' E, in 450 m, Chalutage 115, 2 December 1973;

Paratypes 7, 8, MNHN, 58.17 mm, 58.89 mm, SW Mad-

agascar, 22°17.9’ S, 43°04’ E, in 450 m, Chalutage 105,

29 November 1973; Paratypes 9, 10, MNHN, 63.11 mm,

74.05 mm, SW Madagascar, 23°36.4’ S, 43°31.1’ E, in

450-460 m, Chalutage 66, 29 February 1973; Paratype

11, Ritter collection, trawled off Tuléar, Madagascar, in

600-799 m, dead collected, June 2002.

Distribution (Figure 7): Coluzea madagascarensis

new species occurs off the southwestern coast of Mad-

agascar. The confirmed bathymetric range is 415-470 m,

although dead-collected specimens have been taken at

depths between 600 and 799 m.

Etymology: madagascarensis = Madagascar + en-

sis—Latin, belonging to.

Remarks: Coluzea madagascarensis is intermediate in

Page 95

THE NAUTILUS, Vol. 118, No. 3

morphology between C. eastwoodae from off southeast-

em Africa and C. distephanotis (Melvill, 1891) from off

northwestem Australia, all occurring at comparable

depths. It may be distinguished from C. eastwoodae by

its more elongate, fusiform shell, spiral sculpture that is

stronger, more uniform in size and more uniformly dis-

rralbmnierd (C. eastwoodae lacks spiral sculpture between

suture and periphery), and by having fewer, longer, ax-

ially oriented spines along the periphery. Coluzea mad-

agascarensis differs from C. distephanotis in having a

ieee tabulate shoulder, stronger spiral sculpture between

suture and periphery, and spines that are axially rather

than posteriorly directed along the periphery. Both C.

eastwoodae and C. distephanotis have a prominent an-

terior carina, which is barely distinguishable in C. mad-

agascarensis.

Columbarium quadrativaricosum new species

(Figures 7, 18-23)

Diagnosis: A small species with a gradate spire, fusi-

form, white shell, with tabulate, squarish whorls, axial

sculpture of thick, rounded varices [8-9 per whorl] and

spiral sculpture of distinct cords, square in profile, stron-

gest on periphery and anterior carina. Inner lip formed

by a thickly glazed peristomal plate that overlies the col-

umella and proximal siphonal canal.

Description: Shell (Figures 18, 22, 23) small (to 45

mm), thick to moderately thin, Fneilforarn, with squarish

varices. Spire angle 46-51°. Protoconch (Figure 20) of

about 14 smooth, glassy whorls. First whorl inflated, de-

flected from coiling axis by 67—72°. Transition to teleo-

conch distinguished by onset of peripheral keel, rounded

at first, becoming keel-like, with broad, undulating axial

nodes within % whorl. Teleoconch of up to 7 strongly

shouldered, nearly tabulate, squarish whorls. Suture ad-

pressed to previous whorl anterior to anterior carina.

Earliest sculpture, apart from very fine growth striae vis-

ible on all teleoconch whorls, consists of broad, rounded

axial nodes along peripheral keel (8-9 per whorl) that

become narrower, more sharply defined, supported by

axial ribs of increasing prominence by third teleoconch

whorl. By fourth teleoconch whorl, axial ribs form broad,

solid varices, thickest between peripheral keel and an-

terior carina, producing a square whorl profile, with

nearly tabulate shoulder. Spiral sculpture first appears

on second teleoconch whorl, as single cord below pe-

ripheral keel. By third teleoconch whorl, 2-3 weak cords

or threads appear above peripheral keel. Body whorl

with 4—5 strong, square, equally spaced cords between

suture, peripheral keel; 1 strong cord between major

cords along peripheral keel, anterior carina; 4-5 cords

between anterior carina, siphonal canal; 7-10 cords on

proximal % of siphonal canal. Finer threads between ad-

jacent cords, suture to keel (0 threads); keel to carina

(1-3 threads); carina to siphonal canal (1-2 threads), si-

phonal canal (0-1 threads). Aperture broadly ovate,

nearly rounded, tapering anteriorly, deflected from shell

axis by 23-29°. Outer lip thickly glazed, with furrows

beneath suture, peripheral keel, anterior carina extend-

ing to nearest varix. Inner lip smooth, thickly glazed,

peristomal plate overlaying columella, proximal portion

of siphonal canal. Siphonal canal long, axial, stout,

straight. Shell color uniformly white. Periostracum (Fig-

ure 21) thick, amber colored, lamellose. Operculum

(Figure 19) thin, ovate, broadly rounded posteriorly,

with terminal nucleus. Inner surface with rounded at-

tachment area. Soft tissues, radula, unknown.

Type Locality: Off Mendu Point, Transkei, South AF

rica, 32°21.8’ S, 29°00.0’ E, in 300 m, on coarse sand,

R/V Merrinc NaubE, sta. R 10, 12 July 1984.

Type Material: Holotype, NM C 6279, 36.87 mm,

from the type locality; Paratype 1, USNM 1018396,

39.79 mm, off Qora River, Transkei Region, Eastern

Cape, South Africa, 33°33.6’ S, 28°48.8’ E, in 300 m,

coarse sand, some broken shell, R/V Merrinc Naupb#£,

sta. U 10, 11 July 1984; Paratype 2, NM C 1802, 28.67

mm, off Bulungula River, Transkei Region, Eastern

Cape, South Africa, 32°13.7' S, 29°08.7' E, in 250-270

m, muddy sand, old shell debris, R/V MerriNc NAUDE,

sta. 17 July 1982; Paratype 3, NM C 6377, 27.52 mm,

off Mendu Point, Transkei Region, Eastern Cape, South

Africa, 32°24.0’ S, 28°59.0’ E, in 250 m, coarse sand,

rubble, few sponges, R/V Metrinc Naupgr, sta. R11, 12

July, 1984; Paratypes 4, 5, NM C4911, 44.77 mm, 35.81

mm, off Mendu Point, Transkei Region, Eastern Cape,

South Africa, 32°22.6’ S, 29°00.4’ E, in 250-260 m,

dredged on coarse sand; R/V MEIRING NauDE, sta. R 7,

§ June 1983; Paratype 6, NM C9337, 26.75 mm, off

Bulungula River, Transkei Region, Eastern Cape, South

Africa, 32°14.0’ S, 9°08.6’ E, in 250-300 m, dredged on

coarse sand, R/V MeErtriInc Naupe, sta. O10, 5 June

1985; Paratype 7, NM C6304, 20.02 mm, off Qora River,

Transkei Region, Eastern Cape, South Africa, 33°34.2’

S, 28°48.1' E, in 270 m, dredged on old shell bottom,

R/V Merrinc Naupé, sta. U 11, 11 July 1984; Paratype

8, NM C6573, 28.40 mm, off Qolora River, Transkei Re-

Bee Eastern Cape, South Africa, 32°47.6’ S, 28°36.6'

E, in 510 m, dredged on sandy mud, R/V MErRING

NaAupbE, sta. Y 12, 14 July 1984: Paratype 9, NM C6468,

16.46 mm, off Shixini Point, Transkei Region, Eastern

Cape, South Africa, 32°31.4’ S, 28°52.5' E, in 400-420

m, dredged on coarse sand, fine shell rubble, R/V Meir-

ING NaAupk, sta. T 16, 12 July 1984; Paratypes 10, 11,

NM C8668, 38.33 mm, 29.35 mm, off Nthlonyane River,

Transkei Region, Eastern Cape, South Africa, 32°18.2'

S, 20°06.2' E in 550 m, dredged on sand, stones, broken

Dendrophyllia, B/V Metrinc Naupg, sta. P 13, 5 July

1985; Paratype 12, NM C8946, 33.39 mm, off Mgazi

River, Transkei Region, Eastern Cape, South Aree

31°44.3' S, 29°32. or E, in 250 m, dredged on muddy

sand, R/V Mrtrinc Naupk, sta. J 11, 4 July 1985.

Distribution (Figure 7): Columbarium quadrativar-

icosum has been collected at multiple stations, all in a

narrow range [31°34'—33°34' S] off the Transkei Region,

M. G. Harasewyc

Figures 18-23. Columbarium quadrativaricosum new species. 18-21. Holotype [NM C 6279], off Mendu Point, Transkei South

Africa, 32°21.8’ S, 29°00.0’ E, in 300 m. 18. Apertural, right lateral, and dorsal views of shell. 19. Outer and inner surfaces of

operculum. 20. Lateral view of the protoconch. 21. Periostracum. 22. Paratype 6 [NM C9337], off Bulungula River, Transkei,

South Africa, 32°14.0’ S, 9°08.6' E, in 250-300 m. 23. Paratype 1 [USNM 1018396], off Qora River, Transkei, South Africa, 3

S, 28°48.8' E, in 300 m. Scale bar (1 cm) applies to all entire shells. Abbreviation: ac, anterior carina.

Page 100

Eastern Cape, South Africa, at depths ranging from 250

to 550 m. Specimens from depths in excess of 420 m

were all dead collected and extremely worn.

Etymology: quadrativaricosum = quadratus—Latin,

squared + varicosus—Latin, varicose or ridged.

Remarks: Columbarium quadrativaricosum is related

to the group of southern African congeners character-

ized by the presence of thick, rounded varices, including

C. subcontractum (Sowerby, 1902), C. formosissimum

Tomlin, 1928, and C. natalense Tornalten: 1928. It most

closely resembles C. subcontractum, which inhabits a

similar bathymetric range, but appears to be allopatric,

based on limited aeons that indicate a range to the

north of Durban Bay. While these two species reach

comparable shell length, C. quadrativaricosum has a

thinner shell, a proportionally shorter, more gradate

spire, a smaller, rounder aperture, a longer, laine si-

phonal canal, a tabulate shoulder, and varices that are

squarish rather than triangular. It also closely resembles

C. natalense, which has a comparable geographic range,

but inhabits shallower depths [90-160 m]. Columbarium

natalense can be distinguished from C. quadrativaricos-

um based on its pigmented shell (reddish brown with

lighter spiral cords and distal portion of siphonal canal),

spiral sculpture of cords that are not continuous, but

posteriorly recurved along each varix, the presence of a

long, open spine at the shoulder of each varix, and an

extremely pronounced cord along the anterior carina.

The much larger C. formosissimum, which has a more

southerly distaibution [Cape St. Blaize to Port Alfred]

and shallower bathymetric range [121-165 m], may also

be distinguished by it heavier - proportions more similar

to C. subcontractum, its reduced or absent spiral sculp-

ture, and its weaker varices that develop at a larger shell

size.

Coluzea naxa new species

(Figures 24-30)

Diagnosis: A small species with thin, narrowly fusi-

form, white shell with gradate spire. Sculpture of strong

axial ribs and open, recurved, posteriorly directed spines

along shoulder. Spiral sculpture of strong cords that

overlay axial ribs, 3 between suture and periphery, 2 be-

tween periphery and anterior carina, 4 between anterior

carina and siphonal canal, and 12-15 along proximal %

of siphonal canal. Inner lip of thin glaze deposited after

columellar surface of previous whorl resorbed.

Description: Shell (Figures 24, 29) small (to 52 mm),

thin, narrowly fusiform; spire angle 33-34%2°. Proto-

conch (Figure 26) of about 14% worn bulbous whorls,

first whorl Fate Gtecttad from coiling axis of shell by 80—90°.

Transition to teleoconch marked by onset of weak keel

along shoulder, forming axial ribs and flanked by spiral

ears above and Ihallowe within first teleoconch whorl.

Teleoconch of 9 % convex, strongly shouldered whorls.

Suture abutted to previous whorl just anterior to anterior

THE NAUTILUS, Vol. 118, No. 3

carina (Figure 24, ac). Axial sculpture initially of 11-13

strong ribs per whorl that originate at the shoulder but

do not extend to the anterior carina. Ribs become heavi-

er, increase in number to 18-19 on final whorl, and de-

velop open, curved, posteriorly-directed spines along

shoulder by 3rd postnuclear whorl. Spiral sculpture of 3

strong cords between suture and periphery, 2 major

cords between periphery and anterior carina, 4 strong

cords between anterior carina and siphonal canal, and

12-15 major cords along proximal *% of siphonal canal.

One to three weaker cords present between adjacent

major cords anterior of shoulder. Aperture broadly

ovate, tapering anteriorly, deflected from shell! axis by

17°. Outer lip glazed, slightly reflected, forming poste-

riorly directed, open spine at shoulder. Weak furrows on

inner surface of outer lip correspond to shoulder and

major spiral cords. Inner lip smooth, surface sculpture

resorbed prior to deposition of thin porcellaneous glaze.

Siphonal canal long, axial, stout, straight. Shell color uni-

formly white. Periostracum (Figure 25, per) thin, of

overlapping axial lamellae, straw-colored. Operculum

(Figure 27) thin, wedge-shaped, broadly rounded pos-

teriorly, sharply tapering anteriorly, slightly concave

along right side leading to a terminal nucleus.

Holotype a mature, poorly preserved, desiccated fe-

male, with mantle cavity spanning ~%4 whorl, kidney ~%

whorl, upper whorls not recovered. General orientation

and morphology of mantle cavity and cephalic hemocoel

organs as in Coluzea aapta (see Harasewych, 1986:161).

Proboscis, long, tightly folded in proboscis sheath, which

occupied posterior half of cephalic hemocoel. Radula

(Figure 28) short (3.2 mm), narrow (132 ym), with 142

rows of teeth. Lateral teeth monocuspid, recurved, ta-

pering to a point from 20 wm wide attachment area.

Rachidian teeth with 3 well-defined, closely spaced, pos-

teriorly-directed cusps emanating from the center of a

semicircular basal plate that expands laterally.

Type Locality (Figure 30): Off Westem Australia,

North West Shelf, 33 nautical miles S of Bedwell Island,

Clerke Reef, 17°46.76’ S, 119°24.6’ W to 17°45.97’ S,

119°25.6’ W, in 250 m, AIMS Survey R/V Lapy BASTEN

station LB8, 18 August 1995, Sled dredge, substrate of

calcareous mud ang worm tubes.

Type Material: Holotype, 2, WAM S14314, 52.1

mm; Paratype, WAM S14394, 40.1 mm, (with hermit

crab), both from the type locality.

Distribution (Figure 30): This species is presently

known only from its type locality.

Etymology: —naxa—Latin, wicker basket with a narrow

neck.

Remarks: The narrow, fusiform shell, presence of

strong axial ribs that originate at the shoulder but do not

extend to the anterior carina, and open, posteriorly re-

flected spines of Coluzea naxa serve to distinguish it

from all congeners. Coluzea naxa is conchologically most

similar to the New Caledonia species, C oluzea ‘faceta

M. G. Harasewy Page 101

Figures 24-29. Coluzea naxa new species. Dredged off Western Australia, North West Shelf, 33 nautical miles S of Bedwell

Island, Clerke Reef, 17°46.76’ S, 119°24.6’ W to 17°45.97' S, 119°25.6’ W, in 250 m. 24-28. Holotype [WAM S14314]. 24.

Apertural, right lateral and dorsal views of shell. 25. Detail showing periostracum and open spines along shoulder. 26. Protoconch.

27. Outer surface of operculum 28. Dorsal view of mid-section of radular ribbon. 29. Paratype [WAM $14394]. Scale bar (1 cm)

applies to all entire shells. Abbreviations: ac, anterior carina; per, periostracum.

THE NAUTILUS, Vol. 118, No. 3

105°E 110°E 115°E 120°E 125°E 130°E

15°S

20.

25°S ig

30°S

Figure 30. Geographic distribution of Coluzea naxa new

species, open triangle = type locality.

Harasewych, 1991, which differs in having a larger,

heavier shell, with stronger axial ribs that extend beyond

the anterior carina, short, laterally directed spines, a

broader aperture and narrower shoulder.

Coluzea naxa is easily distinguished from the three

geographically more proximal, bathymetrically zoned

species C. distephanotis (Melvill, 1891), C. icarus Har-

asewych, 1986, and C. aapta Harasewych, 1986 [from

off Rowley Shoals, Western Australia] by its smaller,

thinner, shell, prominent rounded axial sculpture, and

posteriorly directed spines. Although C. distephanotis

was originally described from a depth of 31 m (Melvill,

1891), more recent, better-documented samples indicate

that this species inhabits depths in excess of 300 m (Har-

asewych, 1986), while C. icarus and C. aapta live at even

greater depths along the continental slope. As Coluzea

naxa is presently known only from a single station at a

depth of 250 m, additional sampling will be required to

determine the geographic and bathymetric boundaries

among these species.

ACKNOWLEDGMENTS

I am grateful to Drs. Dick Kilburn and Dai Herbert

(NM), Dr. Philippe Bouchet (MNHN), and Dr. Fred

Wells and Correy Whisson (WAM) for making available

most of the specimens on which this study was based.

Special thanks are due to Dr. Martin Avery Snyder for

bringing to my attention the specimens that comprise

the type series of C. naxa.

LITERATURE CITED

Darragh, T. A. 1969. A Revision of the Family Columbariidae

(Mollusca: Gastropoda). Proceedings of the Royal Society

of Victoria 83: 63-119, pls. 3-6.

Harasewych, M. G. 1986. The Columbariinae (Gastropoda:

Turbinellidae) of the eastern Indian Ocean. Journal of the

Malacological Society of Australia 7: 155-170.

THE NAUTILUS 118(3):103-111, 2004

Genetic differentiation in two c

Page 103

tic species of Ostreidae,

Ostrea edulis (Linnaeus, 1758) and Ostreola stentina

(Payraudeau, 1826) in Mar Menor Lagoon, southwestern

Mediterranean Sea

M. Gonzalez-Wangitiemert'

A. Pérez-Ruzafa

Departamento de Ecologia e

Hidrologia

Facultad de Biologia

Universidad de Murcia

30100 Murcia

SPAIN

[email protected]

M. J. Rosique

A. Ortiz

30100 Murcia

SPAIN

Departamento de Biologia Animal

Facultad de Veterinaria

Universidad de Murcia

ABSTRACT

Ostrea edulis is a target species for aquaculture but its hatchery

has suffered as a result of the lack of morphological differen-

tiation between individuals with a low growth and those that

reach commercial size. Two sympatric species of oysters, Os-

trea edulis and Ostreola stentina, have been reported at the

Mar Menor Lagoon, Spain. A third nominal species, Ostreola

parenzani, is now considered a synonym of O. stentina. The

external morphology of O. edulis and O. stentina is very similar

and this prevents their differentiation at the morphological lev-

el, except for maximum size. Oysters were collected from 3

locations along the Mar Menor Lagoon and examined for var-

iation at the PGI locus. Principal component analysis of allo-

zyme data revealed the existence of two groups, which confirms

the presence of two species: Ostrea edulis and Ostreola sten-

tina. The genetic variability of the glucose-6-phosphate isom-

erase (PGI) locus was also compared in Ostrea edulis and Os-

treola stentina from the Mar Menor Lagoon. Ostrea edulis has

high levels of homozygosis and shows an important deviation

from the Hardy-Weinberg equilibrium. Ostreola stentina shows

high heterozygosis and significant differentiation among coastal

lagoon samples. The allele frequencies at the PGI locus can be

used as a diagnostic character at the species level.

INTRODUCTION

Molecular techniques, including cytogenetics (Thiriot-

Quiévreux, 1994) and flow cytometry (Partensky et al.,

1997), provide a range of methods for quantifying the

phylogenetic relationships between species and higher

taxa, defining species limits, and identifying and quan-

tifying cryptic species (Féral, 2002).

Biochemical methods helped demonstrate that many

abundant and ecologically important “species” are, in

' Author for correspondence

fact, groups of species or species complexes (Avise,

1974).

Genetic studies have indicated a remarkably high in-

cidence of cryptic speciation in marine invertebrates

(Knowlton, 1993; Thorpe and Solé-Cava, 1994) includ-

ing marine bivalves (Koehn, 1991; André et al., 1999;

Daguin, 2000) and gastropods (Munksgaard, 1990;

Palmer et al., 1990; Liu et al., 1991; Corte-Real et al.,

1996a; 1996b) and sometimes even in comparatively well

studied commercially important species (Yeatman and

Benzie, 1994; Chan and Chu, 1996; Thorpe et al., 2000).

These genetically differentiated groups often show mi-

nor differences in shell morphology that are not always

consistent with genetic (allozyme) characters (Sarver et

al., 1992). This has important implications for studies on

the biology of the involved species. The overlooked pres-

ence of cryptic species may produce unexpected varia-

tion in physiological or ecological studies.

The demand for high-quality protein, especially from

aquatic sources, is rising dramatically. Increased aqua-

culture production is clearly needed to meet this de-

mand (Dunham et al., 2000). However, aquaculture pro-

ductivity cannot be optimized if the biological potential

of cultured species is not realized. Due to the above-

mentioned difficulties in differentiating some species on

the basis of external morphological characters, the ge-

netic identification and discrimination of aquaculture

stocks and species is a fundamental requirement in any

culture program (Ferguson, 1994).

Oysters have been exploited since the time of the Ro-

man Empire (Magenis et al., 1983), but harvesting on a

large scale began in France around 1850. Spat have been

collected from natural beds and cultured with varying

success due to epizootic diseases (Jaziri et al., 1987).

In the Mar Menor Lagoon, Ostrea edulis have un-

dergone rapid expansion since the early 1980s after the

artificial enlargement of one of the inlets that connect

the lagoon caatin the Mediterranean. In 1992, 177 million

indiv sara were tallied in a survey with a mean density

of 2 oysters/m? reaching 22 oysters/m2 in the most pop-

ulated areas (Rosique and Garcifa-Garcia, 1997). As a re-

sult of these high densities, several attempts at hatching

were made and the spats of this species have been col-

lected from natural beds and cultured with varying de-

grees of success due to epizootic diseases and the irreg-

ular growth of percentile individuals, which does not al-

low for profitable exploitation. Blanc et al. (1986) cited

a similar finding in Nador Lagoon (Morocco). They stud-

ied two populations of oysters that belonged to the same

cohort: 49 individuals of a normal growth population and

49 individuals of slow growth. They concluded that the

fast-growing sample was Ostrea edulis. Of the slow-

growing oysters, only 19% were considered to be Ostrea

edulis, while 81% belonged to another species. More-

over this second species differed from Ostrea edulis by

three loci and appeared to be a dwarf sibling species of

Ostrea edulis with similar larvae and spat.

A second species (Ostreola stentina) inhabits the Mar

Menor Lagoon (Murillo and Talavera, 1983; Olmo and

Ros, 1984; Pérez-Ruzafa, 1989) and is undifferentiated

from Ostrea edulis except for the maximum size reached

by each species. A third species, Ostreola paranzani, has

been reported at the lagoon (Murillo and Talavera, 1983)

although it is considered a synonym for Ostreola stentina

by some authors (Parenzan, 1974).

Ostrea edulis can reach 94 g and 95 mm in weight

and size respectively after thirteen months in culture,

Ostreola stentina does not exceed 20 g in weight and 45

mm in size (Rosique et al., 1995).

Ostrea edulis has a high commercial value and its pop-

ulations have suffered a strong decline due to overex-

ploitation (Yonge, 1960). It is a hermaphroditic, infrali-

toral species with a wide geographical distribution along

the Atlantic coastline from Norway to Morocco, and all

along the Mediterranean as well as the Black Sea (Yonge,

1960; Launey et al., 2002). It has also been introduced

into many other parts of the world (e.g., United States,

Canada and Japan) due to its aquaculture potential (Kor-

ringa, 1976; Launey et al., 2002). Its life history is char-

asternad by fertilisation occurring inside the pallial cav-

ity and the brooding of larvae (Yonge, 1960). As a result

of a brooding period of 8 to 10 days, the length of the

plankton larval phase is reduced compared to that of

other oyster species (Buroker, 1985).

Ostreola stentina is small to medium in size and lives

in shallow subtidal waters to a few meters depth, in trop-

ical and temperate seas (Harry, 1985).

The systematic position of Ostreidae has been studied

in several works (Pasteur-Humbert, 1962; Harry, 1985;

Orton, 1928; Nelson, 1938; Montero, 1971; Stenzel,

1971; Parenzan, 1974) but most of them have not re-

solved all the taxonomic problems. Harry (1985) pre-

sented a good synopsis of the supraspecific classification

of living oysters in which he considered not only the

structure of the flesh and shells but also the environ-

THE NAUTILUS, Vol. 118, No. 3

ments, geographic range, and behavior of oysters. The

author concluded that the intraspecific variation of oys-

ter shells, which is probably greater than in any other

group of living bivalves, precluded the preparation of a

simple and satisfactory taxonomic key. The use of mo-

lecular genetic techniques in oyster systematics has in-

creased over the past several years, largely due to the

increased availability of techniques and increased aware-

ness of the value of genetic data (Littlewood, 1994; Hare

and Avise, 1998; Jozefowick and O Foighil, 1998; Lee et

al., 2000).

Variation in enzyme coding genes has been studied in

recent years in several species of marine bivalves, pro-

viding differentiation among similar species and infor-

mation regarding genetic structure in populations of

these organisms. Several studies of variations at enzyme

loci in Ostrea edulis have been made (Wilkins and Math-

ers, 1973; Buroker, 1982; Maggenis et al., 1983; Johan-

nesson et al., 1989; Le Pennec et al., 1986; Blanc et al.,

1986: Saavedra et al., 1987; 1993; 1995: Alvarez et al.,

1989). Electrophoretic studies have been mainly restrict-

ed to Atlantic populations, which have been very much

affected by human harvesting activities (Yonge, 1960;

Maggenis, et al., 1983). These studies indicated high ge-

netic uniformity, covering restricted areas of the total

range of the species’ distribution (Le Pennec et al., 1986;

Jaziri et al., 1987; Saavedra et al., 1987). Saavedra et al.

(1995) showed that broad macrogeographical clines are

a major feature of allozyme interpopulation variability in

this species. The origin of these clines probably implied

the contact of two Atlantic and Mediterranean oyster

stocks that became differentiated in allopatry. Launey et

al. (2002) studied the genetic differentiation in Ostrea

edulis by means of variation at five microsatellite loci.

The results showed a mild but significant isolation-by-

distance profile, a noticeable between-sample variance

in expected heterozygosity, and a tendency for Atlantic

populations to be less variable than Mediterranean ones.

Comparison with data on allozyme variation in relevant

literature confirms this view.

MATERIALS AND METHODS

Stupy AREA

The Mar Menor is a hypersaline coastal lagoon with a

surface area of about 135Km2. It is located in a semi-

arid region of the southeast of Spain (37°44" N, 0°47’ W)

on the Mediterranean coast. The mean depth is 3.5 m

with a maximum depth of 6 m (Pérez-Ruzafa, 1996). It

has five open inlets, which permit the interchange of

water with the Mediterranean Sea. In the 1970s, one of

these channels (E] Estacio) was dredged and widened,

inducing important changes in the hydrodynamics and

biological communities of the lagoon, including coloni-

zation by new species (Pérez-Ruzafa et al., 1987; 1991).

SAMPLING

In order to analyze the causes of the observed differ-

ential growth in oyster populations and to confirm the

M. Gonzalez-Wangtiemert et al., 2004

El Estacio w

By ®

Mar Menor 72)

6 =

Los ie ©

* ®

—s

Z@)

° ®

37 40° =

bea

0 50° 045°

Figure 1. Sampling localities in Mar Menor Lagoon.

existence of the two reported species in the Mar Menor

Lagoon, thus determining their importance in oyster

hatchery, three localities were sampled at the lagoon in

1996 (Figure 1). Two samples were taken in natural oys-

ter beds at Los Urrutias and Ciervo Island. The third

sample was collected as spat at El Estacio in January

and moved to aquaculture installations at Marbella

(southern Spain) where after 8 months they were col-

lected as adult oysters. This ensures that all individuals

belong to the same cohort.

ELECTROPHORESIS

All oysters were transported live to the laboratory where

they were dissected. Portions of adductor muscle were

removed from each individual, homogenized in 1.5M

Tris buffer (pH 9), and centrifuged at 4°C and 13500xg.

They were stored at —40°C until electrophoresis.

Vertical polyacrylamide gel electrophoresis was car-

ried out at a constant voltage (125 V) for 5 hours at 4°C.

Page 105

Gels were stained for PGI activity as described in Harris

and Hopkinson (1976) with some modifications in the

proportion of reagents (see Gonzdlez-Wangtiemert,

1997).

Isozymes were numbered in decreasing order of mo-

bility starting from the most anodal; allozymes were en-

coded according to the mobility of the most common

allele (100).

Population Genetic Analysis: The existence of ho-

mogeneous genetic groups was explored performing a

Principal Component Analysis (PCA) (ter Braak and

Prentice, 1988) on the matrix of genotypes. The results

of the ordination analysis are displayed in a biplot, scal-

ing the axes, adjusting genotype scores to genotype var-

iance: the resulting scores are correlations between ge-

notypes and eigenvectors. All these calculations were

done using the CANOCO vy. 3.15 package (ter Braak,

1990).

The groups identified by the PCA were characterized

morphologically comparing the maximum length of the

shells (L1) using analyses of variance (ANOVA). Accord-

ing to the results, identified groups were assigned to the

species Ostrea edulis and Ostreola stentina.

The genetic variability of the samples was recorded as

expected and observed heterozygosity (H. and H, re-

spectively) and the deviation coefficient were calculated.

Differences in gene frequencies among three samples of

Ostreola stentina were tested using x? test (two degrees

of freedom).

To analyze spatial differences in populations, a second

PCA analysis was performed on the allelic frequency ma-

trix for the two species and genetic variability descriptors

at each locality.

F-statistics following Wright (1951) were calculated to

detect non-random mating within populations (F,;) and

differentiation between populations (F,,). Both statistics

were calculated via the Weir and Cockerham method

(1984). Probabilities of random departure from zero for

F-values, according to the null hypothesis, were read di-

rectly from the distribution of 1000 randomized matrices

computed via permutation of individuals among popu-

lations. This was performed using the “Genetix” F-test-

ing procedure, thus providing a test of significance.

Genetic distance (Nei’s D; Nei, 1978) was computed

between pairwise samples. Probabilities of random de-

parture from zero for Nei’s D-values, according to the

null hypothesis, were read directly from the distribution

of 1000 randomized matrices computed by permutation.

Gene flow between samples was estimated as the

number of migrants exchanged between populations per

generation at equilibrium (N, m). Values for N, m were

derived from one approach with F,; values, following

Wright's island model (1951).

The data was analyzed using the Genetix package (Bon-

homme et al., 1993) (available at: www.univ-montp2.fr/

genome-pop/genetix.htm).

Page 106

o H

; PGI70 i

S i

+ i

1 ;

i s

: PGIOO

| pcies :

eal :

! si

-1.0 7+1.0

Ostreola stentina

Ostrea edulis

Figure 2. Ordination of first two axes of principal compo-

nent analysis (PCA) of individual genotypes that jointly ex-

plained 91.7% of the variance in the global data set (numbers

in brackets correspond to number of Andlnidivel in that partic-

ular point).

RESULTS

A total of 168 individual oysters have been analyzed for

the glucose-6-phosphate isomerase (PGI). The electro-

phoretic survey shows nine different genotypes. The first

two axes of the PCA analysis (Figure 2) explain 91% of

the total variance in data. The results shows two well-

differentiated groups along the first ordination axis,

which accounted for most of the variation, explaining

67.2% of the total variance.

The results of the ANOVA performed in order to

THE NAUTILUS, Vol. 118, No. 3

11 ==

10 - LOCALITIES

gL O El Estacio-Marbella

L B Ciervo island

8 A Los Urrutias

eS ft

Fj 6

O edulis © sfentina

SPECIES

Figure 3. Significant differences in maximum diameter of

the shell in the two species of oysters in the three sampling

sites in Mar Menor. All individuals grouped as a species in this

figure correspond to the individuals of same species in Figure

2 above.

characterize these groups morphologically show signifi-

cant differences (p<0.001) in maximum size among ge-

netic groups (Figure 3). pes 1 has a mean size of 6.61

cm (+ 0.21). Group 2 has a mean size of 3.16 cm

(+£0.04). The individuals cultivated in Malaga, belonging

to the same cohort, show the same significant differenc-

es in size which are in line with genetic differentiation.

So the two groups, which do not share any alleles, would

correspond to the two species reported at the Mar Men-

or Lagoon, Ostrea edulis with only three alleles and

three genotypes and Ostreola stentina with three alleles

and six genotypes, respectively. The relative frequencies

of all detected genotypes are shown in Table 1.

PGI was encoded by three alleles in Ostrea edulis (Ta-

ble 2), though only three individuals exhibited the

PGI*115 and one individual showed the allele PGI°110

Table 1. Relative frequencies of PGI genotypes in coastal la-

goon oysters.

Genotypes Ostrea edulis Ostrea stentina

N 57 111

°100/100 0.9452 0

*100/115 0.0410 0

°100/110 0.0136 0

°70/70 0 0.0930

°70/85 0 0.3813

°70/95 0 0.1101

*$5/85 0 0.3644

°85/95 0 0.0762

°95/95 0 0.0084

M. Gonzalez-Wangiiemert et al., 2004

Page 107

Table 2. Allele frequencies at PGI locus of Ostrea edulis and Ostreola stentina (N: number of individuals; EE: El Estacio; IC:

Ciervo Island; U: Los Urrutias).

Species Samples N PGI 70° PGI 85° PGI 95° PGI 100° PGI 110° AGH JUS)”

Ostrea edulis EE 20 0 0 0 0.950 0.025 0.025

IC 19 0 0 0 1 0 0

U 18 0 0 0 0.970 0 0.030

EE 76 0.263 0.645 0.092 0) 0 0

IC 22, 0.386 0.568 0.046 0 0) 0

Ostreola stentina U 13 0.385 0.461 0.154 0 0 0

in heterozygous combination (Table 1). The locus can be

regarded as essentially monomorphic in this species.

In Ostreola stentina three alleles of the phosphoglu-

cose isomerase were expressed with frequencies higher

than 0.10, as such the locus can be regarded as poly-

morphic.

Little difference was detected between the observed

and expected heterozygosity. The highest deviation co-

efficient (D) was 0.0278 for Ostrea edulis and 0.111 for

Ostreola stentina (Table 3). The observed heterozygosity

in Ostrea edulis showed low values (ranging from 0.00

to 0.0952) due to PGI*100 being mainly combined as a

homozygote and only four individuals being heterozy-

gotes. Ostreola stentina has higher ahseaved! heterozy-

gosity than Ostrea edulis, since 56% of the individuals

analyzed were heterozygotes.

Allele frequencies at PGI differed significantly among

the three sampled populations of Ostreola stentina

(x2=5.99; P=0.035).

The PCA analyses performed on the allelic frequency

matrix and genetic variability descriptors at each locality

separate both species along the first axis which explains

97.8% of the total variance in data (Figure 4). Ostrea

edulis samples groups in the positive part of the axis are

characterized by a high homozygosity and a low hetero-

zygote deficit. Ostr -wolle stentina populations in the neg-

ative part are characterized by a higher expected and

observed heterozygosity and a high heterozygote deficit.

The second axis explains an additional 1.7% of the total

variance and discriminates mainly among Ostreola sten-

tina populations, with the El Estacio population, closer

to the Mediterranean, in the positive part, with a dom-

inance of PGI 85* allele, and that of Los Urrutias in the

negative part with a dominance of PGI 95° allele and a

higher heterozygote deficit.

Deviations from Hardy-Weinberg proportions within

samples are shown by means of F,, statistic. F,, values

indicated a significant heterozygote excess ranging from

—0.006 to —0.101 in Ostreola stentina (Table 3). Ostrea

edulis showed a deviation from Hardy-Weinberg expec-

tations within the El Estacio sample, although it was not

significant. (Table 3). Nei’s genetic distances (Nei, 1978)

were estimated using PGI locus (Table 4) in Ostreola

stentina. Values ranged from —0.024 to 0.017. All dis-

tances were not significant at the 0.05 level. Estimates

of genetic subdivision (F,,) in the three samples are giv-

en in Table 4. The minimum positive F,; value derived

from allelic variation was found between the El Estacio

and Ciervo Island samples (0.008), showing low diver-

gence in gene frequencies between the two populations.

In contrast, F.; was considerably higher (0.026) between

E] Estacio and Los Urrutias samples, suggesting the pos-

sible occurrence of restricted gene flow between these

populations. F,; values between samples were significant

at the 0.05 level.

Assuming equilibrium between genetic drift and mi-

gration, we calculated the number of migrants (N.m) per

generation (Table 5), based on Fs; values and according

to the island model. Estimates of the number of mi-

grants ranged from 9.16 (El Estacio-Los Urrutias) to in-

finite (Ciervo Island-Los Urrutias).

DISCUSSION

Allele frequencies at the PGI locus, used as a species-

diagnosing character, allow the differentiation of the two

sympatric oyster species studied. The coexistence of

both species could explain the disastrous oyster hatchery

attempt in the Mar Menor Lagoon (Rosique et al.,

1995).

Table 3. Observed and expected heterozygosities (H, and H.), deviation coefficient (D) and F,, (ns: non-significance; *: p<0.05;

EE: El Estacio; IC: Ciervo Island; U: Los Urrutias).

Species Samples H, H. D Fis

Ostrea edulis EE 0.0952 0.0963 —(0.0106 0.013 ns

IC 0 0 0 =

U 0.0556 0.0540 0.0278 0 ns

EE 0.5132 0.5066 0.0129 —(0.006*

IC 0.5909 0.5258 0.1101 —0.101*

Ostreola stentina U 0.6923 0.6154 0.1111 —(0.085*

fe ay g

Page 108

PGIllI9 ©

OEE!

Ostrea edulis

-1.0 +1.0

Figure 4. Ordination of first two axes of principal compo-

nent analysis (PCA) of allele frequencies that jointly explained

99.5% of the variance in the global data set. (EE:, El Estacio;

IC: Ciervo Island; U: Los Urrutias).

Genetic variation within and between populations has

been demonstrated by the use of electrophoresis. We

now have some information regarding the frequencies

and distribution of alleles in wild populations of Ostrea

edulis and Ostreola stentina in the Mar Menor Lagoon.

Lower levels of genetic variation and heterozygote def-

icit were detected in the Ostrea edulis population.

All the electrophoresis studies on Ostrea edulis pop-

ulations coincide in that this species displays lower levels

of allozyme variation than other bivalves (Buroker, 1982;

Saavedra et al., 1987) and the overall differentiation

THE NAUTILUS, Vol. 118, No. 3

Table 4. Pairwise Nei’s genetic distances (below the diagonal)

and F,, values (above the diagonal) in Ostreola stentina. F.;

and Neis D considered to be significantly different from zero

(°) if they fall within the 5% most extreme values in the per-

mutation test. (ns: non-significance; ° = p<0.05; EE: El Es-

tacio; IC: Cievo Island; U:Los Urrutias).

EE IC U

EE ~ 0.082° 0).0266°

IC 0.0080 ns ~ —(0.0076°

U 0.017 ns —().024 ns ~

among its populations is usually slight (Johannesson et

al., 1989). In fact Saavedra et al. (1993) showed, through

an UPGMA dendrogram based on Nei unbiased ge-

netic distances, two main clusters, one formed by the

eastern Mediterranean samples and the other by the re-

maining populations (western Mediterranean and Atlan-

tic samples).

Two of the Ostrea edulis populations studied (El Es-

tacio and Los Urrutias) have very low observed hetero-

zygosity, though a significant deficit in heterozygotes is

not observed. The Ciervo island population has 100%

homozygotic individuals, so that this population shows

an excessive heterozygote deficit and an important de-

viation from the Hardy-Weinberg equilibrium.

Some researchers have documented a deficit in het-

erozygotes for populations of Ostrea edulis from Atlantic

oyster beds (Buroker, 1982; Maggenis et al., 1983; Jo-

hannesson, et al., 1989; Saavedra et al., 1995; Launey et

al., 2002). The biological origin of these heterozygote

genotype deficiencies may be related to fecundation.

This takes place inside the pallial cavity of the female,

which favors mating between nearest-neighbors. Also,

larvae are brooded for a period of 8 or 10 days before

the plankton phase, which limits dispersal. In addition,

the extremely low levels of variability detected may to

some extent be due to the recent history and exploitation

of these populations (Saavedra et al., 1993).

Apart from chance alone, a number of factors may be

responsible for causing deficiencies in heterozygotes

against the H-W model in allozyme data. These include,

null alleles, the Walhund effect, inbreeding and selection

against heterozygotes or strong directional selection as a

consequence of the geographic isolation of some popu-

lations (Zouros and Foltz, 1984; Mamuris et al., 1998;

Rossi et al., 1998).

The low levels of observed allozyme variation in the

Table 5. Ostreola stentina. Estimates of N.m using F,; values

(Wright, 1951) (EE: El Estacio; IC: Ciervo Island; U: Los Ur-

rutias).

EE IC U

EE =

IC 30.28 a

U 9.16 =

M. Gonzdlez-Wangiiemert et al., 2004

Page 109

Ostrea edulis populations of the Mar Menor Lagoon,

may be due to the recent history and exploitation of

these populations. The current Ostrea edulis oyster bed

in the Mar Menor Lagoon could come from oyster beds

harvested for commercial purposes in NW Spain (Ro-

sique per. com.). The transplantation of farmed stocks

from Atlantic populations to Mediterranean populations

has been a common occurrence (Launey et al., 2002).

This hypothesis is reinforced due to the fact that the

Ostrea edulis population from the Mar Menor Lagoon

showed a lower heterozygosity than Mediterranean pop-

ulations, and similar values to Atlantic populations (Ar-

ousa and Ares, NW Spain; Saavedra et al., 1993) and

those of NW France (Jaziri et al., 1987). Allozyme and

microsatellite studies have shown a lower genetic vari-

ability in Atlantic populations than in Mediterranean

ones. This result could be explained by an overall smaller

evolutionary effective size for Atlantic populations com-

pared to Mediterranean populations and two main ex-

planations have been put forward for such a difference:

variance in effective sizes and oyster parasites (Launey

et al., 2002).

High levels of variation were evident in populations

of Ostreola stentina. This species shows six different ge-

notypes for phosphoglucose isomerase and a high ob-

served heterozygosity. This high variability could be due

to long larval period (Harry, 1985) which could favor the

dispersion of the gene pool. Some authors affirm that

patterns of variability at the PGI locus in bivalves suggest

that species inhabiting temporally variable or spatially

heterogeneous environments exhibit higher levels of ge-

netic variability than those from less variable or more

monotonous environments (Valentine and Ayala, 1978).

This agrees with the fact that the Mar Menor Lagoon

shows a high degree of isolation with respect to the

Mediterranean and highly variable environmental con-

ditions (Pérez-Ruzafa, 1996) explaining the high genetic

variability in Ostreola stentina.

F,, values among Ostreola stentina populations are

always lower than 0.1, and although significant, are in-

dicative that there is little divergence among populations

(Hartl, 2000). The fact that the Ciervo Island and Los

Urrutias localities show infinite rates of interchange of

individuals and negative F,; and genetic distance values,

suggests that both localities have the same Ostreola sten-

tina population. Further genetic studies using several

loci are required to confirm this hypothesis.

The results of this study confirm that there are two

species (Ostrea edulis and Ostreola stentina) in the Mar

Menor Lagoon stock and the alleles at the PGI locus can

be used as a species-diagnosing character. As this situa-

tion can be a common state in the distribution area of

both species, some works related to ecological and phys-

iological adaptations or ecotoxicological responses of any

of them should be reviewed. Some marine molluscs reg-

ulate their body tissue levels of particular trace metals

to constant levels over a wide range of metal levels in

their environment (Rainbow et al., 1990). The laboratory

experiments have also provided evidence that this reg-

ulation is species-specific (Bryan et al., 1985; Rainbow

et al., 1990; Dallinger and Rainbow, 1993) so that the

existence of two cryptic species could change the con-

clusions of some toxicological works in Ostrea edulis

(George et al., 1978; atiret et al., 2002). Similar con-

siderations could be applied to Ostrea edulis physiolog-

ical studies (Beiras et al., 1995; Labarta et al., 1999: Cul-

loty et al., 2001; Culloty et al., 2002) and works on the

resistance of this species to the parasite Bonamia ostreae

(Elston et al., 1987; Culloty and Mulcahy, 1996; Naciri-

Graven et al., 1998; Naciri-Graven et al., 1999).

ACKNOWLEDGMENTS

We are grateful to Dr. J. Cano, IEO (Instituto Espafiol

de Oceanografia, Malaga) and some volunteers who as-

sisted in the laboratory. We also want to thank to Dr. R.

T. Dillon, Jr. (College of Charleston, Charleston, South

Carolina) and to the anonymous referees for their com-

ments and suggestions that have enhanced the final ver-

sion of the manuscript.

LITERATURE CITED

Alvarez, G., C. Zapata, R. Amaro and A. Guerra. 1989. Mul-

tilocus heterozygosity at protein loci and fitness in the Eu-

ropean oyster, Ostrea edulis (L.). Heredity 63: 359-372.

André, C., M. Lindegarth, P. R. Jonsson and P. Sundberg.

1999. Species identification of bivalve larvae using random

amplified polymorphic DNA (RAPD): differentiation be-

tween Cerastoderma edule and C. lamarcki. Journal of the

Marine Biological Association of the United Kingdom 79:

563-565.

Auffret, M., N. Mujdzic, C. Corporeau and D. Moraga. 2002.

Xenobiotic-induced inmunomodulation in the European

flat oyster, Ostrea edulis. Marine Environmental Research

54: 585-589.

Avise, J. C. 1974. Systematic value of electrophoretic data. Sys-

tematic Zoology 23: 465-481.

Beiras, R., A. Pérez-Camacho and M. Albentosa. 1995. Short-

term alterations in the energy budget of young oyster Os-

trea edulis L. in response to temperature. Journal Exper-

imental Marine Biology and Ecology 186: 221-236.

Blanc, F., H. Jazira and P. Durand. 1986. Isolement génétique

et taxonomie des huitres planes dans une lagune du sud

de la Méditerranée occidentale. In: Systematic Animal

Comptes Rendues de la Académie des Sciences, Paris,

303, série 3, 6: 207-210.

Bonhomme, F., K. Belkhir, P. Borsa, E. Mathieu and M. Roux.

1993. GENETIX-Logiciel D’analyse Des Données Du

Groupe de Génétique Des Populations de Montpellier,

Version 0.1. Université Montpellier II, France.

Bryan, G. W., W. J. Langston, L. G. Hummerstone and G. R.

Burt. 1985. A guide to the assesment of heavy metal con-

tamination in estuarines using biological indicators. Jour-

nal of the Marine Biological Association of the United

Kingdom, Occasional Publication 4: 1-92.

Buroker, N. E. 1982. Allozyme variation in three non-sibling

Ostrea species. Journal of Shellfish Research 2: 157-163.

Buroker, N. E. 1985. Evolutionary pattems in the family Os-

treidae: larviparity vs oviparity. Jounal of Experimental

Marine Biology and Ecology 90: 233-247.

Page 110

Chan, T. Y. and K. H. Chu. 1996. On the different forms of

Panulirus longipes femoristriga (Von Marstens, 1872)

(Crustacea: Decapoda: Palinuridae), with a description of

a new species. Journal of Natural History 30: 367-387.

Corte-Real, H. B. S., S. J. Hawkins and J. P. Thorpe. 1996a.

An interpretation of the taxonomic relationship between

the limpets Patella rustica and P. piperata. Journal of the

Marine Biological Association U.K. 76: 717-732.

Corte-Real, H. B. S., S. J. Hawkins and J. P. Thorpe. 1996b.

Population differentiation and genetic confirmation of the

taxonomic status of the exploited limpet Patella candei in

the Macaronesian islands (Azores, Madeira, Canaries).

Marine Biology 125: 141-152.

Culloty, S. C., M. A. Cronin and M. F. Mulcahy. 2001. An

investigation into the relative resistance of Irish flat oysters

Ostrea edulis L. to the parasite Bonamia ostreae (Pichot

et al., 1980). Aquaculture 199: 229-244.

Culloty, S. C., P. F. Dugan, X. Quishi and M. F. Mulcahy. 2002.

Amylase and aspartate aminotransferase in the haemo-

lymph of the European flat oysters Ostrea edulis. Fish &

Shellfish Immunology 12: 367-369.

Culloty, S. C. and M. F. Mulcahy. 1996. Season-, age, and sex-

related variation in the prevalence of bonamiasis in flat

oysters (Ostrea edulis) L. on the south coast of Ireland.

Aquaculture 64: 237-242.

Daguin, C. 2000. Phylogéographie des moules du complexe

despéces Mytilus edulis Thése de l Université Montpellier

II, 103 pp. + annexes.

Dallinger, R. and P. Rainbow. 1993. Ecotoxicology of metals in

fanart nna, CRC Press-Levis Publishers/SETAC. Boca

Raton, 461 pp.

Dunham, R. A., K. Majumdar, E. Hallerman, D. Bartley, G

Mair and G. Hulata, et al. 2000. Review of the status of

aquaculture genetics. http:/Avww.fao.org/DOCREP/003/

AB412E/ab412e03.htm. pp 42.

Elston, R. A., M. L. Kent and M. T. Wilkinson. 1987. Resis-

tance of Ostrea edulis to Bonamia ostreae infection. Aqua-

culture 64; 237-242.

Féral, J. P. 2002. How useful are the genetic markers in at-

tempts to understand and manage marine biodiversity?

Journal of Experimental Marine Biology and Ecology, 268:

121-145.

Ferguson, A. 1994. Molecular genetics and fisheries. Current

and future perspectives. Revision Fish Biology and Fish-

eries 4: 389-392.

George, S. G., B. J. S. Pirie, Ar. Cheyne, T. L. Coombe and P.

T. Grant. 1978. Detoxication of metals by marine bivalves

an iltraestructural study of the compartimentalization of

copper and zinc in the oyster Ostrea edulis. Marine Bi-

ology 45: 147-156.

Gonzalez-Wangiiemert, M. 1997. Variabilidad morfolégica y

del locus PCI de Cardiwm glauwcum en el Mar Menor (SE

de Espafia) y su relacién con las condiciones ambientales.

Thesis of Licenciature, University of Murcia, Spain. 149

PP:

Hare, M. P. and J. C. Avise. 1998. Population structure in the

American oyster as inferred by nuclear gene genealogies.

Molecular Biology Evolution 15: 119-128.

Harris, H. and D. A. Hopkinson. 1976. Handbook of enzyme

electrophoresis in human genetics. Elsevier, Amsterdam.

350 pp.

Harry, H. W. 1985. Synopsis of the Supraspecific Classification

of living oysters, (Bivalvia:Gryphaeidae and Ostreidae).

The Veliger 28: 121-158.

THE NAUTILUS, Vol. 118, No. 3

Hartl, D. L. 2000. A primer of population genetics. Sinauer

Associates, Sunderland, 221

Jaziri, H., P. Durano, P. Pichot and F. Blanc. 1987. Genetic

diversity between and within populations of the European

oyster, Ostrea edulis. Proceedings World Symposium on

Selection, Hybridization and Genetic Engineering in

Aquaculture, Bordeaux. Vol. I. Berlin.

Johannesson, K., M. Rédstrém and H. Aase. 1989. Low ge-

netic variability in Scandinavian populations of Ostrea

edulis (IL). Possible causes and implications. Journal of

Experimental Marine Biology and Ecology 128: 177-190.

Jozetowick, C. J. and D. 6) Foighil. 1998. Phylogenetic analysis

of southern hemisphere flat oysters based on partial mi-

tochondrial 16S rDNA gene sequences. Molecular Phy-

logenetic Evolution 10: 426-435.

Knowlton, N. 1993. Sibling species in the sea. Annual Revision

Ecology Systematic 24: 189-216.

Koehn, R. K. 1991. The genetics and taxonomy of species in

the genus Mytilus. Aquaculture 94: 125-145.

Korringa, P. 1976. Farming the flat oysters of the genus Os-

trea—a multidisciplinary treatise. Developments in aqua-

culture and fisheries science. vol 3. Amsterdam. Elsevier.

Labarta, U., M. J. Femdndez-Reiriz and A. P’rez-Camacho.

1999. Dynamics of fatty acids in the larval development,

metamorphosis and post- metamorphosis of Ostrea edulis

(L.). Comparative Biochemestry and Physiology 123: 249—

254.

Launey, S., C. Ledu, P. Boudry, F. Bonhomme and Y. Naciri-

Graven. 2002. Geographic structure in the European flat

oyster (Ostrea edulis LL) as revealed by microsatellite poly-

morphism. The American Genetic Association 93: 331—

338.

Lee, S. Y., D. W. Park, H. S. An and S. H. Kim. 2000. Phy-

logenetic relationship among four species of Korean oys-

ters based on mitochondrial 16S r DNA and CO! gene.

Korean Journal of Biological Sciences 16: 203-211.

Le Pennec, M., D. Moraga, F. Blanc, P. Pichot and C. Thiriot-

Quievreux. 1986. Recherche de différences morphogé-

nétiques, biochimiques et cytogénétiques entre Ostrea ed-

ulis, sensu stricto et O. edulis “pied de cheval”. Vie Marine

7:19-39.

Littlewood, D. T. J. 1994. Molecular phylogenetics of cupped

oysters based on partial 25S rRNA gene sequences. Mo-

lecular Phylogenetic Evolution 3: 221-229.

Liu, L. L., D. W. Foltz and W. B. Stickle. 1991. Genetic pop-

ulation structure of the southern oyster drill Stramonita

(=Thais) haemastoma. Marine Biology 111: 71-79.

Mamuris, Z., A. P. Apostolidis and C. Trianta-Phyllidis. 1998.

Genetic protein in red mullet (Mullus barbatus) and

striped red mullet (M.swrmuletus) populations from the

Mediterranean Sea. Marine Biology 130: 353-360.

Maggenis, B. A., E. Gosling and N. P. Wilkins. 1983. Irish

oyster populations: a historical and genetic history. Pro-

ceedings of Royal Irish Academy 83B: 291-299.

Montero, I. 1971. Moluscos Bivalvos Espafiles. Anales de la

Universidad de Sevilla 5: 1-358.

Muksgaard, C. 1990. Electrophoretic separation of morpholog-

‘geil similar species of the genus Rissoa (Gastropoda: Pro-

sobranchia). Ophelia 31: 97-104.

Murillo, L., P. A. Talavera. 1983. Aportacién a la malacologia

de una laguna litoral: el Mar Menor (Murcia). Iberus 3:

15-28.

Naciri-Graven, Y., A. G. Martin, J. P. Baud, T. Renault and A.

Gérard. 1998. Selecting the flat oyster Ostrea edulis (L.)

M. Gonzdlez-Wangtiemert et al., 2004

for survival when infected with the parasite Bonamia os-

treae. Journal Experimental Marine Biology and Ecology

294: 91-107.

Naciri-Graven, Y., J. Haure, A. Gérard and J. P. Band. 1999.

Comparative growth of Bonamia ostreae resistant and wild

flat oyster Ostrea edulis in an intensive system: II. Second

year of the experiment. Aquaculture 171: 195-208.

Nei, M. 1978. Estimation of average heterozygosity and genetic

distance from a small number of individuals. Gants,

Austin Texas 89: 583-590.

Nelson, T. C. 1938. The feeding mechanism of the oyster. I.

On the pallium and the branchial chambers of Ostrea vir-

ginica, Ostrea edulis and Ostrea angulata with compari-

sons with other species of the genus. Journal of Mor-

phometry 63: 1-61.

Olmo, R. and J. D. Ros. 1984. Las malacocenosis del Mar

Menor. Estudio y comparacién con comunidades de me-

dios lagunares semejantes. Actas 4° Simp. Ibér. Est. Ben-

thos Marinho, I: 253-260.

Orton, J. H. 1928. The dominant species of Ostrea. Nature

121 (3044): 320-321.

Palmer, A. R., S. D. Gayron and D. S. Woodruff. 1990. Re-

productive, morphological, and genetic evidence for two

cryptic species of northeastern pacific Nucella. The Veliger

33: 325-338.

Parenzan, P. 1974. Carta didentita delle conchiglie del Medi-

terraneo. Volume II. Bivalvi. Prima Parte. Ed. Bios Taras.

Taranto, 277 pp

Partensky, F., L. Guillon, N. Simon and D. Vaulot. 1997. Re-

cent advances in the use of molecular techniques to asses

the genetic diversity of marine photoshyntetic microor-

ganisms. In: Féral, J.-P., Boucher, G (Eds.). Biodiversity

in Dispersive Environments. Vie Milieu-Life Environ-

mental 47 (4), pp. 367-374.

Pasteur-Humbert, C. 1962. Les mollusques marins testacés du

Maroc. Parte II: Les lamellibranches et les scaphopodes.

Travaux de Vinstitut Scientifique chérifen. Série Zoologie

28, Rabat, 245 pp.

Pérez-Ruzafa, A. 1989. Estudio ecoldgico y bionémico de los

poblamientos bentonicos del Mar Menor (Murcia, SE de

Espaiia). Thesis. Universidad de Murcia, 356 pp.

Pérez-Ruzafa, A. 1996. The Mediterranean lagoons. The Mar

Menor, Spain. In Management of Mediterranean Wet-

lands (Murillo, C. and Gonzalez, J. L. eds.). Ministerio de

Medio Ambiente, Madrid, pp. 133-155.

Pérez-Ruzafa, A., C. Marcos, I. M. Pérez-Ruzafa and J. D. Ros.

1987. Evolucion de las caracteristicas ambientales y de los

poblamientos del Mar Menor (Murcia, SE de Espaiia).

Anales de Biologia, 12 (Biologia Ambiental, 3): 53-65.

Pérez-Ruzafa A., C. Marcos and J. D. Ros. 1991. Environmen-

tal and biological changes related to recent human activ-

ities in the Mar Menor. Marine Pollution Bulletin 23: 747—

751.

Rainbow, R. S., D. J. Phillips and M. Depledge. 1990. The

significance of trace metal concentration in marine inver-

tebrates a need for laboratory investigation of accumula-

tion strategies. Marine Pollution Bulletin 21: 321-324.

Rosique, M. J., B. Garcia-Garcia and M. Rosique. 1995. Pri-

mera aproximacion a la identificaci6n del comportamiento

en cultivo de dos especies de ostreidos del Mar Menor.

Actas del V Congreso Nacional de Acuicultura. Ministerio

Page 111

de Agricultura, Pesca y Alimentacion, Cartagena, Murcia,

pp. 106-112.

Rosique, M. J. and B. Garcia-Garcia. 1997. Distribucidn es-

pacio temporal y caracteristicas biométricas de la pobla-

cién de ostra plana (Ostrea edulis) en el Mar Menor. Actas

del VI Congreso Nacional de Acuicultura Ministerio de

Agricultura, Pesca y Alimentacion, Cartagena, Murcia, pp.

353-358.

Rossi, A. R., M. Capula, D. Crosetti, L. Sola and D. E. Camp-

ton. 1998. Allozyme variation in global populations of

striped mullet, Mugil cephalus (pieces Mugilidae). Marine

Biology 131: 203-212.

Saavedra, C, C. Zapata, A. Guerra and G. Alvarez. 1987. Ge-

netic structure of populations of flat oyster (Ostrea edulis,

Linné, 1758) from the NW of the Iberian Peninsula. In-

vestigacién Pesquera 51: 225-241.

Saavedra, C., C. Zapata, A. Guerra and G. Alvarez. 1993. Al-

lozyme variation in European populations of the oyster

Ostrea edulis. Marine Biology. 115: 85-95.

Saavedra, C., C. Zapata and G. Alvarez. 1995. Geographical

patterns at variability at allozyme loci in the European

oyster Ostrea edulis. “Marine Biology 122: 95-104.

Sarver, S. K., M. Katoh and D. W. Foltz, 1992. Apparent over-

dominance of enzyme specific activity in two marine bi-

valves. Genetica 85: 231-239.

Stenzel, H. B. 1971. Oysters. In R. C. Moore, Treatise on in-

vertebrate paleontology. Part.N, vol.3, Mollusca 6, Bival-

via. Geological Society of America, pp. 953-1224.

ter Braak, C. J. F. 1990. Update notes: CANOCO v3.10. Ag-

ricultural Mathematics Group, Wageningen.

ter Braak, C. J. F. and I. C. Prentice. 1988. A theory of gradient

analysis. Advanced Ecology Research 18: 271-317

Thiriot-Quiévreux, C. 1994. Advances in cytogenetics of aquat-

ic organisms. In: Beaumont, A.R (ed.), Genetics and Evo-

lution of Aquatic Organisms, Chapman & Hall, London,

pp. 369-388.

Thorpe, J. P. and A. M. Solé-Cava. 1994. The use of allozyme

electrophoresis in invertebrate systematics. Zoological

Scripta 23; 3-18.

Thorpe, J. P., A. M. Solé-Cava and P.C. Watts. 2000. Exploited

marine invertebrates: genetics and fisheries. Hydrobiolo-

gia 420: 165-184.

Valentine, J. W. and F. J. Ayala. 1978. Adaptative strategies in

the sea. Genetic variation and resource stability in marine

invertebrates. In: Marine Organisms (eds. Battaglia and

Beardmore), pp: 323-346.

Weir, B. S. and C. C. Cockerham. 1984. Estimating F-statistics

for the analysis of population structure. Evolution 38:

1358-1370

Wilkins, N. P. and N. F. Mathers. 1973. Enzyme polymor-

phisms in the European oyster, Ostrea edulis L. Animal

Blood Groups Biochemical Genetics 4: 41-47.

Wright, S. 1951. The genetical structure of populations. Annual

Eugenetics 15: 323-354.

Yeatman, J. and J. A. H. Benzie. 1994. Genetic structure and

distribution of Photololigo spp. in Australia. Marine Bi-

ology 118: 79-87.

Yonge, C. M. 1960. Oysters. 2nd edition. Collins, London, 209

Tae E. and D. W. Foltz. 1984. Possible explanations of

heterozygote deficiency in bivalve molluscs. Malacologia

25: 583-591.

THE NAUTILUS 118(3):112-120, 2004

Page 112

The genus Margarella Thiele, 1893 (Gastropoda: Trochidae) in

the southwestern Atlantic Ocean

Diego G. Zelaya

Division of Invertebrate Zoology

Museo de La Plata

1900 La Plata, Buenos Aires

ARGENTINA

[email protected]

ABSTRACT

The taxonomic status of eight species from the southwestern

Atlantic and adjacent sub-Antarctic and Antarctic waters, pre-

viously referred to either as Margarella or Margarites Gray,

1847, is analyzed. The presence of a first marginal tooth of the

radula forming a “protolateromarginal” plate, a long afferent

membrane in he ctenidium and the presence of antero-lateral

expansions on the foot (horn-like projections) clearly indicate

that southwestern Atlantic species belong to Mar; garella. On

the basis of a morphological analysis of representatives of the

genus type species, Margarella is redescribed and the genera

Margarites, Margarita, Margaritella and Valvatella are exclud-

ed from the Southwestern Atlantic Ocean list of species. The

placement of Margarella within the Gibbulini tribe of the sub-

family Trochinae is re-affirmed based on anatomical and be-

havioral observations.

INTRODUCTION

Margarella Thiele, 1893 is a high-latitude, southern

hemisphere genus of small size trochoid gastropods liv-

ing on brown algae or on hard substrata mainly in the

anita or shallow subtidal. Members of this genus

have been formerly described or reported, among auners

(see below), under the genus Margarita Leach, 1819, a

synonym of Margarites Gray, 1847, a taxon well known

as distributed in the northern hemisphere.

A “not rudimentary innermost lateral plate” (i.e. the

innermost marginal tooth with a laterally expanded base

and a well-developed shaft and cutting edge) was de-

scribed for Margarella expansa (Sowerby, 1838), the

type species of ine genus. This character was also re-

ported by Dembrosi (1969) as present in M. violacea

(King and Broderip, 1832), M. achilles (Strebel, 1908),

M. tropidophoroides (Strebel, 1908) and M. obsoleta

Powell, 1951. Powell (1951) described Margarella bou-

vetia from Bouvet Island, and although he figured the

first marginal tooth as a reduced plate (as it is charac-

teristic in the northern hemisphere genus Margarites),

the species was placed under Margarella. Deambrosi

(1969) described the first marginal tooth as “rudimentary

without cutting edge” in Margarella antarctica (Lamy,

1905), M. bouvetia, and M. expansa, as did Amaud

(1972a) for Margarella refulgens (E. A. Smith, 1907) and

Margarella crebrilirulata (E. A. Smith, 1907). Accord-

ingly, these species were assigned to Margarites. From

previous literature records both Margarella and Mar

garites have been recognized as living in sub-Antarctic

Fat Antarctic waters. In contrast, Hickman and McLean

(1990) questioned the presence of Margarites in the

Southern Hemisphere, and Reid and Osorio (2000), re-

ferring to Margarella sensu lato, pointed out that “con-

fusion surrounds the specific identification, generic as-

signment and relationships of Magellanic and Antarctic

species”.

In the present paper the taxonomic status of species

from the southwestern Atlantic and adjacent sub-Ant-

arctic and Antarctic waters, previously referred to Mar-

garella and Margarites, is analyzed based on the study

at the radula, een, epipodial tentacles, and shell

morphology.

MATERIALS AND METHODS

Representatives of eight Margarella species from the

Magellanic, sub-Antarctic, and Antarctic regions were

studied (Table 1). Radulae of adult specimens were dis-

sected, cleaned by rinsing in a sodium hypochlorite so-

lution and appropriately mounted for scanning electron

microscopy (SEM). Light microscope radula prepara-

tions used by Deambrosi (1969) were re-mounted for

analysis under SEM. Radulae from all studied species

were figured through SEM and described following

Hickman and Miele! s (1990) terminology. Een

anatomy in Margarella expansa, M. violacea, M. antarc-

tica, and M. steineni was studied. In addition, live speci-

mens of Margarella expansa from Ushuaia, Tierra del

Fuego, Argentina, were observed in laboratory conditions.

Voucher specimens are housed at Museo de La Plata

(MLP) and Museo Argentino de Ciencias Naturales

“Bernandino Rivadavia” (MACN). Photographs of two of

the three syntypes of Margarella violacea and the ho-

D. G. Zelaya, 2004

Page 113

Table 1. Species of Margarella examined in this study. (*): only radula preparations made by Deambrosi (1969).

Species Locality Repository

P MLP unnumbered

Margarella achilles (Strebel, 1908) (+)

Margarella antarctica (Lamy, 1905)

Margarella bouvetia Powell, 1951 (*)

Margarella expansa (Sowerby, 1838) (*)

Margarella expansa (Sowerby, 1838)

Margarella obsoleta Powell, 1951

Margarella steineni (Strebel, 1905)

Margarella tropidophoroides Strebel, 1908

Margarella violacea (King and Broderip, 1832)

Melchior Is., Antarctica

Ushuaia, Tierra del Fuego

South Georgia Islands

Inutil Bay, Magellan Strait

MLP 5623

MLP unnumbered

MLP 25241

MLP 7349

MACN 13526

MACN 13573

MACN 18939

MLP 7350

lotype of Margarella jason from the Natural History

Museum, London (BMNH) were used for comparative

purposes.

SYSTEMATICS

Margarella Thiele, 1893

Margarella Thiele, 1893, p. 406 (replacement name for Mar-

garitella Thiele, 1891, p. 259).

Type species by subsequent designation (Thiele, 1924): Tro-

chus (Photinula) expansus Sowerby, 1838.

Diagnosis: Margarella is characterized by having the

first marginal tooth of the radula forming a protolater-

omarginal plate, dimorphic neck lobes, four pairs of epi-

podial tentacles emerging from a simple epipodial tissue,

and foot expanded in two antero-lateral horn-like pro-

jections. The shell is small to medium size, low spired,

expanded at the last whorl; the columella is thick, with-

out denticles and interior of shell nacreous.

MORPHOLOGY AND ANATOMY OF SOUTHWESTERN

ATLANTIC SPECIES:

Shell: In the species of Margarella examined (Table

1), the shell is small to medium size (up to 22 mm wide),

trochoid, usually globose, solid, of up to 4 whorls, with

a relatively high whorl increment rate. The protoconch,

about 700 wm wide, is smooth, whitish, glossy, with one

whorl (Figure 21). The spire is low, with rounded

whorls, the last expanded, somewhat globose, sometimes

angulate (Figures 23-28). The surface is glossy or dull,

smooth or spirally sculptured (Figures 23-28). The ap-

erture is large, rounded, or obliquely ovate. The aper-

ture lip is simple and smooth, and the columella is thick,

without denticles. The interior is nacreous. Adult spec-

imens have a wide white columelar callus with a median

longitudinal depression. The umbilicus is open in young

specimens and may be open or closed in adults (Figures

DR),

In M. expansa the shell is composed of two well-dif-

ferentiated ultrastructural layers: the outer layer of pris-

matic crystals, the inner, thicker, with a laminar structure

(Figure 22).

All studied species have a circular, multispiral, with a

short growing edge, fully corneous, thin and brownish

operculum, that can be deeply retracted into the shell.

Radula: All species of Margarella examined (Table 1)

showed the same general radular morphology: the large,

regularly arcuate, bilaterally symmetrical radula is of the

rhipidoglossate type; rachidian, lateral, and marginal

teeth show characteristically well-developed, overhang-

ing cutting edges (Figure 1). The base of the rachidian

tooth has a pyriform outline, sometimes ovate or greatly

expanded at the base (Figures 2-4), The M-shaped top

of central tooth has a large central cusp, rounded at the

tip, and 6—8 narrower and smaller pointed cusps on

each side (Figure 5). Each of the five lateral teeth is

elongated and of similar shape; their laterally expanded

bases overlap; the basal profile is equivalent to the half

of that of the central tooth (Figures 2, 3). The tongue-

like cutting edge on the lateral tooth is serrated on both

sides and slightly asymmetrical, nearly reaching the tip

of the cutting edge only on the inner margin (Figure 6).

The first marginal tooth has a laterally enlarged base,

and a shaft and cutting edge similar to that of the re-

maining marginal teeth (this tooth element corresponds

to the so called protolateromarginal plate) (Figures 7—

14). The remaining marginal teeth are numerous (about

40 on each half), long and narrow, with a serrated cusp

and a sickle—like-shaped distal end when viewed laterally

(Figures 15, 16, 18). Within a row, the marginal teeth

are steeply graded in solidness and shape: the innermost

are stronger, with well-developed serrated edges and the

outermost delicate with more sparsely serrated cutting

edges (Figures 15, 16, 18). In posterior view, the bases

of the marginal teeth bend in a finely serrated heel; the

outer edge of the shaft is serrated, bearing small spines

(Figure 17).

Anatomy: Specimens of Margarella expansa, M. vio-

lacea, M. antarctica, and M. steineni were examined.

The snout is broad, short, and tubular, with a split ven-

tral lip and the distal end expanded to form an oral disk;

the snout projects laterally as two flaps. Small and simple

cephalic lappets are present. Long, narrow, and cylin-

drical cephalic tentacles arise between the base of the

cephalic lappets and the stout eyestalks. The right eye-

stalk bears a long and narrow appendix at its base. The

neck lobes are dimorphic: the right one is simple, form-

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THE NAUTILUS, Vol. 118, No. 3

Figures 1-6.

Details of the central tooth. 3

= 10 pm.

ing the exhalant siphon, the left one bears 1-3 small

alee proj jections (sometime called “small tentacular fil-

aments’). The epipodium is simple, with four pairs of

epipodial tentacles; the tentacles are narrow and long,

highly contractile, with a micro-papillated surface; some-

times an additional left unpaired tentacle may be present

(Figures 19-20).

The foot has an anterior truncated end, expanded into

two short lateral hom-like projections, which are usually

visible also in preserved specimens (Figure 19). In Mar-

garella expansa the entire upper surface of the body is

pigmented black, while the ventral foot surface is cream.

The bipectinate ctenidium has a relatively long dorsal

afferent membrane that surpasses the transverse pallial

vein and connects the ctenidial axis to the mantle skirt.

Radula of Margarella species. 1. M. bouvetia: general morphology. 2. M. expansa: central and lateral teeth. 3-4.

3. M. steineni. 4. M. expansa. 5. M. violacea: detail of the cutting edge of central tooth. 6

detail of the cutting edge of a lateral tooth. Scale bars: Figure 1 =

6. M. steineni:

1 mm; Figures 2, 3 = 100 pm; Figures 4, 6 = 20 wm; Figure

Biological Observations: Specimens of Margarella

expansa were collected from subtidal to 50 m depth,

living mainly on blades and holdfasts of the brown alga

Macrocystis pyrifera, as well as on rocky or crushed shell

substrates.

Laboratory observations revealed that they are able to

move floating in upside-down position with the expand-

ed foot gliding onto the inner side of the air-water in-

terface. It was also observed that the foot is able to wrap

around algal blades and holdfasts.

Egg masses of Margarella expansa were collected

from blades of the kelp Macrocystis pyrifera in January

and November. The largest egg-mass (29 mm in diam-

eter) consisted of a ribbon with about 1600 eggs, mea-

suring 0.51+0.03 mm diameter (n = 33), which adhered

D. G. Zelaya, 2004

Page 115

Figures 7-14. Details of first inmermost marginal tooth in Margarella species. 7. M. expansa. 8. M. antarctica. 9. M. tropido-

phoroides. 10. M. violacea. 11. M. steineni. 12. M. achilles. 13. M. obsoleta. 14. M. bouvetia. Scale bars: Figures 7-14 = 100 wm.

to the blade surface via a gelatinous matrix, and were

arranged in a spiral pattern (Figure 29). Embryos de-

veloped within the egg capsule until the stage of crawl-

ing juveniles (no free-swimming stage).

Remarks: Margarella resembles the genus Margarites

in shell morphology and in some reproductive features

(both spawning benthic egg masses), but in the mor-

phology of the radula and details of external anatomy,

both genera clearly differ. The radula of all Margarella

species studied here consistently differs from that de-

scribed for Margarites, particularly the morphology of

the first marginal tooth, variously referred to as a “pro-

tolateromarginal plate”, “innermost lateral plate” or “la-

teromarginal plate”. Thiele (1929) and Hickman and

McLean (1990) recognized the morphology of this tooth

as a valuable diagnostic character to separate Margarites

and Margarella. From the present study it is clear that

in Margarella the innermost marginal tooth undoubtedly

corresponds to a protolateromarginal plate as defined by

Hickman and McLean (1990) (i.e., a tooth with a base

greatly enlarged laterally, but with a well-developed shaft

and cutting edge). Conversely, in Margarites this tooth

(a true lateral plate) is represented by an expanded plate

without cutting edge.

The constancy in the number of lateral teeth appears

to be an additional character that contributes to the def-

inition of Margarella. All Margarella species studied

here, as well as others for which the radula is known,

have five lateral teeth: Margarella refulgens (Eales, 1923;

Amaud, 1972a; Numanami, 1996), Margarella crebrili-

rulata (Arnaud, 1972a), Margarella gunnerusensis Nu-

manami, 1996 and Margarella whiteana Linse, 2002.

Conversely, in Margarites, the number of lateral teeth

varies: four in Margarites gigantea (Leche, 1878), five in

Margarites argentata (Gould, 1841), six in Margarites

helicinus (Phipps, 1774), Margarites beringensis (E. A.

Smith, 1899) and Margarites albolineatus (E. A. Smith,

1899), and seven in Margarites hickmanae McLean,

1984 (McLean, 1984; Hickman and McLean, 1990; War-

én, 1990). The lateral teeth in Margarites are usually

stronger and wider than in Margarella.

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THE NAUTILUS, Vol. 118, No. 3

Figures 15-18. Detail of marginal teeth in Margarella species. 15-16. Lateral view of the uppermost part of marginal teeth.

15. M. obsoleta: inner marginals. 16. M. expansa: outer marginals. 17. M. expansa: detail of lateral and basal portion of marginal

teeth. 18. M. expansa: detail of the cutting edge. Scale bars: Figures 15, 16 = 20 ym; Figure 17 =

Smith (1879), Pelseneer (1903) and Thiele (1906)

considered the asymmetry of the left and right neck

lobes as an additional diagnostic character for Marga-

rella. All the species studied here showed dimorphic

neck lobes.

The ctenidial morphology of southwestern Atlantic

species of Margarella is studied for the first time. All

species crenata had a ctenidium with a relatively long

afferent membrane, clearly different from that in Meme

garites, in which a short dorsal afferent membrane is

restricted to the region posterior to the transverse pallial

vein (Hickman and McLean, 1990).

100 um; Figure 15 = 10 pm.

Species of Margarella here studied, as well as the

Antarctic and GibL mien species M. crebrilirulata,

M. expansa, M. steineni, M. bouvetia, and M. achilles

studied by Smith (1879), Powell (1951), Arnaud

(1972b) and Numanami (1996), have four pairs of epi-

podial tentacles, while in Margarites the number of

epipodial tentacles ranges from oe to seven pairs (Kie-

ner, 1880; Hickman arn McLean, 1990). However, dur-

ing the present study specimens of Margarella expansa

frequently showed an additional left unpaired tentacle,

as previously reported by Thiele (1906) for M. violacea.

Throughout the present study, an additional unpaired

Figures 19-20. External anatomy of Margarella species. 19. M. violacea. 20. M. steineni. cl: cephalic lappets; ct: cephalic

tentacles; e: eye stalk; f: foot; h: antero- lateral expansions of the food (horn-like projections); 0: operculum; sn: snout; t: epipodial

tentacles. Scale bars: Figure 17-18 = 4 mm.

D. G. Zelaya, 2004

Page 117

Figures 21-28. Shell morphology of Margarella species. 21. Protoconch of M. expansa. 22. Shell structure of M. expansa. 23-

24. Two syntypes of M. violacea (BMNH 20020261). 25. Juvenile specimen of M. violacea. 26. Type specimen of M. jason (BMNH

1961348). 27. M. obsoleta. 28. M. tropidophoroides. Scale bars: Figures 21, 25 = 1 mm; Figure 22 = 100 ym; Figures 23, 24, 26

= 5 mm; Figures 27, 28 = 3 mm.

tentacle was also found in M. steineni and M. antarc-

tica. This observation might explain the discrepancies

in the number of epipodial tentacles reported by Pow-

ell (1951), Arnaud (1972a), and Numanami (1996) for

Margarella antarctica, M. refulgens, and M. gunneru-

sensis. Similarly, in Margarella refulgens, Arnaud

(1972b) reported five and Numanami (1996) four pairs

of epipodia. A similar variability in the number of epi-

podial tentacles is known in the Antarctic trochid An-

timargarita dulcis as reported by Eales (1923) and

Margarites groenlandica by Kiener (1880).

Moreover, the foot shape forming lateral horn-like

projections during locomotion and the ability to enroll

the food around algae in Margarella are characters not

reported for Margarites (Hickman, 1996).

DISCUSSION

Several authors have misinterpreted the morphology of

the first marginal tooth in some species from the South-

erm Ocean (i.e. Margarella expansa, M. antarctica, M.

bouvetia, M. refulgens), describing it as a lateromarginal

plate and, consequently, reporting them under the genus

Margarites. In the present paper it was possible to re-

study and reinterpret from SEM images the morphology

of the radula of Margarella expansa, M. antarctica, and

M. bouvetia, which were previously described by Powell

(1951), Deambrosi (1969), and Troncoso et al. (2001)

from light microscope slide preparations. In all these

species, as well as it was observed in this study for M.

achilles, M. obsoleta, M. steineni, M. tropidophoroides,

Page 118

THE NAUTILUS, Vol. 118, No. 3

Figure 29. Egg masses of Margarella violacea. Scale bar: 10

mm.

and M. violacea, the morphology of the first marginal

tooth clearly corresponded to that of a protolateromar-

ginal plate. It is evident that discrepancy with previous

workers is a consequence of the observational technique

used. The Margarella radula is very difficult to interpret

from light microscope preparations (a technique used

mostly in the past) due to the high number of teeth and

the high degree of juxtaposition. Numanami (1996) also

reported Margarella refulgens as Margarites, althought

it is clear that the morphology of the fr st marginal tooth

also corresponds to a protolateromarginal plate (Numan-

ami, 1996, fig. 12F).

The present paper reveals that previous records of the

genus Margarites (Margaritinae) from the southwestern

Atlantic Ocean originated through misinterpretation of

the radular characters. The evidence discussed here

strongly suggests that the Antarctic species referred to

the genus Margarites actually belongs to Margarella.

OTHER GENUS-LEVEL NAMES APPLIED TO MARGARELLA

As it was mentioned above, southwestern Atlantic and

Antarctic species of Margarella have been frequently re-

ported under Margarites. Margarita, a name under

which some southern species of Margarella were also

reported was shown to be a junior synonym of Marga-

rites (Keen, 1960; Quinn, 1979); similarly Valvatella

Gray, 1857, was wrongly used, being presently consid-

ered a subgenus of Margarites (Hickman and McLean,

1990).

Margaritella, proposed by Thiele (1891) to include

Margarella violacea and M. expansa, is a homonym of

Margaritella Schmidt, 1880, a genus of hexactinellid

sponges. Accordingly, Thiele (1893) proposed Margar-

ella as a replacement name.

Trochus (sensu lato) Linnaeus, 1758 was frequently

and ambiguously used in the old literature to refer to

gastropods with trochoid shells and presently included

in the families Trochidae and Turbinidae. Margarella ex-

pansa was described under Trochus (Photinula), a name

subsequently used by Smith (1879) and Watson (1886).

Trochus sensu stricto is the type genus of the subfamily

Trochinae, and is clearly different from Margarella

(Hickman and McLean, 1990).

Species of Margarella have also frequently been re-

ferred to Photinula H. Adams and A. Adams, 1854 (no-

men novum pro Photina H. Adams and A. Avdaans. 1853,

non Burmeister, 1838) (Ihering, 1902; Strebel, 1905,

1908). Photinula sensu stricto, however, is a member of

the Calliostomatinae, clearly differing from Margarella

in radular morphology, number of epipodial tentacles

and honeycomb-like sculpture of protoconch.

Promargarita was proposed by Strebel (1908) as a

subgenus rai Photinula (s.1.), but in the same work he

considered it as a genus-level name when describing

Promargarita tropidophoroides, which is actually a spe-

cies of Margarella. Powell (1951) and Keen (1960) con-

sidered Promargarita as a subgenus of Margarella that

includes the species from South Georgia Islands with

spirally ornamented shells: M. tropidophoroides, M.

achilles, and M. obsoleta; although, this shell character

is also present in Margarella whiteana from Antarctic

Peninsula, Antarctica (Linse, 2002). The validity of Pro-

margarita as a subgenus of Margarella will be discussed

in a future paper, in the context of a systematic revision

of the species of the genus.

Lamy (1911) wrongly identified specimens of a true

Margarella species, probably M. obsoleta, from South

Georgia Islands as Photinula (Kingotrochus) lahillei var.

carinata.

Submargarita (type species: S. impervia) was de-

scribed by | Strebel (1908), and a number of species were

described under Submargarita: S. notalis (Strebel,

1908), S. strebeli Thiele, 1912, S. similis Thiele, 1912,

S. mamillata Thiele, 1912, and S. unifilosa Thiele, 1912.

Thiele (1929) considered Submargarita a subgenus of

Margarella. Warén (1992) based on radular morphology

of S. notalis transferred all the above-mentioned Ant-

arctic species (including S. impervia) to Lissotesta Ire-

dale, 1915, provisionally allocating the genus to the fam-

ily Skeneideae. Furthermore, a true Margarella species,

M. crebrilirulata from Antarctica, was reported under

Submargarita by Thiele (1912).

THE SUPRAGENERIC PLACEMENT OF MARGARELLA

The higher taxonomy of the Trochidae is mainly based

on characters of the radula and external anatomy (Hick-

man and McLean, 1990). Hickman and McLean (1990)

proposed three informal clades between the family and

subfamily ranks, based on ctenidial morphology, recog-

nizing the radula as diagnostic at subfamily level, and

certain characters of the shell and epipodial structures,

as diagnostic at the tribe rank. Powell (1951) considered

the morphology of the epipodial tentacles as a valuable

character in defining genera within the Trochoidea.

The relatively long afferent membrane of the bipec-

tinate ctenidum, the M-shaped top to cusp rachidian

tooth and the presence of a protolateromarginal plate,

clearly place Margarella in the subfamily Trochinae, as

D. G. Zelaya, 2004

previously suggested by Hickman and McLean (1990),

rather than Margaritinae. The ability to wrap the foot

around algae is a behavioral character only reported as

present in Gibbulini, Trochini, and Cantharidini, but not

in other trochid groups, including Margaritini (Hickman,

1996). The relatively low-spire shell, with the aperture

lacking columellar denticles, and the presence of epi-

podial tentacles emerging from simple epipodial tissue,

as well as the presence of antero-lateral horn-like pro-

jections (according to Hickman (1996) the latter char-

acter is only present in the tribe Gibbulini) indicate that

Margarella belongs to tribe Gibbulini. However, other

behavioral character such as the living habit of Marga-

rella, commonly reported as living on brown algae, ei-

ther on blades or associated to their holdfasts (King and

Broderip, 1932; Arnaud, 1972b; Ojeda and Santelices,

1984; Castilla, 1985; present study) is not a typical fea-

ture among representatives of the Gibbulini (Hickman

and Mc Lean, 1990), such mode of life was reported for

austral Gibbulini of small size, especially in the genus

Notogibbula Iredale, 1924 (Hickman and McLean,

1990).

ACKNOWLEDGMENTS

I wish to express my gratitude to C. Ituarte for his help

and constructive suggestions; C. Hickman and B. Mar-

shall, journal reviewers, kindly provided valuable criti-

cisms that considerably contributed to improve the man-

uscript. Any remaining inaccuracies, however, are solely

the author's responsibility. I thank to L. Simone and R.

Bieler for literature and additional information. The au-

thor is a Doctoral Fellow of the National Research

Council for Science and Technology (CONICET), Ar-

gentina. This work was funded by a student grant from

Conchologist of America (COA).

LITERATURE CITED

Arnaud, P. M. 1972a. Invertebrés marins des XIJéme et XVeme

Expéditions Antarctiques Frangaises en Terre Adélie. 8.

Gastéropodes prosobranches. Tethys, supplément 4: 105—

134.

Arnaud, P. M. 1972b. Sur une petite collection de gastéropodes

prosobranches et pélécypodes de |’Ile Petermann (An-

tarctique). Tethys 4 (2): 429-436.

Castilla, J. C. 1985. Food webs and functional aspects of the

kelp, Macrocystis pyrifera, community in the Beagle

Channel, Chile. In: Siegfried W. R., P. R. Condy and R.

M. Laws (eds). Antarctic nutrient cycles and food webs.

Springer-Verlag, Berlin, pp. 407-414.

Deambrosi, D. F. de. 1969. La rédula como cardcter diagnés-

tico en los géneros Margarella y Margarita y nueva ubi-

cacién de algunas especies—(Mollusca). Acta Zooldégica

Lilloana 24: 49-55.

Eales, N. 1923. Mollusca. Part V. Anatomy of Gastropoda (ex-

cept the Nudibranchia). British Antarctic Terra Nova Ex-

pedition, Natural History Report, Zoology, 7 (1): 146.

Hickman, C. S. 1996. Phylogeny and patterns of evolutionary

radiation in trochoidean gastropods. pp. 177-198. In: J.

Page 119

D. Taylor (ed.) Origin and evolutionary radiation of the

Mollusca. Oxford University Press, Oxford, 392 pp:

Hickman, C. S. and J. H. McLean. 1990. Systematic revision

and suprageneric classification of trocheacean gastropods.

Natural History Museum of Los Angeles County, Science

Series 35: 1-169.

Thering, H. Von. 1902. Die Photinula-Arten der Magellan-

Strasse. Nachrichtsblatt der Deutschen Malakozoologisch-

en Gesellschaft 34: 97-104.

Kiener, L. C. 1880. Spécies général et iconographie des co-

quilles vivantes comprenant la collection du Muséum

d'Histoire Naturelle de Paris. Genres Calcar, Trochus,

Xenophora, Tectarius et Risella. J. B. Bailliere et Fils, Par-

is, 480 pp + 120 pls.

King, P. P. and W. J. Broderip. 1832. Description of the Cir-

rhipeda, Conchifera and Mollusca, in a collection formed

by the officers of H.M.S. Adventure and Beagle employed

between the years 1826 and 1830 in surveying the South-

em coasts of South America including the Strait of Ma-

galhaens and the coast of Tierra del Fuego. The Zoological

Journal 5 (19): 332-349.

Keen, M. 1960. Systematic descriptions (Archaeogastropoda).

In: R. C. Moore (ed.), Treatise on invertebrate paleontol-

ogy, ed., Part. I, Mollusca 1, 169-310. Lawrence, Kansas:

Geological Society of America and University of Kansas

Press, xxiii + 351 pp.

Lamy, E. 1911. Sur quelques mollusques de la Géorgie du Sud

et des iles Sandwich. Bulletin du Muséum d’ Histoire Na-

turelle 17 (1): 22-27.

McLean, J. H. 1984. New species of Northeast Pacific archeo-

gastropods. The Veliger 26: 233-239.

Numanami, H. 1996. Taxonomic study on Antarctic gastropods

collected by Japanese Antarctic research expeditions.

Memoirs of National Institute of Polar Research, Series

E (Biology and Medical Science) 39: 1-244.

Ojeda, F. P. and B. Santelices. 1984. Invertebrate communities

in holdfasts of the kelp Macrocystis pyrifera from southem

Chile. Marine Ecology Progress Series 16: 65-73.

Pelseneer, P. 1903. Mollusques (Amphineures, Gastropodos et

Lamellibranches). Results du Voyage du S.Y. “Belgica”.

Expedition Antarctique Belge 1897-1899, Zoologie 3, 85

pp. + 9 pls.

Picken, G. B. 1979. Non-pelagic reproduction of some antare-

tic prosobranch gastropods from Signy Island, South Ork-

ney Islands. Malacologia 19: 109-128.

Powell, A. W. B. 1951. Antarctic and Subantarctic Mollusca:

Pelecypoda and Gastropoda collected by the ships of the

Discovery Committee during the years 1926-1937. Dis-

covery Reports 26: 49-196.

Quinn, J. F. Jr. 1979. Biological results of the University of

Miami Deep-Sea Expeditions. 130. The systematics and

zoogeography of the family Trochidae collected in the

Straits of Florida and its approaches. Malacologia 19: 1—

62.

Reid, D. G. and GC. Osorio. 2000. The shallow-water marine

Mollusca of the Estero Elefantes and Laguna San Rafael,

southern Chile. Bulletin of the Natural History Museum,

London (Zoology) 66 (2): 109-146.

Smith, E. A. 1879. Mollusca. An account of the petrological,

botanical and zoological collections made in Kerguelen’s

Island and Rodriguez during the Transit of “Venus” Ex-

pedition in the years 1874-75. Philosophical Transactions

of the Royal Society of London 168: 167-192.

Strebel, H. 1905. Beitriige zur Kenntnis der Molluskenfauna

Page 120

THE NAUTILUS, Vol. 118, No. 3

der Magalhanaen-Provinz. 2. Die Trochiden. Zoologische

Jahrbiicher. Supplement 8: 121-166.

Strebel, H. 1908. Die Gastropoden. Wissenschaftliche Ergeb-

nisse der Schwedischen Siidpolar-Expedition, 1901-1903

unter Leitung von Dr Otto Nordenskjold 6 (1): 1-112,

Is.1-6.

Thiele, J. 1891. Das Gebiss der Schnecken zur Begriindung

einter Nattirlichen Classification. Untersuch von Professor

Dr. F. H. Troschel, Fortgesetz von Dr J. Thiele, vol. 2,

part 7. R. Sricker, Berlin, pp. 249-334, plates 25-28.

Thiele, J. 1893. Das Gebiss der Schnecken zur Begriindung

einter Natiirlichen Classification. Untersuch von Professor

Dr. F. H. Troschel, Fortgesetz von Dr J. Thiele, vol. 2,

part 8. R. Sricker, Berlin, pp. 337—409, plates 29-32.

Thiele, J. 1906. Bemerkung iiber die Gattung Photinula. Nach-

richtsblatt der Deutschen Malakozoologischen Gesell-

schaft 38: 12-15.

Thiele, J. 1912. Die Antarktischen schnecken und muscheln.

Deutsche Siidpolar Expedition 1901-1903, 13: 183-285.

Thiele, J. 1929. Handbuch der systematischen Weichtierkunde.

Gustav Fischer Verlag, Jena. Teil 1, pp. 1-376.

Troncoso, N., J. L. Vangoethem and J. S. Troncoso. 2001. Con-

tribution to the marine molluscan fauna of Kerguelen Is-

lands, South Indian Ocean. Iberus 19: 83-114.

Warén, A. 1990. Ontogenetic changes in the trochoidean (Ar-

cheogastropoda) radula, with some phylogenetic interpre-

tations. Zoologica Scripta 19 (2): 179-187.

Warén, A. 1992. New and little known “Skeneimorph” Gastro-

pods from the Mediterrean Sea and the adjacent Atlantic

Ocean. Bolletino Malacologico 27 (10-12): 149-248

Watson, R. B. 1886. Report on the Scaphopoda and Gastero-

poda collected by the H.M.S. Challenger during the years

1873-76. Report of the Scientific Results of the Voyage

H.M.S. Challenger during the years 1873-76 under the

command of Captain George S. Nares, R. N., F. R. S. and

the late Captain Frank Tourle Thomson, Zoology 15 (41):

608 pp: (42): 609-756, pls. 1-50.

THE NAUTILUS 118(3):121-126, 2004

Page 121

Comparative resistance to starvation among early juveniles of

some marine muricoidean snails

Carlos S. Gallardo

Cristian Manque

Marcela Filtin

Instituto de Zoologia, Universidad

Austral de Chile

Casilla 567, Valdivia

CHILE

[email protected]

ABSTRACT

Trophon geversianus is a muricid snail in which juveniles show

abundant internal yolk reserves, as well as a 2-3 mm size at

hatching which is double that of most of the species within the

family. It is thus possible that the resistance of these juveniles

to the effects of food deprivation would be significantly greater

than that typical of other species in the family. This hypothesis

was evaluated experimentally by studying the tolerance of these

juveniles to starvation in comparison with juveniles of Chorus

giganteus and Acanthina monodon, which represent the more

generalized characteristics found among the muricoideans. Re-

cently hatched snails of the three species were established in

aquaria with flow-through seawater and no food sources. Three

replicate systems containing 100 individuals each were run.

Mortality rates and growth in columellar length were evaluated

over an experimental period of 135 days. The results showed

a clear survival advantage in T. geversianus under food depri-

vation when compared with the other species. This species also

showed some growth over the first 45 days of observation. Sur-

vival of this species was nearly 90% at 45 days, with mortality

increasing at 60 days and about 20% survival present at 105

days, with a few even surviving after 4 months without food.

Chorus giganteus and A. monodon juveniles demonstrated an

earlier, continuous mortality, with only 50% survival at 45 days,

and 10% survival at 75 days, with none surviving beyond 90

days. There was no increase in size in these juveniles during

the study period. The high resistance to fasting in T. geversi-

anus juveniles appears due to their high levels of energetic

reserves, obtained from high levels of intracapsular extra-em-

bryonic nutrients. This is interpreted as an adaptation to critical

seasonal conditions in their natural habitat during their first

months of life in the Straits of Magellan.

INTRODUCTION

Benthic marine invertebrates might suffer mortalities of

over 90% as juveniles or in their first days of life as they

settle into the benthos, as documented by Thorson

(1966) in a number of specific cases. More recently, Gos-

selin and Qian (1997) summarized information on di-

verse invertebrates, which confirmed an exponential

drop in juvenile survival during the first days or weeks

post-settlement, with less than 20% survival after four

months of life in the benthos. This pattern was also ob-

served by Moran (1999). Attention has been centered

on environmental factors, both biotic and abiotic, in de-

termining early mortality in marine organisms as they

initiate post-settlement life in the benthos. When mor-

tality factors persist, however, under various environ-

mental conditions, uniformity in mortality of recently

settled organisms suggests an intrinsic factor producing

a common vulnerability to mortality in these organisms

(Gosselin and Qian, 1997). Therefore, some of their

structural and functional attributes at the time of settle-

ment need to be ascertained, as these may be included

among the factors producing vulnerability in recently

settled organisms.

Among these noteworthy intrinsic determinants of

vulnerability are their post-settlement size and growth

rates upon starting benthic life, as well as the amounts

of energy reserves with which they are supplied in this

critical stage of their lives. The gradual leveling off of

juvenile survival in this early phase is a result of their

reaching critical sizes at which their vulnerability to en-

vironmental stress is substantially reduced. Unpredict-

able critical conditions in the environment which may

impede these organisms from reaching optimal growth

and size for survival may be overcome by the presence

of adequate energy reserves in the juveniles at settle-

ment. As a result, evolution has produced initial juvenile

sizes and levels of reserves that may be very diverse and

varied among species within a given taxon. Thus, em-

bryonic encapsulation, which is characteristic of many

marine gastropods, seems to have opened new evolu-

tionary possibilities for the regulation of both optimal

sizes of individuals and the nutritional reserves among

early juveniles within given cohorts, according to the en-

vironmental conditions present in each specific habitat.

Spight’s (1976) analysis of data available for muricid

snails concluded that probabilities of survival among ju-

veniles increased with growth, where there was an op-

timal size at hatching determining the corresponding

size-specific survival. Selection for larger size at hatching

favored survival under environmental stress, reduced

problems of dehydration, lowered susceptibility to pre-

dation, and provided greater tolerance to temporary con-

ditions of starvation. According to Spight (1976) an op-

timal hatching size of about 1 mm is prevalent among

muricoidean snails with direct development. However,

the regularity of this pattern is clearly altered among

muricoideans on the coast of Chile, specifically among

those at extreme southern latitudes as in the case of the

Magellanic species of Trophon. These juveniles are be-

tween 2.1 to 3.0 mm at hatching (Santana, 1998; Zaixso,

1973), which is more than double the size at hatching

and settlement observed in this family as predicted by

Spight (1976). This condition is achieved through the

unusual mechanism of extra-embryonic feeding on dis-

aggregated nutritive eggs and albuminous fluid durin

intracapsular development (Penchaszadeh, 1976; Santa-

na, 1998). Microscopic inspection of recently hatched

Trophon juveniles has shown large relative size (Santana,

1998; Zaixso, 1973) and content of exceptionally high

levels of nutritional reserves, indirectly suggesting the

potential for survival over long periods of starvation. In

order to test this prediction the present study experi-

mentally examined the capacity for survival and growth

of Trophon geversianus (Pallas, 1769) maintained in the

absence of food. The response is compared with parallel

observations on two Chilean species in the same family,

the snails Chorus giganteus (Lesson, 1829) and Acan-

thina monodon (Solander, 1786) whose juveniles, al-

though fed through the typical intracapsular mechanism

of provision of nutritive eggs, have a size at hatching of

1 mm, which fits the predicted pattern for snails in this

family.

MATERIALS AND METHODS

ORIGIN AND HANDLING OF EGG CAPSULES

Egg capsules of Chorus giganteus and Trophon gever-

sianus were obtained from adults acclimated in the Ma-

rine Culture Center of the Universidad de Los Lagos

(CEACIMA) located on Metri Bay (41°36’ S, 72°42’ W).

Capsules from Acanthina monodon were obtained from

the intertidal zone at Mehuin (39°25’ S, 73°13’ W) be-

tween July and August 2002.

Once obtained aad separ: ated by species, the capsules

were placed into culture in the CEACIMA laboratory,

Metri, in order to obtain post-hatching juveniles of each

species for experimentation. The egg capsules were

maintained in a flow-through seawater bath in cylindrical

tubes 15 cm in diameter by 20 cm high, having tops,

bottoms, and side ports covered with nylon screening

(600 wm mesh for C. giganteus, A. monodon, and 1000

wm mesh for T. geversianus). In advanced stages of mat-

uration, the capsules were cleaned manually every week

to eliminate fouling and accumulation of sediment.

THE NAUTILUS, Vol. 118, No. 3

EXPERIMENTAL DESIGN

Once hatching began, the juveniles obtained were main-

tained in the containers until the numbers required for

the experiments had accumulated. One hundred individ-

uals of each species were selected randomly and depos-

ited into new containers, with three replicates per spe-

cies. These containers were plastic, with screening both

on the walls and over the upper part (top), and arranged

in a culture system receiving a constant flow of 30 m-

filtered seawater at about 2 L/min, at ambient seawater

temperature. Experimentation was carried out from July

2002 to February 2003. The water temperature in the

experimental tanks during the experimental period

ranged from 10.5°C (Austral winter) to 11.8°C (spring),

reaching a high of 16°C in December and January (sum-

mer). The three species were exposed to the same tem-

perature regime in parallel throughout the experiments.

In order to comparatively estimate the tolerance of the

juvenile snails of the three species to fasting conditions,

measurements were made on survival and growth of the

individuals. For this, the number and mean size of the

hatched individuals of each species were recorded ini-

tially. Observations were made every 15 days, recording

survival in each replicate and the mean value by species,

as well as measuring the size of each individual. The

fragile snails were handled with a fine camel-hair brush,

fine dissecting forceps, Pasteur pipettes, always within

Petri dishes kept on a cold surface so as to reduce any

stress due to manipulation and unusual temperature

changes. Evidence of mortality was based on the pres-

ence of empty shells or shells containing decaying tissue

residues. Growth of the snails was determined by mea-

suring the columellar length of each living specimen us-

ing an ocular micrometer in a stereoscopic microscope.

A daily temperature record was maintained in the lab-

oratory tanks where the experiment took place.

The ideal experimental design for this study would

have included a control set of each of the three species

of snails maintained under the same conditions as those

above, but with abundant food supplies available. In this

way, the laboratory-expected mortalities of fed speci-

mens could have been subtracted from those obtained

for the fasting specimens. This arrangement was impos-

sible to deploy because of the logistical difficulties in

obtaining the large numbers of post-hatching snails re-

quired, which included having them in the laboratory in

the same season of the year in order to be able to expose

the three species to uniform conditions of seawater tem-

perature, oxygen content, photoperiod, and other envi-

ronmentally dependent factors in the experimental

tanks. Based on our previous experience in culturing C.

giganteus, A. monodon, and, more recently, T. geversi-

anus, mortalities of fed individuals was low, and we as-

sumed a priori that it would not be a factor in the com-

parative observations among mortalities of the three spe-

cies in the above described experiment.

Figure 1. Post-hatching juveniles of T. geversianus. Note the

important nutrient reserve visible through the apical region of

the shell indicated by the arrow.

C. S. Gallardo et al., 2004

Figure 1. Post-hatching juveniles of T. geversianus. Note the

important nutrient reserve visible through the apical region of

the shell indicated by the arrow.

STATISTICAL ANALYSES

In order to carry out an analysis of the eventual growth

exhibited by the three species, a stage was selected in

which the survival was above the population minimum

(mean “n”), which could be statistically evaluated as a

representative sampling size. In this case, the stage was

represented by the first 45 days of the experimental

treatment.

Survival and growth in the experimental treatments

were examined using analysis of variance (ANOVA; Sokal

and Rohlf, 1981). For determination of the significance

between differences observed we employed an a poster-

iori Tukey test. All the analyses were carried out using

a significance level of 5% (P < 0.05). Computer statis-

tical packages for these tests were the SYSTAT 9 ® and

Microsoft Excel 97.

RESULTS

MorPHOLOGICAL EVIDENCE OF NUTRITIONAL RESERVES

IN POSTMETAMORPHIC JUVENILES

A considerable quantity of yolk reserves may be ob-

served through the translucent globose apical region of

the shell of post-metamorphic juvenile T geversianus

(Figure 1). This material occupies the digestive gland

(“hepatopancreas”) region, producing a yellow-orange

mass visible like large transparent cells. Similar reserves

in juveniles of the other species, as observed under the

microscope, appear to be markedly smaller than those

of the Trophon.

SURVIVAL RATES DURING FASTING

Results of the laboratory observations presented in Fig-

ure 2 and corresponding analysis of variance, clearly

demonstrate the greater tolerance and survival in juve-

niles of T geversianus under fasting conditions when

Page 123

100

cal

80-4

70+ GC. giganteus

$ 60+ GA. monodon

2 OT. geversianus

3) 505)

12)

eS S|

1S mM) a

T. geversianus

Time of Control (Days)

Figure 2. Comparison of survival rates for post-metamorphic

juveniles of C. giganteus, A. monodon, and T. geversianus sub-

mitted to fasting under experimental conditions over successive

time periods.

compared with the data from C. giganteus and A. mon-

odon under the same experimental treatment. Within T.

geversianus, the mortality observed in the first 15 day

stage was relatively low, at 5-11%; survival continued to

be very stable up until day 45. Mortality continued to

be more intense and continued to rise after the 2-month

measurement. Indeed, at this point the survival was 67%

of those starting in the experiment. Over 20% of these

snails survived to day 105, and the last few individuals

survived fasting past four months of observation.

The C. giganteus and A. monodon juveniles demon-

strated a markedly lower tolerance to fasting, and a par-

allel relationship, with no significant differences detect-

ed by the ANOVA employed among mortality rates

throughout the experiment. Beginning with the first 15-

day period, the survival of these species was significantly

lower than that of Trophon, with an intensity that per-

sisted over the subsequent observational periods. There

was a greater deterioration in survival capacity in fasting

beginning in the early weeks, with no stabilization of

mortality as observed in Trophon lasting to 45 days. By

45 days, about 50% of both the C. giganteus and A.

monodon populations had died, while at 75 days they

showed less than 10% survival. Neither of these species

survived past 90 days in the experiment.

GrowTH RATES

Together with showing comparatively better survival

than the other two species during fasting, the T. gever-

sianus juveniles also showed significant growth in the

sizes of their shells during their period of survival under

the experimental conditions (Figure 3). These juveniles,

beginning at hatching with a mean columellar length of

2.2 mm, showed a significant increase in length in the

ca. 50% of the survivors at 45 days (ANOVA, Table 1).

After this period there was no significant growth. In con-

trast, neither the juveniles of C. giganteus, nor those of

Page 124

THE NAUTILUS, Vol. 118, No. 3

IC. giganteus

A. monodon

OT. geversianus

Shell length (mm)

1 15 30 45 60 75 90 105 120

Days of development

Figure 3. Mean length of post-metamorphic juveniles of C.

sicanteus, A. monodon, and T. geversianus submitted to ex-

g » ee :

perimental fasting over successive ime periods.

A. monodon showed any significant shell growth during

the time they were maintained without food.

DISCUSSION

The results of the present study clearly confirm the

greater viability and survival capacity of post-metamor-

phic juveniles of T. geversianus in absolute absence of

food, in comparison with the other two muricids studied.

The high resistance to starving allowed high rates of sur-

vival for at least two months, with survival declining

gradually after this period. To the initiation of this de-

cline may have been added the stress of increased en-

ergy expenditure due to the expected rise in the water

temperature in December and January. It is possible that

within its natural distributional range, the viability of ju-

veniles of T. geversianus (attributable to their levels of

energy reserves at hatching) may be greater than that

shown in our experiments. The high degree of survival

of these juveniles is comparable to, or even greater than

that reported for newly hatched Nucella emarginata,

which may survive 50—120 days under fasting conditions

(Gosselin and Chia, 1994). This species has extra-~em-

bryonic nutrition in the form of nutritive eggs during its

embryonic development.

There are at least two properties or attributes in the

development of juveniles of T. geversianus which may

account for its greater resistance to fasting under exper-

imental conditions, including; (a) Large size at hatching

of the juveniles (as occurs in T. geversianus) which is

nearly double that of the other two species, and which

is typical of many species in this family (Spight, 1976)

and (b) the accumulation of abundant energy reserves

due to the existence of a special extra-embryonic nutri-

tive mechanism which maximizes the assignation of

raised levels of reserves to each juvenile hatching from

the capsules.

Regarding the first attribute, the greater size at hatch-

ing of the juveniles of a species, as occurs in T. gever-

sianus, increases their relative survival capacity. It is well

known that relative body size has a direct metabolic re-

lation to standard metabolic rate in poikilotherms (Pe-

ters, 1983; Schmidt-Nielsen, 1984) such that the meta-

bolic cost is greater per unit tissue in smaller organisms

than in those of larger size. Spight (1976) noted that an

advantageous attribute for survival conferred by larger

juvenile size in muricoideans, was that these showed lon-

ger periods of resistance to fasting given that in these

conditions larger individuals respired less per unit body

weight thus prolonging the duration of their nutritional

reserves. The size at hatching of T. geversianus is among

the largest known for the Muricidae, similar only to that

of Murex senegalensis (with similar numbers of nutritive

eggs per embryo) and Murex angularis (sensu Knudsen,

1950), although there are no data available on the effects

of fasting in these species useful in comparison with

present results.

The large accumulation of nutritional reserves ob-

served in the morphological examination of T. geversi-

anus post-hatching juveniles is a characteristic of the

species that readily distinguishes it from the other spe-

cies presently studied. This attribute was recognized in

previous studies describing the intracapsular develop-

ment of this species (Zaixso, 1973), and specifically note-

worthy is the accumulation of yolk reserves, which can

be seen through the apical, translucent sector of the

shell. This accumulation allowed characterization of this

species as a muricoidean snail whose juveniles had ex-

ceptionally high levels of nutritional reserves at the time

of their hatching and initiation of life in the benthos.

The embryos are provided with abundant food and nu-

tritional reserves prior to hatching by prolonged inges-

tion of disaggregated yolk in the albuminoid fluid of the

egg capsule throughout their long developmental period.

Table 1. Mean size as shell columellar length in postmetamorphic C. giganteus, A. monodon, and T. geversianus subjected to

f=)

fasting in successive measurement periods. Analysis of variance (ANOVA) is included for growth in length, as well as the index of

relative importance (Tukey = 0.05). Not significant at P > 0.05; significant at P < 0.05.

Time Length (mm) by species ANOVA

(days) C. giganteus n A. monodon n T. geversianus n F Pp Tukey

1 1 + 0.130 300 0.978 + 0.110 300 2.209 + 0.247 300

15 1.008 + 0.136 220

30 1.021 + 0.134 183

45 1.049 0.124 147

0.985 = 0.117 205

0.992 0.113 169

1.018 + 0.107 146

2.264 £ 0.225 278

2.472 + 0.421 268 9.589 0.014

2.879 + 0.434 267

717.410 0.000

470.929 0.000

CG. S. Gallardo et al., 2004

Data on intracapsular development of the species (Car-

celles, 1947; Zaixso, 1973; Santana, 1998) suggest there

are large numbers (900-2000) of nutritive eggs depos-

ited in each capsule that disaggregate at an early stage

and become mixed with the albumin to form a nutritive

suspension (deutolecital material). About 30 to a maxi-

mum of 45 embryos per capsule are fed by this material

for an intracapsular period of about four to five months;

they measure 2.13 to 3.9 mm at hatching, according to

the authors cited above. Thus the embryos may ingest

at least 30—50 nutritive eggs each, accompanied by un-

determined amounts of the albuminoid mass. This sug-

gests a high degree of nutrient intake which is much

greater than that estimated for C. giganteus and A. mon-

odon with a mean of 10-13 mania eggs per embryo

(Gonzalez and Gallardo, 1999; Gallardo, 1979: and re-

cent unpublished data). Uptake of albumin by T. gev-

ersianus may represent an important nutrient uptake as

found by Rivest (1986) for Urosalpinx cinerea, the em-

bryos of which, not having nutritive eggs, are fed by the

capsular fluid that contains important nutrients in some

species (Bayne, 1968; De Mahieu et al., 1974; Bramach-

ary, 1989).

Some specific characteristics of the environment oc-

cupied by T. geversianus may explain some of the selec-

tive pressures that favor its size at hatching and high

content of yolk reserves. The high resistance to fasting

conferred by these attributes without doubt increases

the survivability of these juveniles in situations of limited

food availability or in conditions where their access to

food is restricted in some way upon initiation of their

free-living existence. This may represent selection ori-

ented to maximize early juvenile survival under unfavor-

able conditions prevalent in the natural habitat occupied

by these snails after hatching. This is supported by field

observations carried out in the Magellan Straits, which

is the normal habitat of the species. Periodic surveys for

egg-capsule deposition at low tide in the area have dem-

onstrated the occurrence of an extensive spawning pe-

riod in spring-summer, with a long intracapsular devel-

opmental period that retards hatching of the juveniles

to late summer, with most hatching occurring in the fall

(Santana, 1998). Thus the first months of life of these

juveniles occur during unfavorable fall conditions, and

when climatic conditions are extreme during the follow-

ing winter. Intertidal areas remain exposed at very low

temperatures and probably suffer high degrees of des-

iccation during aerial exposure at low tide.

Although juveniles of C. giganteus and A. monodon

fit the general pattern of many muricoideans by mea-

suring about 1 mm in length, their tolerance to fasting

places them at levels that may be considered interme-

diate to relatively important among the few cases in this

family that have been studied (Gosselin and Qian, 1997).

The above-cited case of Nucella emarginata, whose em-

bryos are also supplied with nutritive eggs, show an im-

portant resistance to food deprivation. It should be not-

ed here that muricids having juveniles of similar size (ca.

1 mm) but no provision of extra-embryonic nutrition

Page 125

(e.g. nutritive eggs), survive for very short periods when

fasting. In Urosalpinx cinerea, for example, the juveniles

exhibit direct development without the benefit of nutri-

tive eggs (Spight, 1976), and only survive a week or less

in the absence of food (Rittschof et al., 1983). It is very

interesting to explore this type of relation with examples

from other species inasmuch as this might reveal the

possible selective advantages implicated in the evolution

of new nutritional strategies for embryos such as nurse

egg feeding. Such strategies function in the delicate

equilibrium between optimal size at hatching and max-

imization of the levels of nutrient reserves that the ju-

veniles of different species require for survival of ex-

treme conditions during their early benthic life.

The great similarity shown between juveniles of C.

giganteus and A. monodon relative to survival when fast-

ing is reflected in the similarity between their mecha-

nisms and levels of provision of nutritive eggs during the

intracapsular development of their embryos (Gallarder

1979: Gonzélez and Gallardo, 1999). Thus, comparison

between the intracapsular development of the two spe-

cies shows a strict similarity indicative of clear differ-

ences with the intracapsular conditions shown by other

muricids of the region, which supports the hypothesis of

a common ancestral origin in the early ontogeny of the

two species presently studied (Gallardo, in prep.). The

results also support the hypothesis that the levels of nu-

tritional reserves in the juveniles are similar between the

two species, explaining their evolution-derived similarity

in survival and growth rates under experimental condi-

tions of severe food limitation.

ACKNOWLEDGMENTS

This study was financed by Project DID-UACH 2001-

02 to the first author. Our thanks also to Universidad de

los Lagos and members of CEACIMA (Metri) for pro-

viding facilities at this marine laboratory.

LITERATURE CITED

Bayne, C. J. 1968. Histochemical studies on the egg capsules

of eight gastropod molluscs. Proceedings of the Malaco-

logical Society of London 38: 199-212.

Brahmachary, R. L. 1989. Mollusca. In: K. G. Adiyodi and R.

G. Adiyodi (eds.) Reproductive Biology of Invertebrates,

Volume 4, Part A, Fertilization, Development and Paren-

tal Care. Oxford and IBH Publishing Co., New Delhi, pp.

1-41.

Carcelles, A. 1946. Observaciones sobre algunas especies ac-

tuales y fésiles de Trophon de la Republica Argentina. No-

tas del Museo de La Plata 11 (93): 59-89.

De Mahieu, G. C., P E. Penchaszadeh and A. B. Casal. 1974.

Algunos aspectos de las variaciones de proteinas y ami-

nodacidos libres totales del liquido intracapsular en relacién

al desarrollo embrionario en Adelomelon brasiliana (La-

marck, 1811) (Gastropoda, Prosobranchia, Volutidae). Ca-

hiers de Biologie Marine 15: 215-277.

Gallardo, C. S. 1979. Developmental pattern and adaptations

for reproduction in Nucella crassilabrum and other mur-

icacean gastropods. Biological Bulletin 157: 453-463.

Page 126

Gonzalez, K. A. and C. S. Gallardo. 1999. Embryonic and larval

development of the muricid snail Chorus giganteus (Les-

son, 1829) with an assessment of the developmental nu-

trition source. Ophelia 51: 77-92.

Gosselin, L. A. and F. S. Chia. 1994. Feeding habits of newly

hatched juveniles of an intertidal predatory gastropod, Nu-

cella emarginata (Deshayes). Journal of Experimental Ma-

rine Biology and Ecology 176: 1-13.

Gosselin, L. A. and P-Y. Qian. 1997. Juvenile mortality in ben-

thic marine invertebrates. Marine Ecology Progress Se-

ries. 146: 265-282.

Knudsen, J. 1950. Egg capsules and development of some ma-

rine prosobranchs from tropical West Africa. Atlantide Re-

port 1: 85-130.

Moran, A. L. 1999. Size and performance of juvenile marine

invertebrates; potential contrasts between intertidal and

subtidal benthic habitats. American Zoologist 39: 304-312.

Penchaszadeh, P. E. 1976. Reproduccién de gastrépodos pro-

sobranquios del Atlantico sur-occidental. El género Tro-

phon. Physis, Seccién A, 35 (90): 69-76.

Peters, R. H. 1983. The ecological implications of body size.

Cambridge University Press., Cambridge, 329 pp.

Rittschof, D., L. G. Williams., B. Brown and M. R. Carriker.

THE NAUTILUS, Vol. 118, No. 3

1983. Chemical attraction of newly hatched oyster drills.

Biological Bulletin 164: 493-505.

Rivest, B. R. 1986. Extra-embryonic nutrition in the proso-

branch gastropod Urosalpinx cinerea (Say, 1882). Bulletin

of Marine Science 39: 498-505.

Santana, M. 1998. Estudios sobre la época de desoves en la

naturaleza y desarrollo intracapsular en laboratorio del

caracol Trophon geversianus (Pallas, 1769) (Gastropoda:

Muricidae). Anales del Instituto de la Patagonia, Serie

Ciencias de la Naturaleza 26: 31-40.

Schmidt-Nielsen, K. 1984. Scaling: Why is Animal Size so Im-

portant. Cambridge University Press, Cambridge, 241 pp.

Sokal, R. R. and F. J. Rohlf. 1981. Biometry: The Principles

and Practices of Statistics in Biological Research. 24 Edi-

tion. W.H. Freeman, San Francisco, 859 pp.

Spight, T. M. 1976. Ecology of hatching size for marine snails.

Oecologia 24: 283-294.

Thorson, G. 1966. Some factors influencing the recruitment

and establishment of marine benthic communities. Neth-

erlands Journal of Sea Research 3: 267-293.

Zaixso, H. E. 1973. Observaciones sobre el desove y embrio-

logia de Trophon geversianus (Pallas) 1774. Neotropica

19: 156-162.

THE NAUTILUS 118(3):127-128, 2004

Page 127

Euglandina rosea (Férussac, 1821) is found on the ground and

in trees in Florida

Elizabeth C. Davis

Department of Ecology and

Evolutionary Biology

University of Kansas

1200 Sunnyside Ave.

Lawrence, KS 66045-7534 USA

[email protected]

Box 870345

Kathryn E. Perez

Department of Biological Sciences

University of Alabama

Tuscaloosa, AL 35487 USA

[email protected]

Daniel J. Bennett

Division of Entomology

Natural History Museum and

Biodiversity Research Center

Department of Ecology and

Evolutionary Biology

University of Kansas

1460 Jayhawk Blvd.

Lawrence, KS 66045-7523 USA

[email protected]

Euglandina rosea (Férussac, 1821) is a carnivorous snail

native to the southeastern United States, including Flor-

ida (Hubricht, 1985: Wolfe and Brooks, 1968). It is

known as a ground-dwelling terrestrial snail that feeds

on terrestrial gastropods (Burch, 1962; Pilsbry, 1946).

Introduced to Hawaii in 1955, it is one of the worst cases

of an intended biological control agent attacking non-

target species (Cowie, 2001). Exclosures keep it away

from consuming remaining endemic snails, including the

endangered Achatinella species (Stone, 1999).

On a collecting trip in Marianna, Jackson County,

Florida (June 2003), individuals of E. rosea were found

between 0.3 and 2.0 m above ground on both trees and

vines. The majority of individuals were found ~1 m

above ground (Figure 1). Collections were made after

heavy rains and in high humidity (>80% R.H.—Marian-

na Airport). Snails were located above the flood plain of

the Chipola River. They were found in a clearing within

a mixed forest dominated by salt cedar (Tamarix gallica

L.) and overgrown with vines.

Of the 17 living specimens of E. rosea collected, three

were found in the leaf litter. Most were found along the

trunks of trees or on the underside of leaves. Snails were

found immobile either attached with their foot or aes-

tivating, as determined by presence of a mucus epi-

phragm over their aperture (Burch, 1962). Euglandina

rosea were found within a 20-m? area, coexisting with

individuals of the potential prey snails: Mesodon thyro-

idus (Say, 1816), Stenotrema maxillatum (Gould, 1848),

Ventridens demissus (A. Binney, 1843), and Oligyra or-

biculata Say, 1818. Eggs of E. rosea were collected a few

centimeters below the surface of the leaf litter.

The literature suggests that E. rosea is found primarily

in leaf litter (Burch, 1962: Griffiths et al., 1993; Mace

et al., 1998; Pilsbry, 1946), although individuals have

been known to climb trees or walls to capture prey

(Cowie, 2001; Davidson, 1965; Gerlach, 1999; Hadfield

et al., 1993; van der Schalie, 1969: Voss, 1976). Davis

and Butler (1964) stated that E. rosea fed on tree snails

in its native habitat. However, these authors did not

mention that the introduction of E. rosea could threaten

native Hawaiian snails. In addition, E. rosea is known to

Figure 1. Euglandina rosea on vegetation in Marianna, Flor-

ida about 1 m above ground. Photograph by DJB.

Page 128

THE NAUTILUS, Vol. 118, No. 3

forage underwater for aquatic snails in Hawaii (Kinzie,

1992).

The behavior of aestivating above ground has serious

implications for the use of E. rosea as a biological control

agent. The ability of E. rosea to utilize both trees and

leaf litter may allow it to out-compete native species for

these resources and allow better access to non-target

species (Gerlach, 1999). The use of both ground and

tree habitat by E. rosea in its native range and aquatic

habitat in Hawaii (Kinzie, 1992) makes it a very danger-

ous species to use in biological control.

ACKNOWLEDGMENTS

We thank two anonymous reviewers, B. A. Wilson, and

Dr. C. Loudon for comments on the paper, the Florida

Fish and Wildlife Conservation Commission for permis-

sion to collect E. rosea, and the Conchologists of Amer-

ica research grant (to ECD), and the Entomology Pro-

gram of the University of Kansas for funding.

LITERATURE CITED

Burch, J. B. 1962. How to know the eastern land snails. Du-

buque, Iowa: WM. C. Brown Company publishers, 215

Coe R. H. 2001. Can snails ever be effective and safe bio-

control agents? International Journal of Pest Management

47(1): 23-40.

Davidson, T. 1965. Tree snails, gems of the Everglades. Na-

tional Geographic 127(3): 372-387.

Davis, C. J., G. D. Butler. 1964. Introduced enemies of the

giant African snail, Achatina fulica Bowdich, in Hawaii

(Pulmonata: Achatinidae). Proceedings of the Hawaiian

Entomological Society 28(3): 377-389.

Gerlach, J. 1999. The ecology of western Indian Ocean carniv-

orous land snails. Phelsuma 7: 14-24.

Griffiths, O., A. Cook and S. M. Wells. 1993. The diet of the

introduced camivorous snail Euwglandina rosea in Mauri-

tius and its implications for threatened island gastropod

faunas. Journal of Zoological Society of London 229: 79-

89.

Hadfield, M. G., S. E. Miller and A. H. Carwile. 1993. The

decimation of endemic Hawai‘ian tree snails by alien pred-

ators. American Zoologist 33: 610-622.

Hubricht, L. 1985. The distributions of the native land mol-

lusks of the Eastern United States. Fieldiana Zoology (24):

1-191.

Kinzie, R. A., III. 1992. Predation by the introduced camivo-

rous snail Euglandina rosea (Férussac) on endemic aquat-

ic lymnaeid snails in Hawaii. Biological Conservation 60:

149-155.

Mace, G. M., P. Pearce-Kelly and D. Clarke. 1998. An inte-

grated conservation programme for the tree snails (Par-

tulidae) of Polynesia: a review of captive and wild ele-

ments. Journal of Conchology Special Publication 2: $9—

96.

Pilsbry, H. A. 1946. Land Mollusca of North America (North

of Mexico). Monographs of the Academy of Natural Sci-

ences of Philadelphia 3(2.1): 1-512.

Stone, R. 1999. Keeping Paradise Safe for the Natives. Science

285(5435): 1837.

van der Schalie, H. 1969. Man meddles with nature—Hawaiian

style. The Biologist 51: 136-146.

Voss, R. S. 1976. Observations of the ecology of the Florida

tree snail, Liguus fasciatus (Miiller). The Nautilus 90: 65—

Wolte, D. A. and J. H. Brooks. 1968. A colony of Euglandina

rosea at Beaufort, North Carolina, Sterkiana 30: 23-24.

THE NAUTILUS 118(3);129-130, 2004

Page 129

Rediscovery of the syntypes of Doriopsilla pelseneeri D’Oliveira,

1895

Gonealo Calado!

Centro de Modelagao Ecolégica

IMAR, FCT/UNL

Quinta da Torre 2825-114, Monte da

Caparica

PORTUGAL

[email protected]

The nudibranch mollusk Doriopsilla pelseneeri

D Oliveira, 1895, was described by the Portuguese nat-

uralist Paulino D’Oliveira, from Sines (37°57 N, 8°53’

W), Portugal, in a paper concerning a number of opis-

thobranchs from his own collection (D’Oliveira, 1895).

There were no illustrations, but the detailed description

was sufficient to characterize this distinctive species. In

a recent revision of the Atlantic species of the genus

Doriopsilla, Valdés and Ortea (1997) considered D. pel-

seneeri as a valid name, and redescribed this species

adding new relevant morphological, anatomical, and

geographical data. Since the type material was untrace-

able, these authors designated a neotype, collected on

May 16, 1992, in Muros de Nalén, northern Spain, and

deposited at the Muséum national d'Histoire naturelle

(MNHN), Paris, France.

A visit to the Museu Zooldgico da Universidade de

Coimbra (MZUC), Portugal, after a major reorganization

of this institution, made it possible for me to locate Pau-

lino D’Oliveira’s opisthobranch collection. It includes

five syntypes of D. pelseneeri divided into two jars. The

larger one contains three specimens (lengths of pre-

served specimens 20 mm, 20 mm, and 21 mm), the

smaller jar contains the remaining two (lengths of pre-

served specimens 8 mm and 14 mm). Both jars are filled

with 70° ethanol. All the specimens bear the same ref-

erence number. An old number (160) is still present in

a label inside the larger flask, but a new classification of

the entire collection of the Museum was implemented

in 1945 (Carvalho, 1945), after which both jars were ex-

ternally labelled with the numeral “20”, which corre-

sponds to species number twenty of the “Order Acoela”

as stated in the published catalogue (Carvalho, 1945).

Hence, the reference number should be indicated as

MZUC-Acoela-20.

All specimens are well preserved and certainly were

relaxed prior to fixation. The 14 mm specimen has its

front partially destroyed, possibly due to the extraction

of the buccal bulb. A longitudinal incision between the

rhinophores is still visible. The other four remain intact.

The color of the preserved specimens is whitish but,

1 Address for correspondence: Instituto Portugués de Malacol-

ogia. Zoomarine, E.N. 125, km 65, Guia, 8200-864, Albufeira,

PORTUGAL

drawing from D’Oliveira’s description, the animals were

reddish or yellow when alive. Recently collected speci-

mens from Sines (personal data) confirm this assump-

tion. As pointed out by Valdés and Ortea (1997), in D.

pelseneeri the whole mid-portion of the dorsum is cov-

ered by large irregular tubercles, a feature that is not

present in any other Atlantic species of this genus. These

tubercles are easily recognizable in all the syntypes. One

of them is illustrated in Figure 1.

According to the fourth edition of the International

Code of Zoological Nomenclature (ICZN, 1999), Article

75.8, the rediscovered syntypes automatically become

the name-bearing types; the neotype should be set aside.

This substitution will cause neither confusion nor insta-

bility since the syntypes and neotype belong to the same

biological species.

ACKNOWLEDGMENTS

I thank Drs. Isabel Carreira and Rolanda Albuquerque

de Matos for the opportunity to examine the types of

Paulino D’Oliveira’s collection in MZUC. I hold a grant

from the Fundagao para a Ciéncia e Tecnologia, Portugal

(BPD7133/2001). This work was carried out under the

scope of the project REN2001-1956-C17-02/GLO

(Spanish Ministry of Science and Technology).

Figure 1. One of the syntypes of Doriopsilla pelseneeri

D Oliveira, 1895 (preserved length 20mm).

Page 130

THE NAUTILUS, Vol. 118, No. 3

LITERATURE CITED

Carvalho, R. N. 1945. Catélogo da colecgao de Invertebrados

de Portugal existentes no Museu Zoolégico da Universi-

dade de Coimbra. Memorias e Estudos do Museu Zoo-

légico da Universidade de Coimbra 167: 1-50.

D Oliveira, M. P. 1895. Opisthobranches du Portugal de la col-

lection de M. Paulino D’Oliveira. O Instituto 42: 574-592.

International Commission of Zoological Nomenclature (ICZN),

1999. International Code of Zoological Nomenclature, 3"

edition. International Trust for Zoological Nomenclature,

London, 306 pp.

Valdés, A. and J. Ortea. 1997. Review of the genus Doriopsilla

Bergh, 1880 (Gastropoda: Nudibranchia) in the Atlantic

Ocean. The Veliger 40: 240-254.

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CONTENTS

IN volte 05) iy BR is:

Volume 118, Number 4

December 21, 2004

ISSN 0028-1544

Néstor E. Ardila

Angel Valdés

Francisco J. Garcia

Jesus S. Troncoso

Pilar Carmona Zalvide

Victoriano Urgorri

Francisco J. Garcia

Richard Duerr

Geerat J. Vermeij

Greg S. Herbert

Luiz R. L. Simone

Diego G. Zelaya

Cristian Ituarte

The genus Armina (Gastropoda: Nudibranchia: Arminidae)

in the southern Caribbean, with the description of a new

SDE CLES ay cwei erin race Cenc mi peine nle mow Se Saale ce eat eCye creat 131

A new species of the genus Anetarca Gosliner, 1991

(Gastropoda: Opistobranchia: Facelinidae) from the

AVES ECTINPAC] ATI El @A@ COaTan Hey are Ren eer epee mew tr ea an ats cn ire 139

Two new species of Leptochiton Gray, 1847

(Polyplacophora) from the Iberian Peninsula (eastern

INTE TINE OXORGLE eee ts gary fescue 2 is leks Dok Tihs oh, Unga de LR seg Ra 144

Cirsotrema (Gastropoda: Ptenoglossa: Epitoniidae) in the

Miocene Chipola Formation of northwestern Florida. ................... 152

A new species of Stramonita (Gastropoda: Muricidae) from

ine Late Pitoeane OF IMGHCA.. .s0ccacccccsossoososdenes oo oben sooner 157

A new Orbitestella (Gastropoda: Heterobranchia:

Orbitestellidae) from Tierra del Fuego, Argentina....................... 160

Sphaeriidae (Bivalvia) from Peruvian Amazon floodplains,

with the description of Pisidium iquito new species..................... 167

Note

Paul Callomon

Dates of publication of Yochiro Hirase’s Kai Chigusa.................... 175

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THE NAUTILUS 118(4):131-138, 2004

Page 131

The genus Armina (Gastropoda: Nudibranchia: Arminidae) in

the southern Caribbean, with the description of a new species

Néstor E. Ardila

Museo de Historia Natural Marina de

Angel Valdés

Natural History Museum of Los

Colombia Angeles County

Instituto de Investigaciones Marinas 900 Exposition Blvd.

INVEMAR Los Angeles, CA 90007 USA

Santa Marta, A.A. 1016 [email protected]

COLOMBIA

[email protected]

ABSTRACT

The southern Caribbean species of the nudibranch genus Ar-

mina Rafinesque, 1814, are reviewed. Armina juliana Ardila

and Diaz, 2002, and Armina muelleri (Ihering, 1886), two pre-

viously described species, are redescribed and their reproduc-

tive systems illustrated. The two North American species Ar-

mina abbotti Thompson, Cattaneo and Wong, 1990, and Ar-

mina wattla Marcus and Marcus, 1967, are regarded as syno-

nyms of A. muelleri, which is reported from dhe Caribbean for

the first time. A new species, Armina elongata, is described

based on a single specimen collected from Colombia. The

number of notal ridges, the size and the shape of the radular

teeth, the features of the masticatory processes, and the mor-

phology of reproductive system are the main distinguishing

characteristics between the new species and other Atlantic spe-

cies of Armina.

INTRODUCTION

The Arminidae constitutes a poorly known group of liv-

ing nudibranchs, with nearly 75 described species divid-

ed into six genera (Kolb and Wagele, 1998). Arminids

arelcharacterized by having an elongated, flattened body,

narrower posteriorly, with longitudinal ridges or pustules

on the dorsum. The notum bears marginal sacs along its

edge. Arminids have a distinct oral veil and retractile

rhinophores with a caruncle situated next to them; the

radula typically has a broad, denticulated rachidian

tooth, and the lateral teeth are falciform (Kolb, 1998).

The majority of species in all other genera than Armina

Rafinesque, 1814 (Dermatobranchus van Hasselt, 1824,

Histiomena Morch, 1860, Linguella de Férussac, 1822

Pleurophyllidella Eliot, 1903, and Pleurophyllidiopsis

Tchang-Si, 1934) are geographically restricted to areas

in the Indo-Pacific (Kolb and Wigele, 1998).

Armina is the most species-rich genus of the family,

with more than 50 nominal species; it includes the most

derived forms of the group (Kolb and Wagele, 1998).

Features shared by members of this genus are a contin-

uous anterior mantle margin, the presence of branchial

and hyponotal lamellae, and the close distance between

the rhinophores. Although Armina has a world-wide dis-

tribution, only five species have been recorded in the

western Atlantic (see Marcus and Marcus, 1960: 1967;

Kolb and Wagele, 1998, Ardila and Diaz, 2002).

Armina juliana Ardila and Diaz, 2002, was the only

species previously known from the southern Caribbean.

The present paper describes two additional species of

Armina collected from materials trawled off the north-

ern Caribbean coast of Colombia.

MATERIALS AND METHODS

Specimens were collected using a bottom trawl (9X1 m

opening, 16 m length) during “the cruises INVEMAR-

MACROFAUNA I wand II, 1999-2001, aboard the R/V

ANCON, working off the Caribbean coast of Colombia at

depths of 20-500 m. Specimens of Arminidae were sort-

ed and preserved in 70% ethanol. Subsequently, jaws

and radulae were dissected and examined using a scan-

ning electron microscope (SEM). The specimens were

dissected for study of the reproductive system.

The material examined is deposited at the Museo Na-

cional de Historia Natural Marina de Colombia, INVE-

MAR (MHNMC) and the Natural History Museum of

Los Angeles County (LACM).

SYSTEMATICS

Family Arminidae Rafinesque, 1814

Genus Armina Rafinesque, 1814

Armina juliana Ardila and Diaz, 2002

(Figures 1-2)

Armina juliana Ardila and Diaz, 2002:

1-7.

27-30, text figs

Description: The living animals are bright red with

contrasting white longitudinal notal ridges; the anterior

Page 132

THE NAUTILUS, Vol. 118, No. 4

10 mm

Figure 1.

animal. B. Rachidian teeth. C. Lateral teeth. D

margin of the oral veil and the apical portion of the rhi-

nophores are also white (Figure 1A). Preserved speci-

mens are pale pink. The body is elongated, flattened,

narrowing posteriorly. The notum Tees 34 longitudinal

dorsal ridges, whose margins do not exhibit any black

spots of seamen pigment. The oral veil is small but

clearly distinct, with the lateral extension curved back-

wards and shorter than the widest part of the body (Fig-

ure 2A). There are two large club-shaped rhinophores,

each one with about 10 werteall lamellae. The rhino-

phores are situated dorsally on the notum. The eyes are

visible through the epidermis at the outer base of each

rhinophore. The caruncle is also distinguishable anterior

to the rhinophores. The genital opening is located on

the right side, anterior to and below the branchial la-

mellae (Figure 2B). The anal papilla is located slightly

behind the midpoint (3/5 of total length, from the an-

terior end of the body). There are twenty-one branchial

lamellae on the right side of the body and nineteen on

the left. Fifteen hyponotal lamellae are present on each

side of the body. A deep groove is present along the

center of the foot sole. The pedal gland is w he and

located at the posterior end of the fat sole (Figure 1A).

Mantle glands (each about 0.7 mm in diameter) are

Armina juliana Ardila and Diaz, 2002, holotype, MHNMC INV MOL1598. A. Dorsal and ventral views of the living

D. Jaws. E. Detail of the jaw elements on the masticatory processes.

barely distinguishable macroscopically on both sides just

behind the Rvemcliiell lamellae (Ardila and Diaz, 2002).

Jaws and Radula: The radular formula is 27 X 33.1.33

(holotype). The rachidian tooth is broad and bears 12

elongated denticles on each side of the median cusp

(Figure 1B). The first lateral tooth is short, bulky and

denticulated on the outer side of the cusp. The remain-

ing lateral teeth are elongated and also bear denticles,

which become progressively smaller towards the margin-

al tooth rows. The jaws have a denticulated masticatory

border with four rows of denticles (Figures 1D—E).

Reproductive System (Figure 2C): The reproductive

system is diaulic. The ampulla is very long, wide and

convoluted. The prostate is short and convoluted, com-

posed of two regions: a glandular, proximal region that

narrows into the distal, muscular ejaculatory region,

which connects directly to the penis. The vagina is short

and connected directly to the rounded bursa copulatrix.

Holotype (MHNMC INV MOL1598),

41 mm length alive, from type locality; Par atype (LACM

2908), off ‘Palonaiiaa, Colombia (11°26’ N, 73°32’ W),

306-312 m depth, muddy bottom, 14 mm length alive.

Type Material:

N. E. Ardila and A. Valdés, 2004

(aves

aes

a Se —J

a p aS)

Figure 2. Armina juliana Ardila and Diaz, 2002, paratype,

LACM 2908. A. Dorsal view of the anterior end of the body.

Scale bar as in B. B. Lateral view of the body. Scale bar = 1

mm. C. Reproductive system. Scale bar = 1 mm. Abbrevia-

tions: am, ampulla; ap, anal pore; be, bursa copulatrix; bl, bran-

chial lamellae; ca, caruncule; fg, female glands; gp, genital

pore; hl, hyponotal lamellae; le, Tateral extension of velum; n,

notum; pad, post-ampullary duct; pn, penis; pr, prostate; rh,

rhinophore; v, vagina; vl, velum.

Page 133

Type Locality: Off Cabo de La Vela, Colombia

(12°19' N, 72°42’ W), 460 m depth, muddy bottom.

Distribution: Off Cabo de La Vela and Palomino, Co-

lombia (Ardila and Diaz, 2002).

Remarks: This species was recently described in de-

tail by Ardila and Diaz (2002); a de scription is repeated

here to allow for quick comparisons with the other spe-

cies treated in this paper. Armina juliana can be easily

distinguished from other species here described by the

position of the genital papilla, which is located anterior

to the branchial lamellae, whereas in the other species

it is located just posterior to the lamellae. For a com-

prehensive discussion of A. juliana versus other Atlantic

species see Ardila and Diaz (2002).

Armina muelleri (Ihering, 1886)

(Figures 3-4)

Pleurophyllidia miilleri Thering, 1886: 223-228, pl. 9, fig. 1.

Armina abbotti Thompson, Cattaneo and W ong, 1990: 403.

Armina wattla Marcus and Marcus, 1967: 213-216, figs.

16—20. ‘

Description: The animal is red with contrasting yel-

low longitudinal notal ridges. The anterior margin Of the

oral veilland the edge of the notum is also yellow (Figure

3A). There are two hin yellow lines surrounding the foot

als and the external edge of the oral tentacles. Pre-

served specimens are pale grayish. The body is wide,

elongated, flattened, with a small notch on the posterior

tip. The notum bears 54 longitudinal dorsal ridges, with

wider, entire ridges, and Thera broken ridges inter-

calated. The oral veil is dlistiine? amd hes favo lane ge, ten-

es lateral extensions lacking any projections (Figure

4A). The extensions are as ratdle as the notum. amhere. are

two club-shaped rhinophores, each one with about 30

vertical lamellae. The rhinophores are situated in a notch

covered by the anterior end of the notum in the pre-

served specimens. The eyes are visible through the epi-

dermis at the outer base of each rhinophore. The car-

uncle is also distinguishable anterior to the rhinophores.

The genital opening is located on the right side of the

body, slightly posterior to and below the branchial la-

mellae (Figure 4B). The anal papilla is located behind

the midpoint (2/3 of total animal length, from the an-

terior end of the body). There are 23 branchial lamellae

and the same number of hyponotal lamellae on each side

of the body. The pedal gland is white and located at the

posterior end of the foot sole (Figure 3A). The mantle

edge is surrounded by a number of small mantle glands

distinguishable macroscopically on both sides.

Jaws and Radula: The radula formula is 36 X 35.1.35

in the single specimen examined. The rachidian tooth is

very broad and bears 3-4 large denticles on each side of

the median cusp (Figure 3B); the innermost denticle lies

very close to median cuspid (Figure 3C). The lateral

teeth are elongated and lack denticles (Figure 3D). The

jaws have a denttouleved masticatory border with four to

Eve rows of denticles (Figures 3E— F),

Page 134

THE NAUTILUS, Vol. 118, No. 4

Figure 3. Armina muelleri Thompson, Cattaneo and Wong, 1990 (MHNMC INV MOL3901). A. Dorsal and ventral views of the

living animal. B. Rachidian teeth, lateral view. C. Rachidian teeth, upper view. D. Lateral teeth. E. Jaw elements on the masticatory

processes. F. Jaws.

Reproductive System (Figure 4C—D): The reproductive

system is diaulic. The ampulla is very long, wide and

convoluted. There is no post- ampullary duct andl the am-

pulla connects directly to the prostate and the female

glands. The prostate is very long and convoluted, com-

posed of two regions: a glandular, proximal region that

narrows into the distal, muscular ejaculatory region,

which connects directly to the penis. The vagina is short,

irregular, and connected directly to the orale! bursa

copulatrix.

Type Material: Holotype (MHNMC INV MOL3901),

32 mm length alive, from type locality.

Type Locality: Off Salamanca Island, Colombia

(11°5'46"” N, 74°40'35" W), 20 m depth, hard bottom

with pennatulaceans (Renilla reniformis and Renilla

muelleri).

Distribution: This species is widespread in the North

America and the Caribbean, including records from

North Carolina, South Carolina, East Florida, West Flor-

ida, Texas, Mexico (Abbott, 1954; Eyster, 1981), and Co-

lombia (present study).

Remarks: Armina muelleri was originally described

from Brazil by Ihering (1886) as a species with a dark

notum covered with light yellow or pale brown ridges.

Marcus and Marcus (1960) re-described this species

based on specimens much lighter than the type material,

but collected from the same area. They argued that in-

traspecific variation was the cause of differences in color

between their material and the original type material.

Marcus and Marcus (1960) also conducted the first an-

atomical examination of this species. The anatomical de-

scriptions by Marcus and Marcus (1960) are very similar

to the material here examined. The radula has a rachi-

dian tooth with a central cusp and 3-4 strong denticles

on each side, and the reproductive system has a long

and convoluted ampulla that connects to an elongate and

curved prostate; the vagina is long and straight, connect

to an oval bursa copulatrix. itarsanellbe Botts specimens

are also very similar, having a wide and flattened body,

with small rhinophores emerging from below the ante-

rior end of the notum, and the genital opening situated

slightly posterior to and below “ine branchial lamellae.

The only consistent difference is the absence of denti-

N. E. Ardila and A. Valdés, 2004

Figure 4. Armina muelleri Thompson, Cattaneo and Wong,

1990 (MHNMC INV MOL3901). A. Dorsal view of the an-

terior end of the body. Scale bar = 1mm. B. Lateral view of

the body. Scale bar as in A. C. Reproductive system. Scale bar

= 1 mm. D. Detail of some reproductive organs. Scale bar as

in C. Abbreviations: am, ampulla; ap, anal pore; be, bursa co-

pulatrix; bl, branchial lamellae; ca, caruncule; fg, female glands;

gp, genital pore; hl, hyponotal lamellae; le, lateral extension of

velum; n, notum; pad, post-ampullary duct; pn, penis; pr, pros-

tate; rh, rhinophore; v, vagina; vl, velum.

cles on the inner lateral teeth of the Caribbean speci-

men. Attempts to locate the type material of Armina

muelleri failed, so the comparisons to our material are

solely based on the original description by Ihering

(1886), and subsequent re-descriptions by Ihering

(1915) and Marcus and Marcus (1960). The absence of

denticles on the inner lateral teeth of the Caribbean

specimen is probably due to variability within the spe-

cies, but the possibility that it belongs to an undescribed

species cannot be discarded. Since the rest of the anat-

omy and external morphology are identical to those of

A. muelleri, the Caribbean specimens is provisionally

placed in this species until more material becomes avail-

able.

The species name Armina abbotti was introduced by

Thompson, Cattaneo and Wong (1990) for the North

American specimens identified by Abbott (1954) and

Eyster (1981) as Armina tigrina Rafinesque, 1814. The

publication of the name is accompanied by bibliographic

references to descriptions and therefore it meets the re-

quirements of the International Code of Zoological No-

Page 135

menclature, ICZN (1999: Article 13.1.2). Examination of

the descriptions of specimens of A. tigrina in the papers

of Abbott (1954) and Eyster (1981), aad the original de-

scription of Armina muelleri and subsequent re- -descrip-

tions by Ihering (1915) and Marcus and Marcus (1960)

revealed that the external morphology and anatomy of

all these specimens are virtually identical, and there is

no doubt they belong to the same species. Again, the

type material of Armina abbotti is untraceable, so com-

parisons are based on the literature available. The radula

of the material studied by Eyster (1981) from South Car-

olina has a rachidian tooth with a central cusp and 4

strong denticles on each side, and the inner lateral teeth

bear some denticles, very similar to those of the speci-

mens from Brazil. The illustrations of the living animals

by Eyster (1981) show a species with a broad velum and

the rhinophores emerging ventrally to the anterior end

of the notum.

Nijssen-Meyer (1965) described one specimen of Ar-

mina from Surinam that he identified as belonging to

the tropical Indo-Pacific species Armina semperi (Bergh,

1861). This identification was based on the fact that the

specimen from Surinam had more rows of denticles in

the masticatory process than specimens of Armina muel-

leri and the innermost lateral teeth had numerous den-

ticles instead of a few or none. These differences in the

number of rows seem to be due to intraspecific variation.

The external morphology and the shape of the rachidian

tooth and the lateral teeth of the Surinam material are

identical to those of other references to A. muelleri

(Ihering, 1915; Marcus and Marcus, 1960). Thus the

specimen described by Nijssen-Meyer (1965) is here re-

garded as A. muelleri, concurring with the earlier sug-

gestion by Marcus and Marcus (1967).

A third species name introduced for the Western At-

lantic is Armina wattla Marcus and Marcus (1967), orig-

inally described from Georgia, USA. This species is Also

similar to Armina muelleri in radular and reproductive

morphology. Both species have wide rachidian teeth with

a few strong denticles on each side of the cusp and in-

nermost denticulate lateral teeth. The reproductive only

differs in the more elongate vagina of A. wattla; we as-

sume that the elongate duct interpreted by Marcus and

Marcus (1967) as the albumen gland is actually the am-

pulla. Externally, A. wattla is similar to A. muelleri by

having a wide and flattened body, lamellated rhino-

phores emerging from below the anterior border of the

notum, and the genital opening situated posterior to and

below the branchial lamellae. Marcus and Marcus (1967)

argued that A. wattla differs from A. muelleri by the

shape of the caruncule (with two lobes in the former)

and the size of the radular teeth. However, examination

of the re-description of A. muelleri by Marcus and Mar-

cus (1967) shows no significant differences in the shape

of the caruncle.

Because of the absence of consistent and distinctive

differences, we regard A. wattla and A. abbotti as junior

synonyms of A. muelleri.

Page 136

THE NAUTILUS, Vol. 118, No. 4

Figure 5.

Armina elongata new species, holotype, MHNMC INV MOL3902. A. Dorsal and ventral views of the living animal.

B. Rachidian teeth and inner lateral teeth. C. Detail of the rachidian tooth. D. Detail of the innermost lateral tooth. E. Jaw

elements on the masticatory processes. F. Jaws.

Armina elongata new species

(Figures 5-6)

Description: The living animal is red with white lon-

gitudinal notal ridges. Tne anterior margin of the oral

“ell and the apical portion of the rhinophores are also

white (Figure 5A). The preserved specimen is pale gray-

ish, almost white, with the spaces between the longitu-

dinal notal ridges pigmented with dark gray. The body

is elongated, f deienecl and narrower posteriorly. The no-

siren eens 24 longitudinal dorsal ridges. The oral veil is

distinct and has two large, tentacular lateral extensions

without any projections. The extensions are as wide as

the notum. There are two club- shaped rhinophores,

each one with about 20 vertical lamellae situated on a

dorsal notch. The eyes are visible through the epidermis

at the outer base of each rhinophore. The caruncle is

also distinguishable anterior to the rhinophores. The

genital opening is located on the right side, slightly pos-

terior to and below the branchial eanelline. The anal pa-

pilla is located behind the midpoint (2/3 of total animal

Honesto. from the anterior end of the body). There are

2 beanie lamellae and 23 large hyponotal lamellae

on each side of the body. The branchial and hyponotal

N. E. Ardila and A. Valdés, 2004

Figure 6. Armina elongata new species, holotype, MHNMC

INV MOL3902. A. Dorsal view of the anterior end of the body.

Scale bar = 1mm. B. Lateral view of the body. Scale bar as in

A. C. Reproductive system. Scale bar = 1 mm. Abbreviations:

am, ampulla; ap, anal pore; be, bursa copulatrix; bl, branchial

lamellae; ca, caruncule; fg, female glands; gp, genital pore; hl,

hyponotal lamellae; le, lateral euension of velum; n, notum:;

pad, post-ampullary duct; pn, penis; pr, prostate; rh, rhino-

phore; v, vagina; vl, velum.

lamellae show alternation of larger and smaller lamellae.

The pedal gland is white and located at the posterior

end of the foot sole (Figure 5A).

Jaws and Radula: The radula formula is 26 X 33.1.33

in the holotype. The rachidian tooth is narrow, with a

long and pointed median cusp, and bears approximately

20 thin and elongated denticles on either side of the

median cusp (Figure 5C). The lateral teeth are elongat-

ed, curved and also bear denticles that become progres-

sively smaller towards the outermost teeth. The jaws

have a denticulated masticatory border with three to

four rows of denticles (Figures 5E—F).

Reproductive System (Figure 6): The reproductive sys-

tem is diaulic. The ampulla is large and convoluted; it

narrows into a long post-ampullary duct that connects to

the prostate and the female glands. The prostate is short

and convoluted, and connects directly to the penis. The

vagina is very long, curved and connected directly to the

rounded bursa copulatrix.

Type Material: Holotype (MHNMC INV MOL3902),

length 16 mm alive, from type locality.

Type Locality: Off Manaure, Colombia (11°53'5” N

72°36'39" W), 22 m depth, substratum with coralline al-

gae and gravel.

Distribution: This species is only known from the

type locality, off Manaure, Colombia.

Etymology: The specific name refers to the elongate

shape of the body and the masticatory processes of Tikes

species.

Remarks: Armina elongata is clearly distinguishable

from other species of die genus in several regards. Ex-

ternally, the body shape of A. elongata is proportionally

longer and narrower than that of A. juliana or A. muel-

leri. The velum is much wider than that of A. juliana

and as wide as the notum. The genital opening is situ-

ated posterior to the branchial annals, whereas in A.

juliana is anterior to the branchial lamellae.

Other differences include the reproductive system

and the radular morphology. Armina elongata has a

much more elongated vagina than those of A. juliana

and A. muelleri. The rachidian tooth of A. elongata is

most similar to that of A. juliana, but the central cusp

is sharper and more elongated and has more denticles,

almost reaching the apex of the cusp.

The description of A. elongata constitutes the second

record of the genus Armina “aml the family Arminidae

in the Caribbean Sea after the original description of A.

juliana (see Ardila and Diaz, 2002). Additionally, this is

the third arminid known to occur in the western Atlantic

with A. juliana and A. muelleri (see Kolb and Wagele,

1998 and Ardila and Diaz, 2002).

DISCUSSION

Comparisons of the external morphology and anatomy

of Armina elongata, A. muelleri, and A. juliana, exam-

ined in this study, reveal that they are easily distinguish-

able from each other and from other Atlantic species of

the genus by at least one or more external characters.

The numbers of branchial and hyponotal lamellae, which

vary little within the southern Caribbean species, show

a great variation among other Atlantic species, and in

some cases the number of lamellae differs from side to

side. Armina tigrina has the largest number of branchial

Page 138

THE NAUTILUS, Vol. 118, No. 4

lamellae (more than 100 per side), whereas A. maculata

has the largest number of hyponotal lamellae (more than

100 per side). For other Atlantic species, Armina nea-

politana has 18-32 branchial lamellae and 9-19 hypono-

tal lamellae, and Armina loveni has 9-26 branchial la-

mellae and 15-25 hyponotal lamellae (see Kolb, 1998).

Armina wattla has 28 branchial leaves and more than 29

lateral lamellae when all primordial are counted (Marcus

and Marcus, 1967).

The shape of the radular teeth, which appears to show

little intraspecific variability in arminids (Marcus and

Marcus, 1966), is another useful set of characters to dis-

tinguish species (see Kolb, 1998 and Ardila and Diaz,

2002). Several species have a distinct innermost lateral

tooth, frequently bearing small denticles on the sides of

the cusp, whereas the other lateral teeth have elongate

cusps that can be denticulated or smooth. However, the

material of Armina muelleri here examined lacks denti-

cles on the innermost lateral teeth, which are present in

other specimens described in the literature. However,

other anatomical similarities suggest that this variation

in the presence of denticles is likely due to intraspecific

variability. Distinctive radular formulae and rachidian

teeth morphology are found in other Atlantic species of

the genus. In A. tigrina the radular formula is 35 X

56.1.56 and the central cusp of the rachidian tooth is not

as broad as in A. juliana and A. muelleri and bears 15—

18 fine denticles. In A. loveni the radular formula is 35

< 29.1.29 and the rachidian tooth has five coarse den-

ticles on each side.

Finally, the morphology of the reproductive system

has provided useful information to distinguish the three

species here examined. In A. elongata the vagina is very

long, curved and the ampulla is large and narrows into

a long post- -ampullary duct that connects to the prostate,

whereas in A. muelleri the vagina is short and there is

no post-ampullary duct connecting the ampulla directly

to the prostate, which is very long and convoluted. Ar-

mina juliana also has a post-ampullary duct, but the va-

gina is much shorter than in the other two species.

ACKNOWLEDGMENTS

Scott Whitaker and Susan Braden assisted with the pro-

duction of the SEM images, which were improved and

composed with the help of the staff and facilities of IN-

VEMAR, Santa Marta. This paper has been supported

by INVEMAR, COLCIENCIAS (Grant No. 210509-

11248), the Colombian Ministry of the Environment,

and the US National Science Foundation, through the

PEET Grant “Phylogenetic systematics of the Nudibran-

chia” (DEB-0329054) to T. M. Gosliner and the junior

author. This is the Contribution No. 777 of the Research

Marine Institute “José Benito Vives de Andreis” IN-

VEMAR. Material was collected by Adriana Gracia and

Gabriel Navas (MHNMC-INVEMAR).

LITERATURE CITED

Abbott, R. T. 1954. The habits and occurrence of the nudi-

branchs Armina tigrina, in south east United States. The

Nautilus 67: 83-86.

Ardila, N. E. and J. M. Diaz. 2002. Armina juliana (Nudibran-

chia: Arminoidea: Arminidae), a new species from the

southem Caribbean. Boletin de Investigaciones Marinas y

Costeras 31: 25-31.

Eyster, L. S. 1981. Observations on the growth, reproduction

and feeding of the nudibranch Armina tigrina. Journal of

Molluscan Studies 47: 171-181.

ICZN 1999. International code of zoological nomenclature. In-

ternational Trust for Zoological Nomenclature, London.

Ihering, H. von. 1886. Zur Kenntniss der Nudibranchien der

brasilianischen Kiiste. Jahrbiicher der Deutsche Malako-

zoologische Gesellschaft 13: 223-240, pl. 9.

Ihering, H. von. 1915. Die Opisthobranchien der brasilianisch-

en Kiiste. Nachrichtsblatt der Deutschen Malakozoolo-

gischen Gesellschaft 47: 133-143.

Kolb, A. 1998. Morphology, anatomy and histology of four spe-

cies of Armina Rafinesque, 1814 (Nudibranchia, Armi-

noidea, Arminidae) from the Mediterranean sea and the

Atlantic ocean. Journal of Molluscan Studies 64: 355-386.

Kolb, A. and H. Wagele. 1998. On the phylogeny of the Ar

minidae (Gastropoda, Opisthobranchia, Nudibranchia)

with considerations of biogeography. Journal of Zoological

Systematics and Evolutionary Research 36: 53-64.

Marcus, Ev. and Er. Marcus. 1960. Opisthobranchs from

American Atlantic warm waters. Bulletin of Marine Sci-

ence of the Gulf and Caribbean 10: 129-203.

Marcus, Ev. and Er. Marcus. 1966. The R/V Pillsbury deep-

sea biological expedition to the Gulf of Guinea, 1964-65.

9. Opisthobranchs from tropical west Africa. Studies in

Tropical Oceanography 4: 152-208.

Marcus, Ey. and Er. Marcus. 1967. Some opisthobranchs from

Sapelo Island, Georgia, USA. Malacologia 6: 199-222.

Nijssen-Meyer, J. 1965. Notes of a few opishobranchs Mollusca

from Surinam (Guianas). Zoologische Mededelingen 40:

143-150.

Thompson, T. E., R. Cattaneo and Y. M. Wong. 1990. Eastern

Mediterranean Opistobranchia: Dotidae (Dendronoto-

idea), Arminidae and Madrellidae (Arminoidea). Journal

of Molluscan Studies 56: 393-413.

THE NAUTILUS 118(4):139-143, 2004

Page 139

A new species of the genus Anetarca Gosliner, 1991

(Gastropoda: Opistobranchia: Facelinidae) from the western

Atlantic Ocean

Francisco J. Garcia

Departamento de Fisiologia y Zoologia

Facultad Biologfa; Universidad de

Jesus S. Troncoso

Area de Biologia Animal

Facultad Ciencias del Mar

Sevilla Universidad Vigo

Apartado 1095, 41080 Sevilla Lagoas-Marcosende, Vigo

SPAIN SPAIN

figarcia@us. es [email protected]

ABSTRACT

The genus Anetarca Gosliner, 1991, has been known as mono-

typic, with A. armata Gosliner, 1991, described from the Pa-

cific coast of North America, as its type species. In this paper,

a second species of the genus, Anetarca brasiliana new species,

is described from the locality of Buzios, State of Rio de Janeiro

(Brazil). It constitutes the first species of the genus found in

the western Atlantic Ocean. The distinguishing characters of

this species are the color pattern, the presence of lamellate

rhinophores, and the penial duct provided with an apical elon-

gate appendage.

INTRODUCTION

In 1991, Gosliner described the genus Anetarca, with

the type species A. armata, belonging to the family Fa-

celinidae, from the Pacific coast of central Baja Califor-

nia. The present paper describes a new species of Ane-

tarca collected in Buzios (State of Rio de Janeiro, Bra-

zil). It thus constitutes the second known species of that

genus and the first species found in the Atlantic Ocean.

MNCN stands for Nacional de Ciencias Naturales de

Madrid, Spain, and MORG stands for Museu Oceano-

grafico “Prof. Eliézer de Carvalho Rios”, Fundacao

Universidade do Rio Grande, Rio Grande, Brazil.

SYSTEMATICS

Genus Anetarca Gosliner, 1991

Anetarca brasiliana new species

(Figures 1-7)

Description:

External Anatomy (Figures 1-3): Body elongated. Foot

broad and with elongated propodial tentacles. Rhino-

phores long, with 7— 10 lamellae obliquely arranged. La-

mellae connected by a longitudinal ridge on anterior and

posterior faces of rhinophores. Rhinophores join at their

bases. Eyes located behind bases of rhinophores. Oral

tentacles considerably long, their length approximately

half body size. Cerata cylindrical am slightly curved,

cnidosac pointed. Precardiac cerata for ming an arch con-

taining a single row of cerata. Bogtioadhec cerata ar-

ranged in 9 clusters, each containing only a single row.

Number of cerata per cluster in 7.5 mm specimen is:

one precardiac arch with seven cerata and nine postcar-

diac rows with 4, 4, 4, 4, 3, 3, 2, 1, 1 cerata, respectively.

Gonopore ventral to precardiac ceratal arch. Anus cleio-

proct, situated between first two postcardiac ceratal

rows.

Coloration: General body and foot color translucent

orange-brown with numerous white spots more or less

densely concentrated, giving appearance of transverse

white bands along dorsal Gres of notum. Rhinophores

and oral ertecles have same general color pattern, al-

though apical third of rhinophores is almost translucent

white, while on basal two-thirds orange-brown is more

intense. In this portion borders of lamellae are red. Cer-

ata have same superficial color pattern as body, with

translucent-white cnidosac and dark-brown digestive di-

verticulum. Jaws visible externally and rose colored.

Internal Anatomy (Figures 4-11): Buccal apparatus

with a series of oral glands on dorsal and lateral surfaces

of its oral tube. Jaws rose colored, having a rather short

masticatory border with irregular denticles (Figures 5,

8, 9). Radular formula in two of specimens, 7.5 and 8

mm long, is 12 X 0.1.0 and 16 X 0.1.0, respectively.

Teeth arch-shaped, with broad and elongate central cusp

and 9-11 elongate denticles on either side of cusp. Lat-

eral denticles decrease in size toward margins (Figures

10, 11).

Reproductive System (Figures 6, 7): With narrow

preampullar hermaphroditic duct. Duct connected to a

broad and curved ampulla, which divides into a short

oviduct and a narrow and coiled deferent duct. Oviduct

Page 140

THE NAUTILUS, Vol. 118, No. 4

Figure 1.

with a small seminal receptacle located anteriorly to

junction with female gland mass. Deferent duct lacks a

differentiated prostate. Penial papilla broad and curved.

Penial duct shows narrow prolongation at its tip and a

subterminal, curved and elongated chitinous stylet (Fig-

ure 7).

Biological Notes: When the animals are disturbed, the

rhinophores bend forward and touch, crossing each an-

other. In addition, the oral tentacles curve backward,

bending around the rhinophores; the cephalic and pre-

cardiac region of the body move upwards. This display

disappears when the molesting stimulus is removed.

Holotype: MORG 40608, 9 mm length, Francisco J.

Garcia and Jestis S. Troncoso colls., 24 Mar. 1998.

Paratypes: MORG 40609, paratype 1, from type lo-

cality, 6 mm length; MNCN 15.05/33182, paratypes 2

and 3, Praia dos Ossos, Arma¢ao dos Bizios, Brazil, 4

and 4.5 mm length respectively, Francisco J. Garcfa and

Jestis S. Troncoso colls, 26 Mar. 1998.

Type Locality: Collected in the intertidal zone at

Praia da Armagao, Armag¢ao dos Biizios (State of Rio de

Janeiro, Brazil).

Other Material Examined: Two specimens, 7.5 and

§ mm in length, were collected in the intertidal zone at

Praia da Armacao, Armagao dos Buzios (State of Rio de

Janeiro, Brazil) (24 Mar. 1998), Francisco J. Garcia and

Jestis S. Troncoso colls.

Etymology: The name of this species, brasiliana, is

dedicated to Brazil, the country where the specimens

were collected.

Anetarca brasiliana. Living holotype, MORG 40608, 9 mm length.

DISCUSSION

Brazilian opisthobranchs and those from other western

Atlantic areas were intensely studied for more than thir-

ty years by Ernst and Eveline Marcus. However, the

tropical western Atlantic piste panch auna is not as

well known as the Indo-Pacific fauna (Millen and Ha-

mann, 1992) and that from the eastern Atlantic. Marcus

(1977) published a checklist of the warm-water opistho-

branchs found from La Plata (Argentina) to Cape Hat-

teras (USA), in which, eleven species of the family Fa-

celinidae were cited. Posteriorly, Millen and Hamann

(1992) described another species of this family from the

Caribbean Sea, Pauleo jubatus Millen and Hamann,

1992. Garcia and Troncoso recently (2003) named a spe-

cies of Phidiana from Archipelago Fernando de Noro-

nha (off northeastern Brazil). Eight species of Facelini-

dae have been cited from Brazilian waters, Phydiana

lynceus Bergh, 1867, Facelina coenda Marcus, 1958,

Cratena pilata (Gould, 1870), Dondice occidentalis (En-

gel, 1925), Godiva rubrolineata Edmunds, 1964, Nanuca

sebastiani Marcus, 1957, Favorinus auritulus Marcus,

1955 (Marcus, 1977; Rios, 1994) and Phidiana riosi

Garcia and Troncoso, 2003.

The species described in this pepe differs externally

from other facelinid species by the coloration and ar-

rangement of cerata. In Phydiana lynceus, Phydiana rio-

si, and Facelina. coenda all cerata are arranged in rows

(Marcus, 1958; Edmunds and Just, 1983; Garcia and

Troncoso, 2003): in Dondice occidentalis and Godiva

rubrolineata the cerata are arranged in arches with more

than one row of cerata per arch (Marcus, 1958; Ed-

munds, 1964); Nanuca sebastiani has only four groups

F. J. Garcia and J. S. Troncoso, 2004

Page 141

Figures 2-7. Anetarca brasiliana. 2. Diagrammatic right profile showing the insertion of the cerata and position of the anus and

reproductive apertures. 3. Rhinophore. 4. Dorsal view of the buccal apparatus. 5. Jaw. 6. Reproductive system. 7. Detail of the

penial duct. Abbreviations: a, ampulla; an, anus; ens, central nervous system; e, eye; dd, deferent duct; fg, female gland; go, genital

opening; hd, hermaphroditic duct; j, jaw; oe, oesophagus; og, oral glands; pp, penial papilla; s, stylet; sr, seminal receptacle. Scale

lines = 0.5 mm.

of cerata arising from a transversely set base (Marcus,

1957; Edmunds and Just, 1983); in F. auritulus the cer-

ata are arranged in arches (Edmunds and Just, 1983) or

the two first groups of cerata are arched and the hinder

groups form simple rows (Marcus, 1955); finally, Cratena

pilata has the first three groups arched and the three

hinder ones forming oblique rows (Marcus, 1957). In

addition, the rhinophores of our species differ from

those of F. auritulus and C. pilata. In F. auritulus, the

rhinophores are smooth and bear two bulbs (Marcus,

1955; Edmunds and Just, 1983); in C. pilata they are

either covered with small tubercles (Marcus, 1957) or

smooth (Marcus and Marcus, 1967). The penis is un-

armed in all the Brazilian facelinid species except for P.

lynceus, P. riosi, and F: coenda. Phidiana lynceus and P.

riosi have a hook-shaped penial stylet (Marcus and Mar-

cus, 1967; Garcia and Troncoso, 2003) and in F. coenda

the border of the penial leaf bears small protuberances

prolonged into one to four brown spines (Marcus, 1958).

There are six other species of Facelinidae from the

warm western Atlantic waters, Palisa kristenseni (Mar-

cus, 1963), P. papillata Edmunds, 1964, Learchis poica

Page 142

THE NAUTILUS, Vol. 118, No. 4

Figures 8-11. Anetarca brasiliana. Scanning electron micrographs. 8. Jaw. 9. Masticatory border. 10-11. Radular teeth.

Marcus, 1960, L. evelinae Edmunds and Just, 1983, Aus-

traeolis catina Marcus and Marcus, 1967, and Pauleo

jubatus Millen and Hamann, 1992. Our species differs

from both species of Palisa because in these species the

rhinophores are covered with papillae, the radular teeth

lack a prominent central cusp, and the penis is unarmed

(Edmunds, 1964; Marcus and Marcus, 1970). Austraeo-

lis catina has the three anterior groups of cerata arched

and its penis ends on a disc whose edge is beset with

ten broad warts, each bearing a tiny spine; there are two

additional warts on the surface of the disc, also with

spines (Marcus and Marcus, 1967). Pauleo jubatus dif-

fers from our species by the arrangement of the cerata

groups in arches, the shape of the radular teeth, and the

presence of a penial sac (Millen and Hamann, 1992). In

the species of the genus Learchis, the post-pericardial

groups of cerata are arched while the pre-pericardial

ones form oblique rows; the penis is unarmed (Ed-

munds, 1964; Marcus and Marcus, 1970; Edmunds and

Just, 1983).

Learchis poica Marcus and Marcus, 1960, is a variable

Caribbean species. However, there are several anatom-

ical features that allow us to differ L. poica from our

specimens.

Learchis poica has the precardiac cerata arranged in

5-6 rows, and the postcardiac cerata form 6-7 arches

(Marcus and Marcus, 1960; Edmunds and Just, 1983).

In Anetarca brasiliana new species, the precardiac cer-

ata form one arch and the postcardiac are arranged in 9

rows. The radular teeth of L. poica have a prominent

cusp flanked by 5-8 denticles (Marcus and Marcus,

1960), while in A. brasiliana the radular teeth have 9—

11] lateral denticles on each side of the central cusp. In

addition, the deferent duct in L. poica is divided into a

proximal narrow, not-glandular portion and a wide pros-

tatic region that continues within the blunt penis (Mar-

cus and Marcus, 1960); in A. brasiliana there is not a

differentiated prostatic region of the vas deferent.

We have allocated the new species in the genus Ane-

tarca Gosliner, 1991, because of the arrangement of cer-

ata, with a pre-anal arch and single postanal rows, the

shape of the radular teeth with a prominent central cusp,

the male ducts lack a penial gland and the penis has a

subterminal and curved penial spine.

Our specimens coincide with Anetarca in the features

that characterize that genus, except the rhinophores, be-

cause in A. armata (the type species) the rhinophores

are smooth while in our species they have oblique la-

mellae. Thus, we suggest that it should be allocated in

the genus Anetarca, and that the generic definition be

expanded to include species with smooth or lamellate

rhinophores. A similar situation occurs in genus Flabel-

F. J. Garcia and J. S. Troncoso, 2004

Page 143

lina (Flabellinidae), which includes species with smooth,

or ringed (annulate or perfoliate), or papillate rhino-

phores (Gosliner and Griffiths, 1981; Gosliner and Wil-

lan, 1991).

In addition to the ornamentation of the rhinophores,

A. brasiliana differs externally from A. armata because

in the former the oral tentacles are clearly longer than

the rhinophores, while in A. armata they are shorter.

Internally, A. brasiliana has the masticatory border of

the jaws denticulated and the penial duct having an elon-

gated apical appendage.

ACKNOWLEDGMENTS

We wish to express our gratitude to Laboratorio de Ma-

lacologia, Departamento de Zoologia, Universidade Fed-

eral do Rio de Janeiro, Brazil, for their assistance during

the expedition to Cabo Frio and Buzios, Brazil, and to

Dr. Ricardo S. Absalao and Dr. Paulo Marcio Costa for

their constant help. This research was included in a pro-

ject supported by Agencia Espafiola de Cooperacién In-

ternacional (AECT) and Ministerio de Educacién y Cien-

cia, Spain, and has been partially supported by the pro-

ject PHB2002-0045-PC of this latter organization.

LITERATURE CITED

Edmunds, M. 1964. Eolid Mollusca from Jamaica, with de-

scriptions of two new genera and three new species. Bul-

letin of Marine Science of the Gulf and Caribbean 14: 1—

32.

Edmunds, M. and H. Just. 1983. Eolid Nudibranchiate Mol-

lusca from Barbados. Journal of Molluscan Studies 49:

185-203.

Garcia, F. J. and J. S. Troncoso. 2003. Two unknown species

of Mollusca Gastropoda from the Archipelago Fernando

de Noronha (Brazil), with description of a new species

belonging to the genus Phidiana Gray, 1850 and a new

anor! ae Dewdiradlarts senegalensis Bouchet, 1975. Scien-

tia Marina 67: 159-166.

Gosliner, T. M. 1991. Four new species and a new genus of

opisthobranch gastropods from the Pacific coast of North

America. The Veliger 34: 272-290.

Gosliner, T. M. and R. J. Griffiths. 1981. Description and re-

vision of some South African Aeolidacean Nudibranchia

(Mollusca, Gastropoda). Annals of the South African Mu-

seum 84: 105-150.

Gosliner, T. M. and R. C. Willan. 1991. Revision of the Fla-

bellinidae (Nudibranchia: Aeolidacea) from the Tropical

Indo-Pacific, with the description of five new species. The

Veliger 34: 97-133.

Marcus, Er. 1955. Opisthobranchia from Brazil. Boletim da Fa-

cultade de Filosofia, Ciéncias e Letras, Universidade de

Sao Paulo, Zoologia 20: 89-262.

Marcus, Er. 1957. On ‘Opisthobranchia from Brazil (2). Journal

of the Linnean Society of London, Zoology 43: 390-486.

Marcus, Er. 1958. On western Atlantic opisthobranchiate gas-

tropods. American Museum Novitates 1906: 1-82.

Marcus, Er, and Ev. Du B.-R. Marcus. 1970. Opisthobranchs

from Curagao and faunistically related regions. Studies on

the Fauna of Curagao and other Caribbean Islands 33: 1—

129.

Marcus, Ev. Du B.-R. 1977. An annotated check list of the

Western Atlantic warm water Opisthobranchs. Journal of

Molluscan Studies, Supplement 4: 1-22.

Marcus, Ey. and Er. Marcus. 1960. Opisthobranchs from

American Atlantic warm waters. Bulletin of Marine Sci-

ence of the Gulf and Caribbean 10: 129-203.

Marcus, Ev. and Er. Marcus. 1967. Tropical American Opis-

thobranchs. Studies in Tropical Oceanography, University

of Miami, 6: 3-137.

Millen, S. V. and J. C. Hamann. 1992. A new genus and species

of Facelinidae (Opisthobranchia: Aeolidacea) from the

Caribbean Sea. The Veliger 35: 205-214.

Rios, E. C. 1994. Seashells of Brazil. 2"¢ Edition. Editora da

Fundagao Universidade do Rio Grande, Rio Grande, 492

212

THE NAUTILUS 118(4):144-151, 2004

Page 144

Two new species of Leptochiton Gray, 1847 (Polyplacophora)

from the Iberian Peninsula (eastern Atlantic coast)

Pilar Carmona Zalvide

Departamento de Fisiologia y Biologia

Animal

Facultad de Biologia

Universidad de Sevilla

Apartado 1095, 41080 Sevilla

SPAIN

Compostela

SPAIN

Victoriano Urgorri

Departamento de Bioloxia Animal

Facultade de Bioloxia

Universidade de Santiago de

15706 Santiago de Compostela

Francisco J. Garcia!

Departamento de Fisiologia y Zoologia

Animal

Facultad de Biologia, Univ. de Sevilla

Apartado 1095, 41080 Sevilla

SPAIN

[email protected]

ABSTRACT

Two new species of the genus Leptochiton (Polyplacophora)

from the Atlantic Ocean are described: Leptochiton (Lepto-

chiton) troncosoi and Leptochiton (Leptochiton) pepezamorat

were collected off Spain at 753-832 m depth. Both species

have been included in genus Leptochiton, subgenus Leptochi-

ton, due to the presence of thin valves without insertion plates,

tegmentum finely granulated with granules of equal size, and

the dorsal girdle covered with small scales. The new species

are characterized by their elongate-oval shape, tegmentum

sculptured with thin rounded granules, which tend to form

strings, and girdle dorsally covered with long, non-imbricate

scales. The larger lateral tooth of the radula is bicuspid in L.

(L.) troncosoi, and unicuspid in L. (L.) pepezamorai. Among

all the known species of Leptochiton, the new species are more

similar to L. (L.) xanthus, L. (L.) tenuis, L. (L.) geronensis and

L. (L.) thalattius. In this paper the anatomic features of the

new species are compared with all the species of Leptochiton.

INTRODUCTION

The species of the genus Leptochiton Gray, 1847, have

been revised by Kaas (1979, 1981, 1991), Van Belle

(1983), Kaas and Van Belle (1985, 1987, 1990, 1994),

Dell’Angelo and Palazzi (1991, 1986, 1987), Della Bella

and Dell’Angelo (1985), Cesari (1987), and Carmona

and Urgorri (1999). These studies allow us to determine

the actual worldwide diversity of Leptochiton. Neverthe-

less, some specimens collected off northwestern Spain

have features that differ from those of the species de-

scribed in these previous works. We describe herein, us-

ing light and scanning electron microscopes, two new

species of Leptochiton based on their distinctive shell,

radula, and girdle scales.

MATERIALS AND METHODS

The specimens were collected in 1990-1991 during the

“CANGREXO I” campaign off A Quiniela (Galicia, north-

' Author for correspondence.

western Spain) between 753-832 m depth. The speci-

mens were collected using royal crab (Chaeceon affinis)

traps. They were preserved in 70% ethanol. To study the

hard parts (shells, scales, girdle spicules, and radulae)

the animals were macerated in 10% KOH and rinsed

with distilled water. For SEM (Philips XL-20), shells,

radulae, and girdle scales were coated with gold. MNCN

stands for Museo Nacional de Ciencias Naturales, Ma-

drid, Spain.

SYSTEMATICS

Class Polyplacophora Gray, 1821

Order Neoloricata Bergenhayn, 1955

Suborder Lepidopleurina Thiele, 1910

Family Leptochitonidae Dall, 1889

Genus Leptochiton Gray, 1847 (Type species: Chiton

cinereus Montagu, 1803, non Linnaeus, 1767).

Diagnosis: Valves thin, lacking insertion plates and

tegmentum finely granulated. The granules are of equal

size and the girdle is dorsally covered with small scales.

Leptochiton (Leptochiton) troncosoi new species

(Figures 1-20)

Diagnosis: Elongate-oval shape, twice as long as wide,

moderately elevated, the back is evenly rounded and

with a marked apex. The color of tegmentum is dark

beige. The tegmentum is sculptured with thin rounded

granules that tend to form strings among which semi-

circular and concentric growth lines of the shell are

shown. The girdle is narrow, covered with long non-im-

bricate scales. The major lateral tooth of the radula is

bicuspid, and the internal cusp is small.

Description: The head valve has a semicircular ante-

rior border whereas the posterior valve forms such a

blunt angle that it hides the notch of the concave apex.

The strong slope originated at the head valve is convex

(Figure 1). The intermediate valves have rectangular

borders except for the second valve, whose anterior mar-

P. C. Zalvide et al., 2004 Page 145

Figures 1-9. Leptochiton (L.) troncosoi. 1. Valve I. 2. Valve II. 3. Valve IV. 4. Valve VIII. 5. Arrangement of granules on jugal

area. 6. Arrangement of granules on pleural area. 7. Arrangement of granules on lateral area. 8. Arrangement of megalaesthetes

and micraesthetes on central area of the granules. 9. Arrangement of megalaesthetes and micraesthetes on lateral area of the

granules.

Page 146 THE NAUTILUS, Vol. 118, No. 4

Figures 10-20. Leptochiton (L.) troncosoi. 10-11. Radula. 12. Central and first lateral radular teeth. 13. Cusp of the major

lateral tooth and spatulate uncinal tooth. 14. Arrangement of ventral scales and marginal spicules. 15. Ventral scales. 16. Arrange-

ment of dorsal scale. 17. Dorsal scale, dorsal view. 18. Dorsal scale, ventral view. 19. Dorsal corpuscle. 20. Marginal spicules.

P. C. Zalvide et al., 2004

gin is completely convex (Figures 2, 3). However, in the

remainder of valves it tends to be convex although it may

be slightly concave in the jugal sinus. The lateral borders

are rounded while the posterior borders are straight;

they converge on the apex, and a little concave curvature

is noticeable on both sides. The lateral areas are not

raised, but are only visible on the sculpture. The tail

valve is smaller in size than the head valve. The anterior

margin of the caudal valve is similar to the intermediate

valves and the semicircular posterior border. It has a

prominent mucro at an antero-central position, which

forms a strong slope (Figure 4).

The tegmentum has rounded granules forming a

stem-like sculpture. The granules form parallel longitu-

dinal strings in the central area (Figure 2). The granules

are wider and flattened in the jugal area, and exhibit an

apical stem divided into three parts that are usually con-

nected to the next granule (Figure 6). On the pleural

area, the granules are longer and the stems are smooth.

The number of strings on the central area may be be-

tween 63 and 67 in the intermediate valves. At the head

valve, lateral areas, and potancrl area, the strings are

radially oriented (Figures 1, 2, 3, 4). The granules on

both terminal valves are rounded with no stems (Figure

5), while on the lateral areas the shape is wider (Figure

7); arranged on a radially striate tegmentum, three stri-

ations are found on each granule. Eighty-nine strings are

observed on the head valve, 58 on the tail valve, and 11—

14 on the lateral areas.

The aesthetes are arranged in groups of five on each

granule, although sometimes only three aesthetes may

be seen (Figures 8, 9). They are situated at the edges of

the granules except for the central one, which is located

in a subcentral position on the granule.

The articulamentum is white, has fragile appearance,

and lacks insertion plates. The apophyses are widely sep-

arated by the large jugal sinus. It has triangular shape

on the intermediate valves and is trapezoid with rounded

edges on the tail valve.

The dorsal girdle is covered with slightly convex, elon-

gate, curved, pointed scales, up to 90 X 53 ym each

(Figure 16). They are sculptured with about 5-8 longi-

tudinal ribs (Figures 17-19). Along the marginal there

is a fringe of stronger cylindrical spicules up to 200 wm

in length (Figure 14). Ventrally the girdle is lined with

triangular scales of 20-28 j1m; scales become more elon-

gate (70-120 jm) and weakly ribbed toward the outer

margin.

The gills are arranged in rows at both sides of the

foot. They are inserted at the level of valve VII, extend

to the anus with a gradual increase in size, but size de-

creases at the last valve. Thus, the gills are classified as

merobranchial abanal without interspace. The number

of gills on each side is 7.

The central tooth of the radula has rectangular shape

with a narrow flexible blade (Figure 12). The first lateral

tooth, with a similar appearance to the central tooth,

exceeds this one slightly (Figure 12). The major lateral

tooth is strongly developed in bicuspid form, with the

Page 147

internal cusp being clearly smaller than the outer cusp

(Figure 13).

Holotype: MNCN 15.03/482, 8.5 X 3.8 mm, 1990,

“CANGREXO I” expedition. Four valves and the radula

were used for SEM, while the rest of the specimen was

preserved in alcohol 70%; holotype is the only specimen

known.

Type Locality: Off A Quiniela, Galicia, northwesterm

Spain, 43°17'22"—43°18'52” N, 09°36'38’—09°35'45”" W,

753-832 m depth [“Cancrexo I” expedition]. It was

found between 753-832 m. The type locality is in an

area with strong currents, and the holotype was found

attached to a rock from a bottom with ferromanganese

nodules, calcareous plaques, and coal slag.

Distribution: Leptochiton (L.) troncosoi is a deep-wa-

ter species, known only from the type locality, off A Qui-

niela (Galicia, NW Spain).

Etymology: The species has been named Leptochiton

(Leptochiton) troncosoi in honor of Dr. Jestis S. Tron-

coso.

Leptochiton (Leptochiton) pepezamorai new species

(Figures 21-38)

Diagnosis: Characterized by elongate oval shape, dor-

sal region evenly rounded, not careened, and without a

marked apex. The tegmentum is white, although an

ochre-oxide coloration is sometimes present. The teg-

mentum is sculptured with rounded granules. Two or

three stems reach along the frontal area to the basal area

of the adjacent granule. This creates some furrows on

the tegmentum interrupted by the granules and concen-

tric growth lines. The girdle is narrow, covered with rect-

angular non-overlapping scales. The major lateral tooth

of “ine radula is unicuspid. The spatulate tooth is well

developed and it exceeds the major lateral tooth.

Description: The head valve has a semicircular ante-

rior border and the posterior border has a little notch

on the apex. The valve presents a strong convex slope

(Figure 21). The intermediate valves, clearly rounded in

their sides, are dorsally rectangular, except for the sec-

ond valve, which has a triangular shape. The anterior

border of this area is slightly convex, almost straight. The

anterior and posterior borders of the remainder of the

valves are straight. The lateral areas are not raised, only

marked by a change in sculpture (Figures 92, 23). The

tail valve is roughly triangular with size similar to the

head valve. The mucro is antero-central, prominently

marked where the originated slope is slightly convex

(Figure 24).

The tegmentum is sculptured with rounded granules

and two or three stems on the apical zone, showing a

more or less concentric arrangement on the head alvee

lateral areas, and postmucral area (Figures 25, 27). On

the central areas, the granules are arranged in longitu-

dinal rows separated by shallow grooves. The number of

rows varies between 56 and 67.

Page 148 THE NAUTILUS, Vol. 118, No. 4

Figures 21-28. Leptochiton (L.) pepezamorai. 21. Valve I. 22. Valve II. 23. Valve IV. 24. Valve VIII. 25. Arrangement of

pustules on jugal area. 26. Arrangement of pustules on pleural area. 27. Arrangement of pustules on lateral area. 28. Arrangement

of megalaesthetes and micraesthetes on pustules.

P. C. Zalvide et al., 2004 Page 149

Figures 29-38. Leptochiton (L.) pepezamorai. 29. Radula. 30. Spatulate uncinal tooth. 31. Central and first lateral teeth. 32.

Cusp of major lateral tooth and spatulate uncinal tooth. 33. Dorsal scale. 34-35. Dorsal corpuscles. 36. Ventral scale, dorsal view.

37. Ventral scales, ventral view. 38. Marginal spicules.

Page 150

THE NAUTILUS, Vol. 118, No. 4

As a general rule, the stems give striated aspect to the

tegmentum, and are interrupted by the granules. In ad-

dition, the lateral areas, head valve, and postmucral area

always show a variable number of growth marks. Each

granule bears three aesthetes. The megalaesthete is lo-

cated in a central-basal position while the micraesthetes

are located in an upper level at both sides.

The articulamentum is white, lacking insertion plates.

The apophyses are triangular in shape in the interme-

diate valves and trapezoid in the tail valve. The wide

jugal sinus separates them.

The dorsal girdle is lined with pointed, rectangular,

slightly curved scales. They are sculptured with 14 to 16

longitudinal grooves (Figure 33). Their size range is 40—

62 wm in length and 25-40 ym on the base. Scattered

among these scales are smaller ones, up to 47 wm in

length, with § longitudinal ribs. Also, the girdle has

smooth dorsal spicules up to 160 jm in length. The

sharply rectangular ventral scales, wp to 65 zm in length,

have a weak median, longitudinal rib up to 110 wm in

length (Figure 36). The marginal fringe shows pointed

conic spicules, which are sculptured with 4 longitudinal

ribs (Figure 38). Another, less abundant, type of spicules

is present. These have three longitudinal ribs up to 54.4

wm in length.

The gills are located at both sides of the foot, extend-

ing from valve VIII to the anus; the gills are classified

as merobranchial abanal without interspace.

The central tooth of the radula is rectangular having

a well-defined flexible edge. The first lateral tooth, sim-

ilar in shape to the central tooth, also has a well-defined

flexible terminal edge, and exceeds the central tooth

slightly in length (Figures 29 and 31). The major lateral

tooth is unicuspid with a slightly blunt edge (Figure 29).

The spatulate uncinal tooth exceeds the major lateral

tooth (Figure 30).

Type Material: Holotype: MNCN 15.03/484, 1.4 x

0.7 mm; Paratype 1, from type locality, Animal Biology

Department, Santiago de Compostela University un-

numbered, 2.4 X 1.3 mm; Paratypes 2, 3, Animal Biol-

ogy Department, Santiago de Compostela University un-

numbered, 1.75 X 1.2 mm and one broken specimen,

“CaNncREXO I” Expedition, 43°23’31” N; 19°32'19” W,

840 m depth.

Type Locality: 43°17'18" N, 09°36’35" W, 753-786 m

depth. A strong current was present in the area. The

specimens were attached to a rock from a bottom with

ferromanganese nodules, calcareous plaques, and coal

slag.

Distribution: Leptochiton (L.) pepezamorai is known

only from the type locality, A Quiniela (Galicia, NW

Spain). It is a deep-water species, found in a depth of

753-840 m.

Etymology: The species has been named Leptochiton

(Leptochiton) pepezamorai in honor of Mr. José Zamora.

DISCUSSION

According to the diagnoses given by Kaas and Van Belle

(1985) for the genus and subgenus, the species de-

scribed in this paper are classified in genus Leptochiton,

subgenus Leptochiton.

Both new species differ from the species of Lepto-

chiton from the western Atlantic Ocean, South Africa,

Pacific, and Indo-Pacific Oceans by the sculptures of

their tegmentum, girdle scales, and the number of cusps

of their major lateral radular teeth (Kaas and Van Belle,

1985, 1987, 1990, 1994).

Among all the species of Leptochiton worldwide, the

only one that is considered cosmopolitan is L. alveolus

(Lovén, 1846) (Kaas and Van Belle, 1985), since it occurs

in the Atlantic, Pacific, and Indian Oceans. It differs

from L. (L.) troncosoi and L. (L.) pepezamorai by the

sculpture of the tegmentum and by the scales of the

girdle.

According to the descriptions of Kaas (1979, 1981,

1991), Kaas and Van Belle (1985, 1987, 1990, 1994),

Dell’Angelo and Palazzi (1987, 1991), Cesari (1987) and

Carmona and Urgorri (1999), L. (L.) troncosoi and L.

(L.) pepezamorai differ from the remaining species from

the Eastern Atlantic Ocean and Mediterranean Sea by

the features below.

In relation to the animal shape, four types have been

described: oviform, oblong, oval and elongate oval. L.

(L.) troncosoi and L. (L.) pepezamorai are allocated in

the fourth type together with L. alveolus, L. cancellatus

(Sowerby II, 1840), L. cimicoides (di Monterosato,

1879), L. scabridus (Jeffreys, 1880), L. intermedius (von

Salvini-Plawen, 1968), L. leloupi Kaas, 1979, L. tenuis

Kaas, 1979, L. sarsi Kaas, 1981, L. gascognensis Kaas

and Van Belle 1985, L. thalattius Kaas and Van Belle,

1985, L. geronensis Kaas and Van Belle, 1985, L. bedullii

Dell’Angelo and Palazzi, 1986, L. xanthus Kaas and Van

Belle, 1990, L. pseudogloriosus Strack, 1991, and L.

compostellanum Carmona and Urgorri, 1999.

Of all these species, only L. tenuis, L. thalattius, L.

geronensis, L. xanthus, and L. bedullii have the tegmen-

tum sculptured with granules arranged in longitudinal

rows on the central area, and the dorsal scales of the

girdle are longer than wide, like in L. troncosoi and L.

pepezamorai.

In relation to the number of cusps of the major lateral

teeth of the radula, L. tenuis and L. xanthus are similar

to L. pepezamorai and L. geronensis is similar to L. tron-

cosoi. The radula of L. thalattius Kaas and Van Belle

(1985) has not been described, but this species differs

from L. pepezamorai because the tegmentum is not fur-

rowed and the dorsal scales of the girdle are bigger and

have a smaller number of ribs.

Leptochiton thalattius differs from L. troncosoi by the

sculpture of the tegmentum because in the former the

central area of the jugal area has about eight parallel

rows, is weakly developed, the four or five adjoining

rows on the pleurae are curved as result of short inter-

polated rows (posteriorly as well as anteriorly), the re-

P. C. Zalvide et al., 2004

Page 151

maining pleural rows are parallel, somewhat diverging

toward the anterior (Kaas and Van Belle, 1985). The

sculpture of L. troncosoi has parallel and longitudinal

strings on the central area. Besides this, the girdle is

dorsally covered with erect scales; among them, calcar-

eous spicules occur and intersegmental spines are found.

The scales of L. troncosoi are curved and they do not

present either dorsal spicules or intersegmental spines.

Comparing L. pepezamorai to L. tenuis, according to

Kaas (1979), the tail valve is much smaller than the other

valves, while that of L. pepezamorai is of a similar size.

The dorsal scales of the girdle of L. tenuis are bigger

and they have a smaller number of ribs, and the cusp of

the major lateral is sharply pointed. In relation to L.

xanthus, following the description of Kaas and Van Belle

(1990), the dorsal scales of the girdle are of smaller size

and they have a smaller amnialser. of ribs, they lack dorsal

spicules and the cusp of the major lateral tooth is sharply

pointed.

Taking into account the description of L. geronensis

by Kaas and Van Belle (1985) it differs from L. troncosoi

by the tegmentum sculpture of round granules, which

are very separated and by having the two cups of the

major lateral tooth of similar size.

LITERATURE CITED

Carmona [Zalvide], P. and V. Urgorri. 1999. Descripeién de dos

nuevas especies de Moluscos Poliplacéforos de la Penin-

sula Ibérica: Leptochiton (L.) gascognensis Kaas and Van

Belle, 1985 y L. (L.) compostellanus sp. nov. Iberus 17:

97-107.

Cesari, P. 1987. Note sistematiche e geonomiche su alcune spe-

cie mediterranee del genere Leptochiton Gray, 1847 (Po-

lyplacophora, Leptochitonidae). Lavori—Societa Vene-

ziana di Scienze Naturali 12: 3-34.

Dell’Angelo, B. and S. Palazzi. 1986. Considerazione sulla fa-

miglia Leptochitonidae Dall, 1889 (Mollusca: Polyplaco-

phora) con desenzione di due nuovi taxa. Bolletino Ma-

lacologico 22 (1-4): 1-36.

Dell’ Angelo, B. ad S. Palazzi. 1987. Considerazione sulla fa-

miglia Leptochitonidae Dall, 1889 (Mollusca: Polyplaco-

phora). Il Ridescrizione di Leptochiton cimicoides (Mon-

terosato, 1879). Bolletino Malacologico 23 (1-4): 95-105.

Dell’Angelo, B. and S. Palazzi. 1991. Considerazione sulla fa-

miglia Leptochitonidae Dall, 1889 (Mollusca: Polyplaco-

pore). IV. Aggiunte e correzioni. Bolletino Malacologico

7 (Ye B= 38.

Della Bella, G. and B. Dell’Angelo. 1985. Prima segnalizacione

di Leptochiton geronensis Kaas and Van Belle, 1985 lungo

le coste Avalon, Bolletino Malacologico 21 (10-12): 309-

310.

Kaas, P. 1979. On a collection of Polyplacophora (Mollusca,

Amphineura) from the bay of Biscay. Bulletin du Muséum

national d’Histoire naturelle, Paris 4 (1): 13-31.

Kaas, P. 1981. Scandinavian species of Leptochiton Gray, 1847

(Mollusca, Polyplacophora). Sarsia 66: 217-229.

Kaas, P. 1991. Chitons (Mollusca: Polyplacophora) procured by

the CANCAP I-VII expeditions, 1976-86. Zoologische

Mededelingen 65: 89-98.

Kaas, P. and R. A. Van Belle. 1985. Monograph of Living Chi-

tons. 1, Order Neoloricata: Lepidopleurina. E. J. Brill/W.

Backhuys, Leiden, 240 pp.

Kaas, P. and R. A. Van Belle. 1987. Monograph of Living Chi-

tons. 3, Ischnochitonidae: Chaetopleurinae, Ischnochiton-

inae. E. J. Brill/W. Backhuys, Leiden, 302 pp.

Kaas, P. and R. A. Van Belle. 1990. Monograph of Living Chi-

tons. 4, Suborder Ischnochitonina: Ischnochitonidae: Isch-

nochitoninae (continued) Additions to vols. 1, 2 and 3: 1—

298 pp. E. J. Brill, Leiden, 298 pp.

Kaas, P. and R. A. Van Belle. 1994. Monograph of Living Chi-

tons. 5 Suborder Ischnochitonina: Ischnochitonidae: Isch-

nochitoninae (concluded). Callistoplacinae; Mopalidae.

Additions to vols. 14. E. J. Brill/W. Backhuys, Leiden,

403 pp.

Van Belle, R. 1981. The systematic classification of the chitons

(Molluca: Polyplacophora). Informations de la Societe

Belge de Malacologie 11 (1-3): 1-178.

THE NAUTILUS 118(4):152-156, 2004

Page 152

Cirsotrema (Gastropoda: Ptenoglossa: Epitoniidae) in the

Miocene Chipola Formation of northwestern Florida

Richard Duerr

P.O. Box 1055

Okeechobee, FL 34973 USA

ABSTRACT

The genus Cirsotrema is represented in the Lower Miocene

Chipola Formation by two species, Cirsotrema dalli Rehder,

1945, also present in the Pliocene and Pleistocene of southern

Florida mal extant in the western Atlantic, and Cirsotrema cir-

ritum new species, found only in the Chipola Formation. A

search of published records revealed no earlier occurrence of

Cirsotrema dalli other than its presence in the Chipola For-

mation.

INTRODUCTION

The family Epitoniidae dates from the Triassic (Clench

and Turner, 1950) and by the Cretaceous was well es-

tablished with worldwide distribution. Sohl (1964: 317)

created the genus Striaticostatum for seven species from

the Cretaceous of the southeastern United States that

have features similar to those now placed in Cirsotrema

Morch, 1852. The very faint spiral striations on the body

whorl of Striaticostatum separate it from Cirsotrema

which has stronger spiral sculpture. Palmer (1937) as-

signed five species from the Eocene of the southeastern

United States to Cirsotrema from the Eocene of the

same area.

Only two species of Cirsotrema have been found in

the Lower Miocene Chipola Formation, restricted to

that portion of the Chipola Formation in the Chipola

River drainage of Calhoun County, Florida, in the region

of Tenmile Creek, Farley Creek, and the Chipola River,

from State Road 20 north to the mouth of Tenmile

Creek. Cirsotrema dalli Rehder, 1945, a common spe-

cies in the Pleistocene Bermont Formation of southem

Florida (Hoerle, 1970) and extant in the westerm Atlantic

(Abbott, 1974), is represented in the Chipola Formation

by specimens from at least ten separate localities present

in the Invertebrate Paleontology collection of the Flor-

ida Museum of Natural History at the University of Flor-

ida, Gainesville, Florida. Although found throughout the

Chipola Formation, Ciao dalli is uncommon at

any particular locality. A species similar in shell mor-

phology to Cirsotrema dalli from the tropical eastern

Pacific, C. togatum Hertlein and Strong, 1951, ranges

from Baja California south to the Galapagos Islands

(Keen, 1971). Cirsotrema togatum has also been found

in the Pliocene Esmeraldas beds of northwestern Ec-

uador (Pitt, 1981; DuShane, 1988). The new species,

Cirsotrema cirritum, is also distributed throughout the

Chipola Formation within the Chipola River drainage,

but has been collected at only five localities and is rare.

Three species exhibit a similarity to Cirsotrema cirritum.

Cirsotrema acutum (J. Sowerby, 1813), from the Eocene

Barton beds of Great Britain, Cirsotrema crassicostatum

(Deshayes, 1850), found in the Miocene of Belgium, and

Cirsotrema excelsum Garcia, 2003, a Recent species

from the Indo-Pacific.

Institutional abbreviations used are: USNM, National

Museum of Natural History, Smithsonian Institution,

Washington, DC, USA; BMNH, British Museum of Nat-

ural History, iLamdlom. England; RMNH, National Mu-

seum of Natural History/Naturalis, Leiden, The Neth-

erlands; UF, Florida Museum of Natural History, The

University of Florida, Gainesville, Florida, USA; BMSM,

The Bailey-Matthews Shell Museum, Sanibel, Florida,

USA; LACM, Los Angeles County Museum of Natural

History, Los Angeles, California, USA. “P. Diegel collec-

tion” refers to the collection of Phyllis Diegel, West

Palm Beach, Florida, USA. “Sunderland collection” re-

fers to the collection of Kevan and Linda Sunderland,

Sunrise, Florida.

SYSTEMATICS

Superfamily Janthinoidea Lamarck, 1812

Family Epitoniidae Berry, 1910

Genus Cirsotrema Mérch, 1852

Cirsotrema Morch, 1852: 49.

Type Species:

monotypy.

Scalaria varicosa Lamarck, 1822, by

Diagnosis: Shell white to gray, turriculate; body whorl

usually with broad spiral ord. often covered with fine

spiral and axial lines forming microscopic reticulate

sculpture. Raised axial lines forming microscopic retic-

ulate sculpture. Raised axial costae composed of numer-

ous lamellations, frequently foliated, sometimes covering

entire body whorl, with or without prominent varices.

All possess a basal ridge. Aperture circular to oval.

R. Duerr, 2004 Page 153

Figures 1-10. Cirsotrema species. 1. Cirsotrema dalli Rehder, 1945, Recent, Anses d’Arlets, Martinique, height 43.8 mm, width

13. 5 mm, P. Diegel collection. 2. Cirsotrema pilsbryi McGinty, 1940, Recent, taken off St. oe Florida, by Ted Yocius, height

15.3 mm, width 6.4 mm, Sunderland collection. 3. Cirsotrema dalli Rehder, 1945, fossil, 45354, Chipola Formation, Tenmile

Creek, Calhoun County, Florida, height 18.3 mm, width 7.0 mm. 4-7. Cirsotrema ae new species. 4. Holotype, UF 110972,

height 30.1 mm, width 11.8 mm. 5. Paratype, UF 112019, sculpture of teleoconch, scale bar = 5 mm. 6. Paratype, BMSM 15301,

height 54.1 mm, width 22.0 mm. 7. Paratype, USNM 522028, height 50.4 mm, width 18.2 mm. 8. Cirsotrema cf. woodringi Olsson,

1967 (= “C. arcella Rehder’, Woodring, 1959), UF 112142, Gatun Formation, road cut east of Cativa, Province of Colén, Panama,

height 36.0 mm, width 13.6 mm. 9. Cirsotrema acutum (J. Sowerby, 1813), UF 112018, Barton Clay Formation, Barton on Sea,

Hampshire, England, height 16.2 mm, width 7.7 mm. 10. Cirsotrema crassicostatum (Deshayes, 1850), RMNH Lot 497 (No.

65166?), Zanden V. Antwerpen, height 37.3 mm, width 14.9 mm.

Cirsotrema dalli Rehder, 1945 Cirsotrema (Cirsotremopsis) arcella Rehder, 1945: 128; Olsson,

(Figures 1, 3) 1967: 40, pl. 5, figs. 4a.

° Cirsotrema arcella Rehder, Clench and Tumer, 1950: 228, pl.

Cirsotrema (Cirsotremopsis) dalli Rehder, 1945: 128; Olsson, 98, fig.3, [considered by Clench and Tumer to be conspe-

1967: 40, pl. 5, figs. 2-2b. cific with C. dalli].

Page 154

Cirsotrema dalli Rehder, Clench and Turner, 1950: 227-228,

pl. 98, fig. 1, 3; Warmke and Abbott, 1961: 71-78, pl. 14,

fig. H; Morris, 973: 154, pl. 44, fig. 1; Abbott, 1974: 114,

fig. 1188; Humfrey, 1975: 91, pl. 7, fig. 12; Rios, 1975: 57,

pl. 15, fig. 219; Abbott and Dance, 1982: 69: Sunderland,

1990: 14; Rios, 1994: 99, pl. 33, fig, 404,

Description: Shell medium to large, turriculate; te-

leoconch whorls 9 or 10, convex, with 5 or 6 broad spiral

cords overlain with fine spiral and axial lines, forming a

microscopic reticulated sculpture. Suture deep. Fim-

brious axial costae composed of fine, wavy lamellae, with

a crosshatched pattern inclined upwards abaperturally,

hooked at shoulder, sinuous, with alternating waves so

that each succeeding costa contacts abapertural costa,

forming a secondary surface which is almost flat or

slightly convex, leaving only very small holes or pits.

Prominent varices present at irregular intervals. Aper-

ture subcircular; with labral varix.

Holotype: USNM 515240

Type Locality: 29°14’ N, 85°29’ W, off Cape San Blas,

Florida, in 25 fathoms.

Remarks: Some authors consider Cirsotrema dalli to

be a junior synonym of the Recent Cirsotrema cochlea

(G. B. Sowerby II, 1844) from the eastern Atlantic

(Clench and Turner, 1950: 228; Weil, et al., 1999: 14).

Also, Cirsotrema cochlea may be a junior synonym of

Cirsotrema pumicea (Brocchi, 1814). A comparative

study of the relationship between C. dalli, C. cochlea,

and C. pumicea has not been made and is beyond the

scope of this paper.

Cirsotrema togatum may be considered the eastern

Pacific cognate of C. dalli. Both have fimbriated costae

frequently covering the entire surface, similar overall

shapes, and varices at irregular intervals. Both may also

have costae of varying widths due to erosion, especially

on fossil specimens. The costae of dead collected Recent

specimens are frequently eroded, as are the early whorls

of live-taken specimens, exposing the sculpture on por-

tions of the body whorl. A figure of a Recent specimen

of Cirsotrema togatum by Keen (1971, fig. 633) and fig-

ures of Bhocener specimens by Pitt (1981, figs. 2, 3) and

DuShane (1988, figs. 10, 11, 12) show specimens mor-

phologically similar'to ©) dalla

DuShane (1974, figs. 54 and 55) illustrated two spec-

imens of Cirsotrema togatum. DuShane’s figure 54, a

live-taken specimen, is similar to the Recent specimens

examined by this author, with the costae extending up

over the suture and onto the preceding whorl, the suture

subdued by the surface sculpture, and lacking a notice-

able shoulder on the whorls, giving a somewhat straight

appearance to the whorl profile. ‘DuGhanes figure 55,

the holotype, exhibits a definite indentation at the suture

with shouldered costae and some exposed intercostal ar-

eas, very similar to the holotype of Cirsotrema dalli. The

discrepancy between DuShane’s figure 54 and the ho-

lotype of Cirsotrema togatum disineved! in figure 55

THE NAUTILUS, Vol. 118, No. 4

might be resolved by the description of the Recent spec-

imen illustrated in figure 54 as a separate, new species.

Although somewhat similar to Cirsotrema dalli, C

pilsbryi McGinty, 1940, a Recent species endemic to the

western Atlantic, is easily distinguished from this spe-

cies. The wavy costae of Cirsotrema dalli touch each

other to a varying extent, forming a secondary superficial

surface and leaving only small holes or pits between the

adherent portions of the costae, obscuring the surface of

the whorl itself, while the costae of C. pilsbryi are more

sloping at the shoulder, with a subdued hook, giving the

surface of the costae a more rounded appearance than

that of C. dalli. Also, the costae of Cirsotrema pilsbryi

consist of wavy lamellations without the foliations or

crosshatched effect of the costae of C. dalli. Both Cir

sotrema dalli and C. pilsbryi have irregularly spaced var-

ices, have a microscopic reticulated sculpture on the

body whorl, and have broad spiral cords which angle

slightly towards the apex adaperturally, rather than re-

volving parallel to the whorl, although the sculpture on

the body whorl is frequently not visible under the sec-

ondary surface on C. dalli. The principal characteristics

separating Cirsotrema dalli from C. pilsbryi are well il-

lustrated in Sunderland (1989, 1990).

The Recent Indo-Pacific Cirsotrema ernestoilaoi

Garcia, 2001, is easily separated from C. dalli by the

trapezoidal shaped (with wide shoulder) profile of the

whorl formed by the axial costae, and lack of pits or

holes in the surface sculpture of C. ernestoilaoi.

Cirsotrema woodringi Olsson, 1967, a rare species

currently found only fa the Miocene Gatun Formation

of Panama and the Pliocene Tamiami Formation at Sun-

nyland, Florida, has foliated costae similar to C. dalli. It

differs by the straight line separating the costae, which

abut, completely covering the surface of the whorl. Cir

sotrema dalli has wavy costae leaving small pits or open-

ings in the secondary surface formed by the costae. As

Olsson so aptly stated in reference to Cirsotrema wood-

ringi (also applicable to C. dalli regarding costae), “Gen-

eral surface has a finely porous texture resembling that

produced by a linen cloth” (1967: 41). This feature may

be attributed to intritacalx (D’Attilio and Radwin, 1971),

a shell layer not well studied in the Family Epitoniidae.

Olsson (1967: 41) reported a specimen of Cirsotrema

dalli from “McClellan Farm’, a locality on the west bank

of the Chipola River south of Tenmile Creek, which he

stated “was carefully compared with the type of C. dalli,

a Recent species, and no distinguishing difference could

be found”.

Cirsotrema cirritum new species

(Figures 4-7)

Description: Shell medium to large, turriculate. Early

whorls missing on all specimens examined. First existing

whorl of holotype (last protoconch whorl) with sculpture

ranging from smooth to 4 turn with several single mi-

croscopic wavy axial lamellae that gradually enlarge into

foliated costae. No clear distinction between protoconch

R. Duerr, 2004

Page 155

and teleoconch present. Teleoconch with 8 tumid

whorls. Suture deep. Axial costae 13 on last whorl,

strongly recurved, hooked, and angled slightly adaper-

turally on shoulder, extending in a straight line over su-

ture; surface of costae composed of multiple wavy la-

mella with very fine irregular diamond or square pattem;

pattern inclined abaperturally. Intercostal spaces vari-

able, usually wider than costae, with 5 broad, rounded

spiral cords, overlain with S$ to 12 much finer cords,

crossed by equally fine axial growth lines so as to form

a microscopic, reticulated pattern. Reticulated pattern

continue from intercostal spaces to cover adapertural

side of recurved costae. Top of wavy axial costae sharp.

Broad spiral cords on intercostal spaces reproduced on

abapertural surface of costae and angled 45° apically.

Adaptertural surface of costae covered with wavy cross-

hatched lamellae. Fine line of demarcation separates

leading edge of costae where joined by succeeding in-

tercostal space. Anterior reflected projections on axial

ribs of last whorl forming coarse, undulating basal ridge.

Varices absent. Aperture holostomatous.

Holotype: UF 110972, height 30.1 mm, width 11.8

mm.

Paratypes: BMSM 15301, protoconch missing, height

54.1 mm, width 22.0 mm, 30°28.030’ N, 85°09.572’ W

(= Taller University locality TU 458), east bank of Chi-

pola River, above Farley Creek, (SW % Sec. 10, TIN,

R9W), Calhoun County, Florida, Chipola Formation;

USNM 522028, spire and aperture missing, height 50.4

mm, width 18.2 mm, 30°29.44’ N, 85°11.17' Ww. (= tue

lane University locality TU 951, = United States Geo-

logical Survey locality 26578), Tenmile Creek, about 2

km west of Chipola River, (SE % Sec. 12, TIN, R10W),

Calhoun County, Florida, Chipola Formation; UF

67746, spire missing, height 29.8 mm, width 10.3 mm,

30°27.45’ N, 85°08.45’ W, (= Tulane University locality

TU 825), Farley Creek at abandoned mill about 350 m

west of bridge on Florida Highway 275 (SW % Sec. 21,

TIN, R9W), Calhoun County, Florida, Chipola Forma-

tion; UF 112019, body whorl only, with aperture, height

16.5 mm, width 20.1 mm, same locality as previous spec-

imen.

Type Locality: 29°30.05’ N, 85°11.00' W, Tenmile

Creek, at power line crossing about 1.6 km west of Chi-

pola River (SE % Sec. 7, TIN, RLOW), Calhoun County,

Florida, Chipola Formation.

Etymology: The name cirritum is derived from the

Latin cirrus meaning “filamentous”, referring to the

fringed costae of the new species.

Discussion: All specimens of Cirsotrema cirritum ex-

amined lack the apical whorls. It may be assumed that

the missing whorls consist of 1 to 3 smooth whorls as

evidenced by the remainder of a partial smooth proto-

conch whorl on the holotype. The width of the axial cos-

tae and the intercostal spaces appear to be a variable

characteristic on all specimens studied. Features which

the eastern Pacific Cirsotrema togatum and the western

Atlantic C. dalli have in common with GC. cirritum are

turreted shape, sculpture on the surface of the costae,

and spiral cords with microscopic reticulated sculpture

on the body whorl. Cirsotrema cirritum differs from C.

togatum and C. dalli by its larger size, narrower costae,

onl intercostal areas, and leak of varices. Although the

foliated face on the costae of Cirsotrema togatum and

C. dalli presents a flat surface, the foliated adapertural

surface of the costae of C. cirritwm is recurved. Also, a

sharply impressed line of demarcation separates each

costa from the succeeding intercostal space and next cos-

ta on C. cirritum, indicating a pause in growth.

The Cirsotrema species from the Eocene of the

southeastern United States are all under 30 mm, have

narrower spiral cords, and lack the reticulated sculpture

on the intercostal areas and the crosshatched effect pre-

sent on the costae of C. cirritum. The Eocene Cirsotre-

ma acutum (J. Sowerby, 1813) from Great Britain, and

the Miocene C. crassicostatum (Deshayes, 1850) from

Europe are very similar to each other. Both have lon-

gitudinal laminations on the costae without the cross-

hatched effect of Cirsotrema cirritum, have narrower,

more pronounced spiral cords, and lack the axial stria-

tions present on the intercostal spaces of C. cirritum.

Cirsotrema acutum is also smaller and more attenuate

posteriorly than C. cirritum.

There is no ae sculpture visible on Maury’s

(1925: 242, pl. 37, fig. 4) figure of a very eroded single

whorl of the nallotaes of Cirsotrema tamanensis (Maury,

1925) from the Miocene of Trinidad.

Several Recent Indo-Pacific species of Cirsotrema

have features similar to C. cirritum. Cirsotrema plexis

Dall, 1925, and C. fimbriatulum (Masahito, Kuroda and

Habe, 1971) may be distinguished from C. cirritum as

both have varices and have about 20 axial costae on the

body whorl as opposed to 13 on C. cirritum. Varices are

lacking on Cirsotrema rugosum (Kuroda and Ito, 1961)

and C. excelsum Garcia, 3003, but both have a greater

number of axial costae on each whorl and the “eantes

differ in sculpture from those of C. cirritum. The sculp-

ture on the surface of the axial costae of Cirsotrema

richeri Garcia, 2003, most closely resembles that of the

costae of C. cirritum. A greater number of axial costae

on the body whorl and the presence of varices on Cir

sotrema richeri separates it from C. cirritum.

Cirsotrema cirritum has only been recorded from the

Chipola Formation of northwestern Florida. Cirsotrema

dalli, which may have originated in the Lower Miocene

Chipola Formation, extended its range over southern

Florida during the Pliocene and Blectocene epochs, and

survives in the Recent of the western Atlantic Ocean.

ACKNOWLEDGMENTS

The author thanks José H. Leal, BMSM, for general as-

sistance, digital photography, and for preparing the

plate. Roger Portell, UF, kindly allowed access to the

collections under his care, provided the SEM image, and

Page 156

reviewed the manuscript. Gary Schmelz graciously do-

nated the holotype of Cirsotrema cirritum. The gener-

osity of Warren Blow and Mark Florence, NMNH;

Frank Wesselingh, RMNH, and Lindsey Groves,

LACM, for specimen loans from their respective insti-

tutions is greatly appreciated, with a special note of

thanks to Paul Jeffery, BMNH, for donation of speci-

mens of Cirsotrema acutum. Richard Petit made avail-

able a rare specimen of Cirsotrema cf. woodringi, criti-

cally reviewed the manuscript, and provided pertinent

information and reference material. Kevan and Linda

Sunderland are thanked for the loan of a specimen of

Cirsotrema pilsbryi and information regarding Recent

Cirsotrema. Thanks also to two anonymous reviewers

whose suggestions greatly improved the manuscript and

to Paula Mikkelsen, Phyllis Diegel, and Marc Gregis for

helpful information. Burke and Brooks Hayes, Archie

and Vicky Whitling, and William Tatum graciously grant-

ed the author permission to collect on their respective

properties.

LITERATURE CITED

Abbott, R. T. 1974. American Seashells. 2° edition. Van Nos-

trand-Reinhold, New York, 663 pp., 24 pls.

Abbott, R. T. and S. P. Dance. 1986. Compendium of Seashells.

E. P. Dutton, New York, x + 411 pp.

Clench, W. J. and R. Turner. 1950. The genera Sthenorytis,

Cirsotrema, Acirsa, Opalia and Amaea in the western At-

lantic. Johnsonia, Museum of Comparative Zoology 2(29):

291-248.

D’Attilio, A. and G. E. Radwin. 1971. The intritacalx, an un-

described shell layer in mollusks. The Veliger 13: 344-347.

Dockery, D. T. III. 1980. The invertebrate macropaleontology

of the Clarke County, Mississippi area. Mississippi De-

partment of Natural Resources 122: 387.

DuShane, H. 1974. The Panamic-Galapagan Epitoniidae. The

Veliger 16 (Supplement): 84, 154 figs.

DuShane, H. 1988. Pliocene Epitoniidae of the Esmeraldas

beds of northwestern Ecuador (Mollusca: Gastropoda).

Tulane Studies in Geology and Paleontology 21(1, 2): 51-

58, 12 figs.

Hertlein, L. G. and A. M. Strong. 1951. Mollusks from the

west coast of Mexico and Central America. Part X. Zool-

ogica 36: 67-120, pls. 1-11.

Hoerle, S. 1970. Mollusca of the “Glades” Unit of southern

Florida, Part II, List of molluscan species from the Belle

Glade rock pit, Palm Beach County, Florida. Tulane Stud-

ies in Geology and Paleontology 8(1, 2): 56-68.

THE NAUTILUS, Vol. 118, No. 4

Humfrey, M. 1975. Sea Shells of the West Indies. Taplinger

Publishing, New York, 351 pp., 32 pls.

Keen, A. M. 1971. Sea Shells of tropical west America: marine

mollusks from Baja California to Peru. 24 edition. Stan-

ford University Press, Stanford, XIV + 1064 pp., 22 pls.

[1984 reprint with only 12 pls.].

Maury, C. J. 1925. A further contribution to the paleontology

of Trinidad (Miocene horizons). Bulletins of American Pa-

leontology 10(42): 250, 43 pls.

McGinty, T. L. 1940. New marine shells dredged off Palm

Beach, Florida. The Nautilus 54; 62-64.

Morch, O. A. L. 1852. Catalogus Conchyliorum quae Reliquit

D. Alphonso d’Aguitta et Gadea Comes de Yoldi. 1: Ce-

phalophora. L. Klein, Hafniae, 170 pp.

Morris, P. A. 1973. A Field Guide to Shells of the Atlantic and

Gulf Coasts and the West Indies. Houghton-Mifflin, New

York, 330 pp., 76 pls.

Olsson, A. A. 1967. Some Tertiary mollusks from south Florida

and the Caribbean. Paleontological Research Institution,

61 pp.

Palmer, K. V. W. 1937. The Claibornian Scaphopoda, Gastro-

poda and Dibranchiate Cephalopoda of the southern

United States. Bulletins of American Paleontology 7(32):

548, 90 pls.

Pitt, W. D. 1981. Two new gastropod occurrences in the Ec-

uadorian Neogene. Tulane Studies in Geology and Pale-

ontology 16(4): 155-156, figs. 1-3.

Rehder, H. A. 1945. Two new species of Cirsotrema (Epito-

niidae) from Florida. Proceedings of the Biological Society

of Washington 58: 127-130.

Rios, E. C. 1975. Brazilian Marine Mollusks Iconography. Fun-

dac&o Universidade de Rio Grande, Rio Grande, 331 pp.,

91 pls.

Rios, E. C. 1994. Seashells of Brazil. 2:4 edition. Funda¢ao

Cidade [and] Fundacao Universidade do Rio Grande, Mu-

seu Oceanografico “Professor Eliézer de Carvalho Rios”,

Rio Grande, 368 pp., 113 pls.

Sohl, N. F. 1964. Neogastropoda, Opisthobranchia and Basom-

matophora from the Ripley, Owl Creek, and Prairie Bluff

Formations. United States Geological Survey Professional

Paper 331B: iv + 153-344, pls. 19-52.

Sunderland, K. 1989. Caribbean Epitoniidae. American Con-

chologist 17: 15-16.

Sunderland, K. 1990. Caribbean Epitoniidae. Part II. American

Conchologist 18: 14-15.

Warmke, G. L. and R. T. Abbott. 1961. Caribbean Seashells.

Livingston Publishing, Narberth, 346 pp.

Weil, A., L. Brown and B. Neville. 1999. The Wentletrap Book.

Evolver, Rome, 224 pp.

Woodring, W. P. 1959. Geology and Paleontology of the Canal

Zone and adjoining parts of Panama. United States Geo-

logical Survey Professional Paper 306-B: III + 147-239,

pls. 24-38.

THE NAUTILUS 118(4):157-159, 2004

Page 157

A new species of Stramonita (Gastropoda: Muricidae) from the

Late Pliocene of Florida

Geerat J. Vermeij

Department of Geology

University of California at Davis

One Shields Avenue

Davis, CA 95616 USA

vermeij @geology.ucdavis.edu

Greg S. Herbert!

Department of Geology

University of California at Davis

One Shields Avenue

Davis, CA 95616 USA

[email protected]

ABSTRACT

Stramonita penelaevis new species from the Caloosahatchee

Formation of southeastern Florida is a smooth rocky-shore

muricid gastropod. Stramonita penelaevis, the first member of

its clade from Florida, represents a group that has become

geographically restricted to the South Atlantic since the late

Pliocene.

INTRODUCTION

Muricids of the subfamily Rapaninae are common mem-

bers of rocky-shore faunas throughout the tropics, but

they are uncommon as fossils. The known fossils docu-

ment surprising patterns of geographic restriction during

Neogene times (Vermeij, 2001). In tropical America, for

example, the genus Neorapana Cooke, 1918, is known

today only as three eastern Pacific species, but during

the early Miocene the genus was also present on the

Caribbean coast of Venezuela (Gibson-Smith et al.,

1997). A group of species today represented by the

South Atlantic S. bicarinata (Blainville, 1832) occurred

in the Caribbean during the late Miocene (Vermeij,

2001).

The genus Thaisella Clench, 1947, found in the west-

ern Atlantic today from Central America and the south-

ern Caribbean to Brazil, was represented in the late Pli-

ocene Caloosahatchee Formation of Florida by a species

that has since contracted its range to the southern Ca-

ribbean (Petuch, 2004). The eastern Atlantic and Med-

iterranean species Stramonita haemastoma (Linnaeus,

1767) made a brief western Atlantic appearance in the

early Pleistocene Bermont Formation of Florida, tem-

porarily coexisting with native western Atlantic members

of the Stramonita haemastoma group that have existed

there from early Miocene to Recent times (Vermeij,

2001).

1 Current address: Department of Geology, University of South

Florida at Tampa, 4202 East Fowler Ave., SCA 528, Tampa,

FL 33620-5201 USA

In this paper we describe a new species of the rapan-

ine muricid genus Stramonita Schumacher, 1817, S. pe-

nelaevis. This species is the first member of its group

(the S. bicarinata group) known from Florida, and

strengthens the case for post-Pliocene geographic re-

striction of its clade. Despite the worm condition of the

material, the distinctiveness and geographic peculiarities

of the new species prompt us to name and describe the

material.

SYSTEMATIC PALEONTOLOGY

Genus Stramonita Schumacher, 1817

Stramonita penelaevis new species

(Figures 1-5)

Type Species: Stramonita haemastoma (Linnaeus,

1767).

Diagnosis: Ovate-elongate Stramonita with five den-

ticles inside outer lip, obsolete axial waves on last whorl,

and very low undifferentiated spiral cords.

Description: Shell small for genus, maximum length

40.5 mm, not constricted abapically; all specimens heavi-

ly worn; teleoconch consisting of four preserved whorls

separated by indistinct, appressed sutures; axial sculp-

ture of last whorl consisting of about seven very low, very

broad, barely perceptible swellings or waves; traces of

spiral cords present, revealing low, undifferentiated

cords, which do not form nodes; aperture elongate-

ovate; outer lip abraded at edge, thickened, its inner side

bearing five prominent denticles; a second row of den-

ticles situated further inside aperture; inner lip smooth,

its abapertural margin slightly recessed; adapical end of

inner lip with prominent parietal tooth; siphonal fasciole

low, broad, rounded; umbilical slit absent.

Type Locality: Caloosahatchee Formation, Palm

Beach Aggregates (GKK Rock Pit), off State Road 80,

Loxahatchee, Palm Beach County, Florida.

Holotype: UF 114426, shell length 36.5 mm, diame-

ter 24.4 mm.

Page 155

THE NAUTILUS, Vol. 118, No. 4

Figures 1-5.

Stramonita penelaevis new species. 1—3. Holotype, UF 114426, shell length 36.5 mm, shell diameter 24.4 mm. 1.

Ventral view. 2. Dorsal view. 3. Lateral view of the outer lip showing two growth checks in close succession and the terminal lip.

Enlarged image of first growth check behind terminal lip shows well-preserved sharp crenulated (toothy) edge, which has been

worn away on terminal lip. 4-5. Paratype A, UF 114427,

view.

Paratype: UF 114427, shell length 40.5 mm, shell di-

ameter 5.3 mm.

Etymology: Latin pene, almost; and laevis, smooth.

Remarks: Stramonita penelaevis is most similar to the

Recent S. bicarinata (Blainville, 1832) from the South

Atlantic islands of St. Helena and Ascension. The new

species differs from S. bicarinata in having obsolete axial

sculpture and lacking the two rows of ede character-

istic of S. bicarinata.

The new species also resembles S. quadridentata (Vokes,

1989) from the late Miocene of the Dominican Republic

and Panama (see also Vermeij, 2001). The latter species

is even more sculptured than S. bicarinata, and like that

species has four to five denticles on the inner side of the

outer lip.

Together with the early Miocene S. semiplicata Ver-

meij, 2001, from the Cantaure Formation of Venezuela,

these three species—S. bicarinata, S. penelaevis, and S.

quadridentata—torm a group (and probable clade) of

species in Stramonita with distinct denticles instead of

shell length 40.5 mm, shell diameter 2

5.3 mm. 4. Ventral view. 5. Dorsal

continuous lirae on the inner side of the outer lip. As

pointed out by Vokes (1989), similar species occur in the

Neogene of Europe.

Vermeij (2001) suggested that the South Atlantic S.

bicarinata is part of a clade that during late Neogene

times was more widespread in tropical America. He not-

ed that the eastern Pacific genus Acanthais Vermeij and

Kool, 1994 (represented by the single Recent species A.

brevidentata (Wood, 1828)), is related to the Atlantic S.

bicarinata group. Acanthais differs from these species of

Stramonita by possessing a labral tooth and by having a

prominent, central columellar fold. The latter feature is

present but reduced in S. semiplicata from the early

Miocene of Venezuela. The discovery of S. penelaevis in

the Caloosahatchee Formation (late Pliocene) of Florida

provides further evidence that the S. bicarinata group

was more widely distributed in the Atlantic during Mio-

cene and Pliocene times than it is today. The new spe-

cies represents the first and only record of this group of

species in the Caloosahatchian biogeographic province

of Petuch (1982).

G. J. Vermeij and G. S. Herbert, 2004

Page 159

ECOLOGY

The new species of Stramonita described here is unusual

among members of Stramonita in being almost smooth.

The near absence of axial sculpture in S. penelaevis

could be the result of abrasion, but the denticulate

sculpture on the inner side of the outer lip is well pre-

served. We are therefore inclined to the view that the

strongly reduced external sculpture is real. In the Recent

fauna, such major sculptural reduction characterized

species that occupy surf-swept rocky shores. Examples

include Purpura persica (Linnaeus, 1758) from the west-

em Pacific; Thais meretricula Réding, 1798, at the is-

lands of Fernando de Noronha (off Brazil) and Ascen-

sion; and some populations of Acanthais brevidentata in

the eastern Pacific. These observations lead us to sur-

mise that the new species of Stramonita lived in a heavi-

ly waveswept environment. Habitats like this yield few

fossils. Co-occurring with S. penelaevis at the type lo-

cality of S. penelaevis in the Caloosahatchee Formation

are specimens of an almost smooth-shell population that

we tentatively assign to the S. haemastoma (Linnaeus,

1767) complex. Specimens of this population as well as

those of S. penelaevis are worn, indicating probable

transport by currents from a wave-swept shore to depths

where sedimentation takes place.

ACKNOWLEDGMENTS

We thank Edward Petuch for taking Herbert to the lo-

cality from which our new species comes, and Janice

Cooper for technical assistance.

LITERATURE CITED

Blainville, H. de. 1832. Disposition méthodique des espéces

Récentes et fossiles des genre pourpre, ricinule, licorne et

concholépas de M. de Lamarck et description des espéces

nouvelles ou peu connues faisant partie de la collection

du Muséum d'Histoire Naturelle de Paris. Nouvelles An-

nales du Muséum d'Histoire Naturelle de Paris 1: 189-

263.

Clench, W. J. 1947. The genera Purpura and Thais in the west-

ern Atlantic. Johnsonia 2: 61-91.

Cooke, A. H. 1918. On the radula of the genus Acanthina, G.

Fischer. Proceedings of the Malacological Society of Lon-

don 13: 6-11.

Gibson-Smith, J., W. Gibson-Smith and G. J. Vermeij. 1997.

Pacific Mexican affinities of new species of the gastropod

genera Macron (Pseudolividae) and Neorapana (Murici-

dae) from the Cantaure Formation (early Miocene) of Ve-

nezuela. The Veliger 40: 358-363.

Linnaeus, C. 1758. Systema naturae per regna tria naturae:

secundum classis, ordines, genera, species, cum charac-

teribus, differentiis, synonymis, locis. Ed. 10, Reformata.

L. Salvii, Holmiae, 824 + iii pp.

Linnaeus, C. 1767. Systema naturae per regna tria naturae:

secundum classis, ordines, genera, species, cum charac-

teribus, differentiis, synonymis, locis. Ed. 13 (decima ter-

tia), ad editionem duodecimam reformatam holmiensem.

Joannis Thomae, Vindoboni, pp. 533-1347.

Petuch, E. J. 1982. Geographical heterochrony: contempora-

neous coexistence of Neogene and Recent molluscan fau-

nas in the Americas. Palaeogeography, Palaeoclimatology,

Palaeoecology 37: 277-312.

Petuch, E. J. 2004. Cenozoic seas: the view from eastern North

America. CRC Press, Boca Raton, 308 pp.

Réding, P. F. 1798. Museum Boltenianum: catalogus cimilior-

um e tribus regnis naturae quae olim colegerat Joa. Frid

Bolten, M.D.Pd. Pars Secunda. J. Christi, Trippii, Ham-

burgi, vii + 199 pp.

Schumeches C. F. 1817. Essai d'un nouveau systéme des hab-

itations des vers testacés avec XXII planches. Schultz, Co-

penhague, 287 pp.

Vermeij, G. J. 2001. Distribution, history, and taxonomy of the

Thais clade (Gastropoda: Muricidae) in the Neogene of

tropical America. Journal of Paleontology 75: 697-705

Vermeij, G. J. and S. P. Kool. 1994. Evolution of labral spines

in Acanthais, new genus, and other rapanine muricid gas-

tropods. The Veliger 37: 414424.

Vokes, E. H. 1989. Neogene paleontology in the northern Do-

minican Republic 8. The family Muricidae (Mollusca:

Gastropoda). Bulletins of American Paleontology 97: 5—

94.

Wood, W. 1828. Supplement to the Index Testaceologicus; or

a catalogue of shells, British and foreign. W. Wood, Lon-

don, 59 pp.

THE NAUTILUS 118(4):160-166, 2004

Page 160

A new Orbitestella (Gastropoda: Heterobranchia:

Orbitestellidae) from Tierra del Fuego, Argentina

Luiz R. L. Simone

Museu de Zoologia da Universidade

de Sao Paulo

Caixa Postal 42594

04299-970 Sado Paulo

BRAZIL

[email protected]

ARGENTINA

Diego G. Zelaya

Division of Invertebrate Zoology

Museo de La Plata

1900 La Plata, Buenos Aires

[email protected]

ABSTRACT

A new species of Orbitestella is described from Tierra del Fue-

go, Argentina. The morphology of the jaws is the main char-

acter used for the generic allocation. Orbitestella patagonica

new species is characterized by a widely umbilicated shell with

low spire, ornamented with numerous, regularly distributed,

spiral threads and weak sinuous commarginal growth lines. The

entire surface of the protoconch is covered with crisp spiral

threads. Examination of the anatomy revealed the presence of

a relatively small stomach and the posterior region of genital

system bulging into haemocoel space as main distinguishing

characters.

INTRODUCTION

The Orbitestellidae are extremely minute, discoid, and

widely umbilicated gastropods, living mainly in the

Southern Hemisphere (Ponder, 1967, 1990; Powell,

1979). The genus Orbitestella Iredale, 1917 has a vir-

tually worldwide distribution, although most of the

known species occur in Australia and New Zealand (Pon-

der, 1990; Ponder and Keyzer, 1998). In the north At-

lantic, two species have been described by Rolén and

Rubio (1992): O. similis from Cape Verde and O. cubana

from Cuba. The only species of the genus presently

known from off South America is Orbitestella ponderi

Linse, 2002, a species described from the East Beagle

Channel (Tierra del Fuego). Linse (2002) identified as

O. ponderi specimens from southern Chile previously

reported by Ponder (1990) as Orbitestella sp.

Orbitestella has previously been placed in the Tro-

choidea or Truncatelloidea, either in the Omalogyridae

or Orbitestellidae (Iredale, 1917; Thiele, 1929; Knight et

al., 1960). Ponder (1967), based on the structure of the

animals and radula, placed Orbitestella in the Rissooidea

[as Rissoacea]. This point of view was followed by several

subsequent authors (Powell, 1979; Ponder, 1983; Mar-

shall, 1988; Hickman and McLean, 1990). Later on,

Ponder (1990) after having studied the anatomy of two

orbitestellid species [Orbitestella wareni Ponder, 1990,

and Microdiscula charopa (Tate, 1899)] suggested that

Orbitestellidae should not be considered as rissooidean

but as primitive Heterobranchia, related to the north-

em-hemisphere freshwater family Valvatidae. Ponder

and Warén (1988) placed Orbitestellidae in the Valva-

toidea.

In the present paper a new species of Orbitestella

from the Magellanic Region is described on the basis of

shell morphology and general anatomy.

MATERIALS AND METHODS

The studied specimens were collected at the Beagle

Channel, Tierra del Fuego, Argentina (Figure 1). Sam-

ples were taken with a dragnet, fixed in 10% buffered

formalin and transported to the laboratory where mol-

lusks were sorted from sediments using a stereoscopic

microscope, and preserved in 70% ethanol. Shell, radula,

jaws, and operculum morphology were studied and il-

lustrated using scanning electron microscopy (SEM).

For anatomical studies specimens were decalcified in

Railliet-Henry’s solution. Two specimens were processed

for histology. Semi-thin sections (0.3 sm thick) were

stained with Mallory’s trichrome (Gabe, 1968). Addition-

ally, 5 specimens were dissected under stereomicro-

scope. It was not possible to study the right side of these

specimens (i.e. pallial tentacle, penis, and propodial

flap).

vere specimens were deposited in the Museo de

La Plata (MLP) and Museu de Zoologia da Universidade

de Sao Paulo (MZSP).

Abbreviations used in the figures: ad, anterior lobe of

digestive gland; an, anus; ca, capsule gland; ce, cerebral

ganglion; eg, pallial genital gland; em, columellar mus-

cle; es, crystalline style sac; dg, digestive gland; es,

esophagus; ey, eye; fs, foot sole; ft, foot; h, heart; go,

hermaphrodite gonad; gp, pedal ganglion; hy, inner

shell organic layer (hypostracum); in, intestine; jw, jaw

plate; mb, mantle border; mf, mantle U-shaped fold;

mg, mantle gland; mo, mouth; od, odontophore; ol,

L. R. L. Simone and D. G. Zelaya, 2004

Dee

Figure 1. Map showing the type locality (®) of Orbitestella

patagonica new species and new findings of O. ponderi (*).

periostracum; op, operculum; os, osphradium; pa, pos-

terior lobe of pedal gland; pe, periostracum; pg, poste-

rior mucous gland; po, pigmented mantle organ; pr,

propodium; pt, prostate; ro, renal organ; sg, salivary

gland; st, stomach; sv, seminal vesicle; sy, statocyst; te,

cephalic tentacle.

SYSTEMATICS

Genus Orbitestella Iredale, 1917 (Type species:

Cyclostrema bastowi Gatliff, 1906, by original

designation)

Orbitestella patagonica new species

(Figures 2-18)

Diagnosis: Shell minute, discoid, with low spire,

widely umbilicated, ornamented with weak sinuous com-

marginal growth lines and numerous, regularly distrib-

uted spiral threads. Protoconch with crisp spiral threads

covering entire surface. Radula with wide, pectinated

rachidian tooth with sharply pointed cusps; lateral tooth

plate-like, with a main cusp and sparsely serrated outer

margin; marginal tooth slender, sickle-like. Stomach rel-

atively small.

Description: SHELL (Figures 2-6, 8-10): Minute

(maximum diameter 1.1 mm), discoid, widely umbilicat-

ed, thin, translucent, shiny, up to 2.75 rounded and reg-

ularly expanding whorls; spire depressed, slightly raised

(Figures 2-5). Color yellowish-buff. Periostracum thin,

colorless. Protoconch formed by 0.75 whorl, about 200

wm wide; initially only sculptured with low granulations,

followed by 10-12 spiral threads (Figure 6). First 0.25

whorl weakly marked by straight growth disruption (Fig-

ure 6). Teleoconch of up to 2 rounded whorls, suture

incised. Umbilicus deep, wide, representing about 32 %

of maximum shell diameter (Figure 5). Shell surface

with weak, sinuous commarginal growth lines and nu-

merous, regularly distributed and fine spiral threads

(Figure 8), also visible inside umbilicus (Figure 5); first

whorl of teleoconch with about 10 spiral threads, last

whorl with 35-37. Shell aperture nearly circular (about

240 ym high by 270 zm wide) with sinuous peristome

(Figures 4, 5).

Page 161

SHELL ULTRASTRUCTURE: Only one well-defined

crossed-lamellar layer, about 25 jm thick (Figure 9). Or-

ganic layer (hy), a third of periostracum thickness, made

more evident in decalcified shell (Figure 16).

Heab-Foor (FicurEs 11, 16): Color cream-yellowish,

uniform in preserved specimens. Head inlaid into foot

(Figure 11). Pair of cephalic tentacles well developed.

Eyes relatively large, sunken into integument medially

between bases of cephalic tentacles; lens solid, volume

about half that of eyes (Figure 16). A pair of tentacle-

like flaps (sometimes referred to as a “snout”) lying be-

tween tentacle bases, flanking mouth. Foot somewhat

massive, cylindrical when retracted, occupying % whorl;

sole entirely covered by long cilia. Columellar muscle

thick. Posterior mucous gland occupying posterior and

middle region of foot.

OPERCULUM (FicuRE 7): Nearly circular, paucispiral,

up to 3.5 slightly overlapping whorls, horny, thin, trans-

lucent, yellowish; nucleus subcentral.

PALLIAL Cavity (Ficures 16, 17): Expanded by %

whorl, with thin, simple mantle border. Gill absent. U-

shaped low fold, sometimes described as “a short, par-

tially ciliated lobe”, dorsally located (Figure 17, mf). Os-

phradium small, fold-like, ciliated, located on left side.

Pigmented mantle gland (seen by transparency) circular,

thin, relatively small, color dark brown, surface uniform,

located at middle-posterior part of roof of pallial cavity

(Figure 9, po).

VISCERAL MASS (FIGURES 11, 16, 17): Comprising two

whorls. Digestive gland almost fully developed on last

whorl. Stomach and gonad located in first half of visceral

whorls, stomach in right side, gonad in left (Figure 16).

CIRCULATORY AND EXCRETORY SYSTEMS (FIGURE 16):

Heart small, located between stomach and renal organ,

posterior to pallial cavity. Renal organ triangular in section.

DIGESTIVE sYsTEM: Mouth on a small protuberance,

between tentacle-like flaps (Figure 16). Oral tube nar-

row, laterally expanded. Jaw plates large, rounded, lat-

erally located and flanked by well-developed muscles

(Figure 16); with five serrated rows of simple curved

plates, posteriorly concave, on each side of inner surface

(Figure 15). Odontophore similar to jaws in size, with

several small muscles, but no cartilages (Figure 16).

Radula 1:1:1:1:1, somewhat longer than odontophore.

(Figure 12). Rachidian tooth large, trapezoidal, with

peg-like lateral edges and an excavated base; cutting

edge pectinated, wide, strongly curved inwards, bearing

11—13 slender and sharply pointed cusps, gradually be-

coming smaller in outward direction; central cusp

straight; lateral cusps somewhat curved, narrower than

central one (Figure 14). Lateral teeth plate-like with

straight base; cutting edge with a large primary cusp, 4—

5 smaller and regularly separated cusps on outer side,

and a small inner side cusp, on base of primary cusp

(Figure 12). Marginal teeth sickle-like, slender, grad-

ually narrowing to sharply pointed tip; secondary cusps,

Page 162 THE NAUTILUS, Vol. 118, No. 4

Figures 2-9. Orbitestella patagonica new species. 2. Holotype, MLP 6367. 3-7. Paratypes, MLP 6368. 3. Dorsal view. 4. Lateral

view. 5. Ventral view. 6. Detail of the protoconch. 7. Operculum. 8. Shell surface sculpture. 9. Shell ultrastructure. Scale bars:

Figures 2-4 = 300 wm; Figures 5 = 500 wm; Figures 6-8 = 100 wm; Figure 9 = 20 wm.

L. R. L. Simone and D. G. Zelaya, 2004 Page 163

Figures 10-11. Orbitestella patagonica. Paratypes, MZSP 38708. 10. Dorsal view of a specimen seen by transparency. 11. Right

view of a specimen removed from shell. Scale bars = 600 jm.

Figures 12-15. Orbitestella patagonica. SEM micrographs of the radula (MZSP 38708). 12. Rachidian (re), lateral (It) and

marginal (mr) teeth (dislodged). 13. Detail of lateral (It) and marginal (mr) teeth. 14. Detail of rachidian teeth. 15. Jaws. Scale

bars = 1 pm.

Page 164 THE NAUTILUS, Vol. 118, No. 4

Figure 16. Orbitestella patagonica. Semi-diagrammatic longitudinal section. Scale bars = 100 ym. Abbreviations in Materials

and Methods.

SOK

NLAT eH"

Figures 17-18. Orbitestella patagonica. Anatomical details. 17. Left side view of a specimen with pallial cavity sectioned and

deflected; 18. Transversal section in middle region of common glandular genital duct. Scale bars: Figure 17 = 100 wm; Figure 18

= 50 pm.

L. R. L. Simone and D. G. Zelaya, 2004

Page 165

absent (Figures 12, 13). Esophagus originating from

middle dorsal region of odontophore, initially narrow,

wider posteriorly, entering stomach at anterior-ventral

region at side of style sac (Figure 16); esophagus wall

thick and undulated by presence of glands. Salivary

glands small, ventral to esophagus and posterior to odon-

tophore. Stomach large occupying about %4 whorl, some-

what flat, on right side of visceral mass (Figures 10, 16).

Inner gastric surface covered by cuticle. Style sac small,

located in antero-dorsal region of stomach, bearing an

ovoid crystalline style. Stomach lumen continuous with

digestive gland. Digestive gland composed by non-pig-

mented yack ells Intestine short (a half of esoph-

agus length), originating between stomach and style sac,

directed forward (Figure 16). Anus located on posterior

end of pallial cavity.

GENITAL sysTEM: Gonad hermaphrodite, located be-

tween stomach and digestive gland, at left side posterior

to pallial cavity, occupying 0.5 whorl (Figures 10, 16).

Male tissue intermingled between female acini. Seminal

vesicle anterior to gonad, very convoluted, with narrow

walls. Common glandular genital duct broad, located on

right side of pallial roof, bulging ventrally into haemo-

coel space; lumen with few longitudinal glandular folds

and short cilia. Prostatic gland well differentiated, ven-

trally located; capsule gland covering outer region of

prostatic gland (Figure 18). Genital aperture and penis

not observed. Oogonia and oocytes in various degrees of

differentiation, randomly distributed (found in two dis-

sected specimens).

CENTRAL NERVOUS SYSTEM: _ Pair of fused cerebro-pleu-

ral ganglia located posterior to buccal bulb; pedal ganglia

located ventral to oral tube. Statocyst of about half eye

volume, immerse anterior to pedal ganglia into pedal

musculature, with a single and relatively large (similar to

eye in diameter) statolith (Figure 16).

Type Material: Holotype (MLP 6367) (720 wm in di-

ameter). Paratypes: 9 specimens (MLP 6368), 5 speci-

mens (MZSP 38708 + 9 slides of serial section of 2

specimens), all from the type locality, March 1998.

Type Locality: 54°52’ S, 68°12’ W, Isla H, Beagle

Channel, Tierra del Fuego, Argentina.

Distribution: Known only from type locality.

Etymology: The species name derives from Patagon-

ia, region on southern part of South America.

DISCUSSION

Three species of Orbitestellidae were previously report-

ed from the Magellanic Region and adjacent subantarc-

tic waters: Microdiscula subcanaliculata (Smith, 1875)

from South Georgia Islands and South Orkneys Islands

(Ponder, 1983 and 1990), Microdiscula vanhoffeni Thie-

le, 1912, from South Shetlands Islands (Dell, 1990) and

Orbitestella ponderi from East Beagle Channel and

southeastern Pacific Ocean, Chile (Linse, 2002). Orbi-

testella ponderi was also collected during this study along

the Beagle Channel at Ushuaia Bay (~ 54°50’ S, 68° W),

Gable iidkerndl (54°54.6' S, 67°21.4’ W) and San Pio Cape

(55°5.6’ S, 66°28.8’ W): at Sloggett Bay (55°00’ S,

66°20.6' W) and Isla de los Estados (54°46’ S, 64°29’

W). These new records confirm that O. ponderi is par-

tially sympatric with O. patagonica.

Orbitestella patagonica differs from O. ponderi Linse,

2002, by having a more depressed shell, with much

weaker axial sculpture and rounded aperture. Orbites-

tella patagonica differs from the Antarctic species of Mi-

crodiscula Thiele (1912) by having a more depressed,

spirally ornamented shell. The ratio 105 the umbilicus/last

whorl diameter in O. patagonica is similar to that of M.

vanhoffeni and greater than in M. subcanaliculata. Or-

bitestella patagonica also differs from the Australian and

New Zealand congeneric species in having a more

rounded shell aperture, more rounded last whorl, and

weakly defined commarginal growth lines. Orbitestella

patagonica also differs from the North Atlantic O. similis

and O. cubana by lacking nodulose sculpture and by hav-

ing a more rounded ees whorl.

The South American species of Orbitestella (O. pon-

deri and O. patagonica) differ markedly from the Aus-

tralasian Orbitestella species (including the type species)

and the North Atlantic species, by having crisp spiral

threads covering the entire shell surface, including the

protoconch. However, based on jaw morphology and

general shell shape, we prefer to expand the generic set

of characters for Orbitestella instead of a introducing a

new generic taxon for the South American species. Be-

yond these two genera, only Lurifax Warén and Bouchet

(2001) has also Then reported to the family Orbitestel-

lidae (Smriglio and Mariottini, 2002). ). Lurifax is very dif-

ferent from the other genera in having taller spire and

multispiral protoconch.

Orbitestella patagonica shows the general anatomical

features of the orbitestellids described by Ponder (1990).

The main anatomical difference is the proportionally

smaller stomach, occupying only the right side of the

visceral mass. Radula morphology also show some dif-

ferences with those previously described and figured by

Ponder (1990) as characteristic for Orbitestella: the pec-

tinated cutting edge of the rachidian tooth in O. pata-

gonica is wider and not as triangular as in O. wareni

eonded 1990; the cutting edge of the lateral teeth were

described by Ponder (1990) as a wide plate with nu-

merous small cusps in the outer margin, while in O.

patagonica they are narrower and having only few cusps;

marginal teeth are wide and numerously cusped in the

species described by Ponder (1990) and are narrow,

hook-like and without additional cusps in O. patagonica.

The jaws in O. patagonica have only five rows of serrated

plates, while Ponder (1990) described six and seven rows

for the species he studied. Both radula and jaws of O.

patagonica are similar to those of O. ponderi.

Orbitestella patagonica, as it was previously reported

for O. wareni by Ponder (1990), has the general anatom-

ical features of a basal heterobranch, such as the lack of

Page 166

ctenidium, the hermaphrodite genital system mostly

bulging into the haemocoel, the lack of odontophoral

cartilages and the presence of a pigmented mantle gland.

ACKNOWLEDGMENTS

The authors wish to express their gratitude to C. Ituarte

for comments on the manuscript and B. A. Marshall for

specimen loans of Microdiscula vanhoeffeni and other

Australian orbitestellid species in the collection of the

Museum of New Zealand Te Papa Tongarewa, and also

for comments on the paper. Part of this study was de-

veloped with the support of funding to the senior author

by Fundacgao de Amparo a Pesquisa do Estado de Sao

Paulo (Fapesp), Process # 00/11074-5 and 00/11357-7.

Diego Zelaya is a Fellow of the National Research Coun-

cil for Science and Technology (CONICET), Argentina.

LITERATURE CITED

Dell, R. K. 1990. Antarctic Mollusca with special reference to

the fauna of the Ross Sea. Bulletin of the Royal Society

of New Zealand 27: 1-311.

Gabe, M. 1968. Techniques histologiques. Masson Editeur,

Paris, 1113 pp.

Hickman, C. S. and J. H. McLean. 1990. Systematic revision

and suprageneric classification of Trochacean gastropods.

Natural History Museum of Los Angeles County, Science

Series 35: 1-169.

Iredale, T. 1917. More molluscan name-changes, generic and

specific. Proceedings of the Malacological Society of Lon-

don 12: 322-330.

Knight, J. B., L. R. Cox, A. M. Kenn, L. R. Batten, E. L.

Vookalgam and R. Robertson. 1960. Systematics descrip-

tions. In Treatise on Invertebrate Paleontology (R. C.

Moore, Ed), I (Mollusca 1), pp. 1169-1310.

Linse, K. 2002. The shelled Magellanic Mollusca: with special

reference to biogeographic relations in the Southern

THE NAUTILUS, Vol. 118, No. 4

Ocean. Theses Zoologicae vol. 74. A. R. A. Ruggell, Liech-

tenstein, 251 pp., 21 pls.

Marshall, B. A. 1988. Skeneidae, Vitrinellide and Orbitestelli-

dae (Mollusca: Gastropoda) associated with biogenic sub-

strata from bathyal depths off New Zealand and New

South Wales. Journal of Natural History 22: 949-1004.

Ponder, W. F. 1967. The classification of the Rissoidae and

Orbitestellidae with descriptions of some new taxa. Trans-

actions of the Royal Society of New Zealand, Zoology 9

(17): 193-224 + pls. 1-13.

Ponder, W. F. 1983. Eatoniellidae, Rissoidae, Cingulopsidae,

Orbitestellidae and Rissoellidae (Mollusca: Gastropoda) of

Signy Island, South Orkney Islands, with a review of the

Antarctic and Sub-Antarctic species. British Antarctic Sur-

vey, Scientific Report 108: 1-96.

Ponder, W. F. 1990. The anatomy and relationships of the Or-

bitestellidae (Gastropoda: Heterobranchia). Journal of

Molluscan Studies 56: 515-532.

Ponder, W. F. and R. G. de Keyzer. 1998. Superfamily Valva-

toidea. pp. 854-858 in P. L. Beesley, G. J. B. Ross, and

A. Wells (eds.) Molltsca: The Southern Synthesis. Fauna

of Australia. Vol. 5. CSIRO Publishing, Melbourne, part

B viii, pp. 565-1234.

Ponder, W. F. and A. Warén. 1988. Appendix. Classification of

the Caenogastropoda and Heterostropha—a list of the

family-group names and higher taxa. In: W. F. Ponder

(ed.) Prosobrach Phylogeny. Malacological Review, Sup-

plement 4, pp. 188-328.

Powell, A. W. B. 1979. New Zealand Mollusca. Marine, Land

and Freshwater shells. Auckland, Collins, 500 pp.

Rolain, E. and F. Rubio. 1992. Two new species of the genus

Orbitestella Trelade, 1917 from the Atlantic Ocean. La

Conchiglia 23(262): 17-20.

Smriglio, C. and P. Mariottini. 2002. Lurifax vitreus Warén and

Bouchet, 2001 (Gastropoda, Orbitestellidae), first report

from Western Mediterranean Sea. Bollettino Malacologico

38: 45-47.

Thiele, J. 1929. Handbuch der systematischen Weichtierkunde.

Fisher, Jena 1: 1-1134.

Warén, A. and P. Bouchet. 2001. Gastropoda and Monopla-

cophora from hydrothermal vents and seeps: new taxa and

records. The Veliger 44: 116-231.

THE NAUTILUS 118(4):167-174, 2004

Page 167

Sphaeriidae (Bivalvia) from Peruvian Amazon floodplains, with

the description of Pisidiwm iquito new species

Cristian Ituarte

Division Zoologia Invertebrados

Museo de La Plata

1900 La Plata, Buenos Aires

ARGENTINA

[email protected]

ABSTRACT

This article reports on the sphaeriid bivalves collected during

a faunistic survey at the Pacaya-Samiria Reserve, in the Peru-

vian Amazon. A new species of Pisidiwm C. Pfeiffer, 1821, from

floodplain habitats, is described. Pisidium iquito new species is

defined by its large, high, not-inflated shell, with trapezoidal

shell outline, beaks prominent, sub-central; the ligament ex-

ternally visible and tending to be protruded, and the presence

of only one branchial opening and one demibranch, are also

diagnostic. The presence of Pisidium sterkianum Pilsbry, 1897,

from small watercourses in the Peruvian Amazon is reported

for the first time. Eupera simoni (Jousseaume, 1889) is re-

ported from the surveyed area, figured, and compared with

other Peruvian and Brazilian species.

INTRODUCTION

As it is the case for large geographic areas of southern

South America, knowledge of the molluscan fauna from

the Amazonian floodplains is rather poor. Particularly

scant is the information from the Peruvian Amazon.

More extensive surveys on the molluscan fauna have

been done in the Brazilian Amazon (Pain, 1960; Bonet-

to, 1967, 1972; Paraense, 1967, 1975). Relevant results

dealing with the composition of the invertebrate assem-

blages in Brazilian Amazonian rainforest have also been

reported by Dreher-Mansur and Valer (1992) and Volk-

mer-Ribeiro et al. (1998). Among Bivalves, the sphaeriid

fauna of Peru and in particular of the Amazonian flood-

plains is virtually unknown. Previous literature records

account mainly for the sphaeriids from high-altitude en-

vironments of the Central Andes (Kuiper and Hinz,

1984). To date, two Pisidiwm species, two species of Eu-

pera, and three species of Sphaerium have been the only

Sphaeriidae reported as living in Peru (Kuiper and Hinz,

1984: Ramirez and Arenas, 2003).

In the present paper, sphaeriids collected during a

faunistic survey in a small area of the Peruvian Amazon

at the Pacaya-Samiria Reserve, a protected area limited

by the Marafién and Ucayali rivers, are reported, and a

new species of Pisidium is described.

MATERIALS AND METHODS

Materials for this study were obtained in September

2002, during the low- water period of the Amazon flood-

plains at the Pacaya-Samiria Reserve, Loreto, Peru (Fig-

ure 1). Sphaeriids were collected from stagnant water

bodies, the so-called “cochas”, flowing watercourses

locally called “cafios” (both habitats were profusely

covered by floating meadows) and from small muddy-

bottom streams. The local term “cocha” designates an

ancient river branch or meander that, due to high rates

of sedimentation, became separated from the main

course, forming ponds of variable sizes.

Sphaeriids were relaxed for study by a brief rinsing in

warm water (approx. 50°C) and immediately fixed in eth-

anol 70°. Specimens for scanning electron microscopy

were cleaned after dissection of soft parts, with a con-

centrated solution of sodium hypochlorite, adequately

mounted and coated. Linear measurements: shell length

(SL), shell height (SH), shell width (SW); shape indices

and morphometric ratios: height index (HI = SH/SL),

convexity index (Ci = SW/SH), and ratio hinge length

(HiL): shell length (HiL/SL), were calculated according

to the criteria fallomed by Ituarte (1996). For each cal-

culation (N = 10), mean and standard deviation values

are given.

For comparative purposes, the following type material

was examined: syntypes of Pisidium sterkianum Pilsbry,

1897, Academy of Natural Sciences, Philadelphia

(ANSP) and paratypes of Pisidium forense Meier-Brook,

1967, Senckenberg Museum, Frankfurt (SMF), Eupera

kKlappenbachi Mansur and Veitenheimer, 1975, Museo

de La Plata (MLP), and Eupera doellojuradoi Klappen-

bach, 1962, Museo Argentino de Ciencias Naturales

(MACN), were used.

SYSTEMATICS

Pisidium iquito new species

(Figures 2-13)

Description: Shell thin, translucent, large (maximum

observed size: 7.2 mm; mean SL of specimens examined:

Page 168

THE NAUTILUS, Vol. 118, No. 4

76°S TASS

---+_@' ()

4°S ;

! ~~ Did

WE aya

ofion gS Wag

Io RO 70

Samiria R. SC REQUENA

hes

e ay

he q iS) cayall

6°S & S

76°S 74°S

| 100 km

Figure I.

5.4+1.2, n = 10), somewhat high (mean HI = 82+2),

not convex (mean Ci = 68 + 4), shell outline sub-qua-

drangular, tending to trapezoidal in medium-sized and

larger specimens, anterior end produced in a short sharp

curve, somewhat pointed, posterior end short, truncated,

nearly straight, somewhat oblique (Figures 2, 4, 5, 11);

dorsal margin gently curved, short, ventral margin uni-

formly aril markedly curved. Beaks prominent, well

raised from shell surface, wide at base, pointed at the

tip, somewhat directed backward, well visible but not

much projected above dorsal margin, slightly displaced

backward, at about 59% of SL. Shell sane finely and

irregularly striated, glossy, straw-yellowish.

Hinge plate weak, hinge line rather long (HiL/SL

about 57%). Hinge: Right valve (Figures 5, 8, 9): car-

dinal tooth (C3) narrow and straight at anterior half,

curved and enlarged in a slightly sulcated cup at poste-

rior end. Lateral “teeth dlendler delicate, inner anterior

lateral (AI) long, slightly curved, cusp displaced forward;

outer anterior lateral tooth (AIII) very short, cusp distal;

inner posterior lateral (PI) straight, cusp sub-central;

outer posterior lateral (PIII) reduced in size, straight,

with distal cusp. Left valve (Figures 4, 6, 7): cardinal

teeth well developed, the inner (C,) robust, blunt, pro-

jected below hinge line, slightly bent upward at tip, with

base slightly oblique with respect to antero-posterior

axis, the outer (C,) a short, flat, curved lame, with distal

Amazonas R.

4°S

6°S

Location map. Encircled area indicates type locality for Pisidium iquito new species.

part slightly bent upward, quite oblique, overlapping C,

at posterior half; anterior lateral tooth (AII) slender,

straight, cusp well displaced distally; posterior lateral

tooth (PII) relatively short, weak, slightly curved, cusp

distal.

Ligament pit long and slender, lanceolate, not deep,

inner margin slightly sinuous at anterior half, enlarged

and uniformly concave at posterior half (Figures 7, 9).

Escutcheon long, lanceolate, marked by a delicate line

(Figure 10). Ligament external, long (representing about

21% of shell length), delicate, well visible from outside,

moderately elevated over dorsal margin or protruded in

variable degrees (Figures 7, 9, 10).

Anatomy: Only the anal mantle opening present, a pair

of well developed, but not particularly strong, siphonal

retractor muscles present (Figure 13). The mantle fusion

anterior to anal siphon not long g, representing 11 + 2%

of SL (Figure 13). Mantle edge broad. Muscle scars (up

to 10 are usually clearly visible) corresponding to the

inner radial mantle muscles well-marked, lying well

apart from the pallial line (Figure 13), those correspond-

ing to anal siphon retractors are coalescent with poste-

rior adductor muscle scars, the one that would corre-

spond to the absent branchial opening (sr;) stronger than

the remaining (Figure 13) that appear grouped in triads

(Figures 4, 5). Only one demibranch, the inner, present.

C. Ituarte, 2004 Page 169

Figures 2-9. Pisidiwm iquito new species. 2-3. Holotype (MLP 6859-5). 2. Right lateral view. 3. Posterior view. 4-9. Paratypes

(MLP 6859-5). 4. Left valve, inner view. 5. Right valve, inner view. 6. Hinge of left valve. 7. Left valve, detail of cardinal teeth

and ligament. 8. Hinge of right valve. 9. Right wall e, detail of cardinal tooth aad ligament. Scale bars: Figures 2-5 = 1 mm; Figures

6-9 = 500 pm.

Page 170

THE NAUTILUS, Vol. 118, No. 4

Figures 10-11. Pisidiwm iquito new species. Paratypes MLP

6859-5. 10. Posterior view showing the external ligament. 11.

Lateral view of a medium sized specimen. Scale bars: Figure

10 = 1 mm; Figure 11 = 500pm.

Brood pouches developing upward and posteriorly in

each inner demibranch, with up to 24 embryos found in

a specimen of 6.5 mm length. Nephridia of closed type,

dorsal lobe elongate with lateral loop visible in dorsal

and posterior and dorsal views (Figure 12).

Type Locality: “Cocha” Tamara, 05°16'28" S,

74°29'55" W, Pacaya-Samiria Reserve, Department of

Loreto, Peru (Figure 1).

Type Material: Holotype, MLP 6859-5; 3 paratypes:

3 MLP 6859-5, 1 MNHN unnumbered.

Other Material Examined: Unnamed small stream

(05°16'12" S, 74°21’27" W) that flows into the Pacaya

River, at the entrance to the Pacaya—Samiria National

Reserve (Figure 1), on muddy soft bottoms, 20 speci-

mens, MLP 6863-1-1.

Etymology: The species name refers to the Iquitos,

ancient people who inhabited the Peruvian Amazon

floodplains.

Remarks: Pisidiwm iquito new species is defined by

its large maximum size, being the largest South Ameri-

can species of Pisidium, sri quadrangular shell shape,

prominent beaks, markedly weak hinge plate, ligament

external, tending to be protruded, mn by the presence

of only one desnflvamah, the outer one, nephridia of

Figures 12-13. Pisidiwm iquito new species. 12. Nephridi-

um. 13, Mantle muscles. (as: anal siphon; irm: inner mantle

radial muscles; pa: posterior adductor muscle; pr: pedal re-

tractor muscle; sr: anal siphon retractor muscle; sr,: retractor

muscle corresponding to the position of lacking branchial

opening. Scale bars = 1 mm.

closed type, with dorsally visible lateral loop, and only

one mantle opening, the anal. Pisidiwm iquito new spe-

cies and Pisidium sterkianum Pilsbry, 1911, have the

same the number of demibranchs, mantle openings, and

type of nephridia; however, Pisidium iquito new species

has a less convex and higher sub-quadrangular shell, with

more prominent and more centrally located beaks; both

species have an external ligament, although much more

protruded in P. sterkianum. Pisidiwm iquito new species

chiefly differs from Pisidium meierbrooki Kuiper and

Hinz, 1984, from the Peruvian highlands and Pisidium

chiquitanum Ttuarte, 2001, from contra Bolivian low-

lands in both shell characters (being larger with no ovate

shell outline) and soft anatomy (having only one demi-

branch and one mantle opening). Pisidium iquito new

species resembles Pisidiwm forense Meier-Brook, 1967,

from the southwestern Brazilian drainage basin, having

a similar siphonal arrangement and only one pair of dle=

mibranchs; however, Pisin iquito new species differs

by its larger size, relatively larger height, less convexity,

and beaks more displaced backwards. Baker (1930). de-

scribed several sphaeriids from northern South America,

among them Pisidium bejumae Baker, 1930, from a sa-

vanna pond in Venezuela, which is similar to Pisidiwm

iquito new species in having a relatively high shell with

centrally located, full, and prominent beaks; however, P

bejumae has smaller size, having higher and more con-

vex, with short, ellipsoid, not trapezoidal, shell outline;

the ligament also differs, not tending to be protruded as

in P. Iquito new species.

Pisidium sterkianum Pilsbry, 1897

(Figures 14—25)

Pisidium sterkianum Pilsbry, 1897: 291-292, pl. 6, figs. 1-4.

Description: Shell of medium size (maximum ob-

, 2004 Page 171

Figures 14-21. Pisidium sterkianum Pilsbry, 1897 (MLP 6863-1). 14. Right lateral view. 15. Posterior view. 16. Left valve, inner

view. 17. Right valve, inner view. 18. Left valve, detail of cardinal teeth aa ligament. 19. Right valve, detail of cardinal tooth and

ligament. 20. Posterior view with detail of external ligament. 21. Lateral view of posterior and with detail of protruded external

ligament. Scale bars: Figures 14-17 = 1 mm; Figures 18-20 = 500 wm; Figure 21 = 100um.

Page 172

THE NAUTILUS, Vol. 118, No. 4

Figures 22-25. Pisidium sterkianum Pilsbry, 1897. 22-23.

Syntypes (ANSP 70490). 22. Right valve, outer lateral view.

23. Right valve, inner view. 24— 25. MLP 5061. 24. Left valve,

inner lateral view. 25. Left valve, outer lateral view. Scale bars:

Figures 22-25 = | mm.

served SL = 5.7 mm), rather inflated (mean CI =

73+2). Shell outline ovate, somewhat high (mean HI =

82+3); dorsal margin slightly shorter than ventral mar-

gin, gently aumevede ventral margin evenly and widely

GUTAEG Anterior end produced in a somewhat acute

curve, posterior end truncated. Beaks wide at base, full,

nearly central (located at about 58% of shell length), well

visible above dorsal mar gin. Surface finely and ‘regular ly

striated.

Hinge vas narrow, not solid. Hinge line long (HiL/

S = 63431). Hinge: Right valve (Figures 17, 19): a mi-

nute, somewhat sreelk cardinal tooth (C 3), enlarged at

posterior end forming a slightly grooved cup; anterior

right lateral teeth elle dev eloped, the inner (AI) a long,

broadly curved lamella, cusp low, somewhat displaced

forward: the outer (AIII) shorter, cusp distal, bent up-

ward. Posterior lateral teeth straight, the inner (PI) nar-

row, cusp sub-central, the outer (PIII) shorter with cusp

distal. Left valve (Figures 16, 18): two well-developed

cardinal teeth, the inner (C,) short, bent upward at tip,

the outer (C,) slightly longer than C,, slender, evenly

arcuate, slightly oblique with respect to antero-posterior

axis, overlapping C, at posterior half. Anterior (AII) and

posterior (PII) lateral teeth well-developed, the anterior

stronger, cusps high, triangular, displaced distally. Liga-

ment pit slender, long, not deep. Ligament external,

markedly protruded and well visible from the exterior

(Figures 20, 21); ligament length about 21% of shell

length.

Anatomy: Only the anal mantle opening present; only

one, the inner, demibranch present, brood pouch de-

veloping from the upper part of inner wall of descending

lamella, up to 8 embryos per demibranch were found.

Seven weakly marked muscle scars, corresponding to in-

ner radial mantle muscles, located slightly above the pal-

lial line, each bundle formed by few weak muscle fibers.

Nephridia of closed type, with lateral loop visible in pos-

terior view.

Type Locality: Fronia creek in the “Prado” (actually

corresponding to Arroyo Miguelete), Montevideo, Uru-

guay.

Material Examined: Unnamed small stream

(05°16'12” S, 74°21'27” W) that flows into the Pacaya

River, at the entrance to the Pacaya-Samiria National

Reserve (Figure 1), on muddy soft bottoms; 27 syntypes

ANSP 70490; > 50 specimens, Arroyo Miguelin (a small

watercourse flowing into the Rio de ‘La Plata), Ensenada

Buenos Aires, Argentina, MLP 5061.

Remarks: Diagnostic characters of Pisidium sterkian-

um are: shell rather solid, medium-sized, inflated, beaks

wide, sub-central; ovate shell outline, expanded forward,

ligament extemal and protruding, only exhalant mantle

opening and only inner demibranch, present; nephridia

of closed type, with lateral loop visible from dorsal view.

When compared with the syntypes of Pisidium ster-

kianum (ANSP. 70490) (Figures 22, 23) and specimens

from the Rio de La Plata, Argentina (MLP 5061) (Fig-

ures 24, 25), the general shell shape of Amazonian spec-

imens is somewhat higher, the shell outline is less oval

with posterior end consistently higher, markedly trun-

cated, and beaks are more outstanding from shell sur-

face.

Eupera simoni (Jousseaume, 1889)

(Figures 26-32)

Limosina simoni Jousseaume, 1889: 217, pl. 9, figs. 22, 23

Eupera simoni Klappenbach, 1967:110; Dreher Mansur and Valer,

1992: 94: Dreher-Mansur and Meier-Brook, 2000: 5.

Description: Shell medium to large (maximum ob-

served SL = 8 mm), relatively low (mean HI = 712),

not inflated (mean Ci = 705), shell outline moderately

oval, posterior end truncate, slightly curved, oblique in

larger specimens, anterior end ev enly rounded, not

pointing (Figures 26—28). Dorsal margin slightly and

evenly arcuate, ventral margin widely curved. Beale tri-

angular, pointed, well mar hedl off from shell surface, but

(oe)

C. Ituarte, 2004 Page |

Figures 26-32. Eupera simoni (Jousseaume, 1889). 26. MLP 6859-6. Right valve, outer lateral view. 27-30. MLP 6864-1. 27.

Right valve, inner lateral view. 28. Left valve, inner lateral view. 29. Right valve, detail of cardinal tooth and ligament. 30. Left

valve, detail of cardinal tooth and ligament. 31. Periostracum folds. 32. Detail of periostracum folds and papillae. Scale bars: Figures

26-28 = 1 mm; Figures 29, 30 = 500um; Figure 31 = 25 wm; Figure 32 = 5 wm.

not prominent, somewhat directed forward; located at hinge plate; anterior lateral teeth short, the inner (AI)

about 33 % of shell length (Figures 26-28). robust, cusp sub-central, the outer (AIII) quite reduced

Hinge plate weak, extremely narrow below beaks, in size; posterior lateral teeth (PI and PIII) delicate,

hinge line relatively long (HiL/SL about 60%). Hinge: slender, almost straight. Left valve (Figures 28, 30): car-

Right valve (Figures 27, 29): cardinal tooth (C;) weak, a dinal tooth (C,;) minute, not very high, anterior (AII)

straight, narrow blade, very close to ventral margin of and posterior (PII) lateral teeth relatively low, slightly

Page 174

THE NAUTILUS, Vol. 118, No. 4

curve, cusps sub-central in AII, nearly distal in PII. In-

ner shell surface with spots of dark pigment sparsely

distributed in small groups, two larger pigmented areas

above and below scar of posterior adductor muscle, usu-

ally present. Periostracum thin, forming commarginal

folds, periostracum surface entirely covered by very

small papillae, up to 0.5 wm long, distributed without a

definite pattern (Figures 26, 31, 32).

Anatomy: The general anatomy fits into the generic di-

agnosis. Worth mentioning is a somewhat marked de-

velopment of the posterior foot retractor muscle, whose

insertion point is well marked just over the scar corre-

sponding to the posterior adductor muscle. Up to 18

embryos were found within each maternal demibranch.

Type Locality: Laguna de Espino, Caracas, Venezue-

la.

Material Examined: Pacaya-Samiria Reserve (Figure

1): Cocha Tamara, 05°16’28” S, 74°29'55” W (MLP

6859-6): Cocha Yarina, 05°24'42” S, 74°30'23" W (MLP

6865-1), attached to roots of Eichornia sp.; Pacaya Riv-

er, 5°16'55" S, 74°25'45" W (MLP 6886-1), on roots of

floating meadows; Cafio Yarina, 05°21'28" S, 74°30'29"

W (MLP 6864-1).

Remarks: Eupera simoni, a common species in the

Peruvian Amazon, is easily identified by the somewhat

shortened shell outline due to the posterior truncated

margin, beaks small, pointed, relatively low but well dis-

cernible from shell surface, and periostracum raised in

well-defined commarginal folds. Eupera primei Klap-

penbach, 1967, described from water courses close to

the Ucayali River, Peru, not far from Pacaya-Samiria Re-

serve, differs from E. primei in having a higher shell,

with shell outline tending to be circular, dorsal margin

strikingly curved, and very low periostracal folds. Eupera

klappenbachi Mansur and Veitenheimer, 1975, a species

common in western Brazilian drainages, is similar to E.

simoni, consistently differing in being higher and more

convex, and having more prominent and pointed beaks;

Eupera platensis Broly Jurado, 1921, from the Rio de

La Plata has a more solid shell, with more centrally lo-

cated beaks, stronger cardinal teeth and robust laterals.

Eupera simoni is quite different from Eupera guarani-

ana Ituarte, 1994, from the Uruguay River, a large spe-

cies with a striking trapezoidal shell outline, and also

differs from Eupera elliptica Ituarte and Mansur, 1993,

from the Iguazi River, Northern Argentina, a species

characterized by its strikingly elliptic shell outline. Eu-

pera guaraniana and E. elliptica also differ from E. si-

moni in having two types of periostracal papillae, the

larger ordered in radial rows.

ACKNOWLEDGMENTS

Materials for the present study were collected during the

first field trip (September 2002) to the Pacaya-Samiria

Reserve, Peru as a part of the Project “Aquatic Faunal

Survey of a Peruvian Amazon Floodplain” (Florida Mu-

seum of Natural History/Universidad Nacional de la

Amazonia Peruana), funded by National Science Foun-

dation of the United States (NSF-DEB 0215388 to J.

Albert, Principal Investigator). M. C. Dreher Mansur

kindly helped with the identification of Ewpera simoni,

for which the author is greatly indebted. The author is

researcher of the Consejo Nacional de Investigaciones

Cientificas y Técnicas (CONICET), Argentina.

LITERATURE CITED

Baker, B. 1930. The Mollusca collected by the University of

Michigan-Williamson expedition in Venezuela. Occasional

Papers of the Museum of Zoology 210: 1-90.

Bonetto, A. A. 1967. La superfamilia Unionacea en la cuenca

Amazonica. Atas do Simpésio sébre a Biota Amaz6nica 3

(Limnologia): 63-82:

Bonetto, A. A. 1972. A new species of Monocondylaeinae from

the Amazon basin, and some considerations on this sub-

family in the hydrographic systems of South America.

Amazoniana 3: 224-230,

Dreher-Mansur, M. C. and R. M. Valer. 1992. Moluscos bi-

valves do rio Uraricoera e rio Branco, Roraima, Brasil.

Amazoniana 12: 85-100.

Dreher-Mansur, M. C. and C. Meier-Brook. 2000. Morphology

of Eupera Bourguignat 1854, and Byssanodonta Orbigny

1846 with contributions to the phylogenetic systematics of

Sphaeriidae and Corbiculidae. Archiv fiir Molluskenkunde

128: 1-59.

Ituarte, C. F. 1996. Argentine species of Pisidiwm Pfeiffer,

1821, and Musculiwm Link, 1807 (Bivalvia: Sphaeriidae).

The Veliger 39: 189-203.

Jousseaume, F. 1889. Voyage de M. Eugéne Simon au Vene-

zuela. Mollusques. Mémoires de la Societé zoologique de

France 2: 232-259, pl. 239.

Klappenbach, M. A. 1967. Eupera primei sp. n. de la regién

del rio Ucayali, Pert (Mollusca, Pelecypoda). Atas do Sim-

posio sobre a Biota Amaz6nica (Limnologia) 3: 109-115.

Kuiper, J. G. J. and W. Hinz. 1984. Zur Fauna der Kleinmus-

cheln in den Anden. Archiv fiir Molluskenkunde 114

[1983]: 137-156.

Pain, T. 1960. Pomacea (Ampullariidae) of the Amazon River

System. The Journal of Conchology 24: 421-432.

Paraense, W. L. 1967. Moluscos planorbideos da Amazonia.

Atas do Simpésio sébre a Biota AmazOnica 3 (Limnologia):

187-194.

Paraense, W. L. 1975. Estado atual da sistematica dos planor-

bideos brasileiros (Mollusca, Gastropoda). Arquivos do

Museu nacional, Rio de Janeiro 55: 105-128.

Pilsbry, H. A. 1897. New species of mollusks from Uruguay.

Proceedings of the Academy of Natural Sciences of Phil-

adelphia, May 1897: 290-298, 2 pl.

Ramirez, R. C. and J. Arenas. 2003. Moluscos del Peri. Revista

de Biologia Tropical suppl. 3: 225-292.

Wollamanitthetvo, C. M., M. C. Dreher Mansur, P. A. S. Mera

and S. M. Ross. 1998. Biological indicators in the aquatic

habitats of the ha de Maraca. In: Milliken, W. and J. A.

Ratter (eds.) Maraca: The Biodiversity and Environment

of an Amazonian rainforest. John Wiley & Sons, London.

THE NAUTILUS 118(4):175-176, 2004

Note

Page 175

Dates of publication of Yoichiro Hirase’s Kai Chigusa

Paul Callomon

Department of Malacology

Academy of Natural Sciences

1900 Benjamin Franklin Parkway

Philadelphia, PA 19103-1195 USA

[email protected]

Yoichiro Hirase (1859-1925) set himself up in business

from 1887 as a shell dealer in Kyoto, Japan. Over the

following 33 years, he built one of the world’s first shell

museums and produced and published the ‘Concholog-

ical Magazine’ together with a number of books on the

Mollusca. Perhaps the best known among the latter is

Kai Chigusa, a title often erroneously cited as Kai Sen

Shu. Sen shu is the nominal Japanese reading of the

Kanji characters for ‘thousand’ and ‘kinds’, but Chigusa

is a classical reading that is admittedly unfamiliar even

to many modern-day Japanese. Hirase studied Chinese

classics in his youth, however, and frequently used clas-

sical or poetic readings of Kanji characters.

Published between 1914 and 1922, Kai Chigusa was

a four-volume work that portrayed 400 predominantly

Japanese mollusks in hand-colored woodblock prints. It

was bound in the traditional Japanese ‘concertina’ style.

Ten volumes were originally planned, depicting a total

of 1000 species (whence the English title, “The illustra-

tions of a thousand shells’), but straitened economic cir-

cumstances forced a downsizing of the project after 1915

(Callomon and Tada, in prep).

A collation of the work was provided by Higo et al.

(1999: 581). Four new species were figured and named

in it: Mandarina omphalina Hirase, 1915, Calliostoma

akoya Hirase, 1922, Gaza (?) nobilis Hirase, 1922 and

Fulgoraria kaneko Hirase, 1922. Complete sets of all

four volumes are relatively rare, and cognoscenti have

long suspected that the more common earlier volumes

were printed in larger numbers than the later ones.

The author recently came across a collection of cor-

respondence between Alvin R. Cahn and R. Tucker Ab-

bott of the Academy of Natural Sciences of Philadelphia

(ANSP) that spans the years 1947-1968. Cahn was a

fisheries scientist who worked for the Natural Resources

Division of Supreme Headquarters Allied Forces in To-

kyo for several years from 1947. His work brought him

into contact with many of the leading Japanese marine

scientists of the day, including Tokubei Kuroda (1886—

1987), Hirase’s one-time assistant and museum manager.

During a long stay in Japan, Cahn built up an extensive

library and a collection of mollusk specimens, both of

which were donated in a series of gifts to the ANSP that

Figure 1. Cover of the first volume (1914) of Kai Chigusa.

started in 1957. His library included a fine complete set

of Kai Chigusa, together with some notes he had made

regarding the production of this work. In 1955, Cahn

wrote to the Unsodo company, the publishers of Kai

Chigusa, requesting information about its publication.

After a long wait he received a noncommittal reply. In

August 1957 he invoked his authority as a former rep-

resentative of the occupation forces to press Unsodo's

president, Mr Shigeji Yamada, for information. Yamada

replied on September 7" of that year with a full account

compiled from company records. This has apparently

Page 176

THE NAUTILUS, Vol. 118, No. 4

Table 1. Dates of publication and press runs of Kai Chigusa.

Volume 1

First printing 300 copies May 1, 1914

Second printing 100 copies December 24, 1914

Third printing 100 copies January 17, 1917

Fourth printing 100 copies September 6, 1919

Fifth printing 100 copies August 17, 1921

Sixth printing 100 copies April 2, 1925

Seventh printing 50 copies June 12, 1930

Highth printing 30 copies March 28, 1937

Total: 880 copies

Volume 2

First printing 300 copies January 10, 1915

Second printing 100 copies August 22, 1917

Third printing 100 copies September 6, 1919

Fourth printing 100 copies February 28, 1922

Fifth printing 100 copies February 21, 1926

Sixth printing 50 copies March 13, 1935

Total: 750 copies

Volume 3

First printing 300 copies November 10, 1915

Second printing 100 copies September 6, 1919

Third printing 100 copies July 28, 1922

Fourth printing 100 copies February 21, 1926

Fifth printing 50 copies November 20, 1933

Total: 650 copies

Volume 4

First printing 300 copies January 15, 1922

Second printing 50 copies October 26, 1929

Third printing 50 copies March 13, 1935

Total: 400 copies

never been published, and the main information is re-

produced here. Interestingly, Yamada also used the er-

roneous reading Kai Sen Shu.

From the figures provided to Cahn, it is clear that

demand for Kai Chigusa persisted long after its author's

death in 1925. Nevertheless, the suspicion that the num-

ber of copies printed of each volume varies is clearly

Figure 2. A plate from the first volume (1914) of Kai Chi-

gusd.

grounded in fact. There are more than twice as many

copies of the first volume than of the last. Yamada con-

firmed that despite Kai Chigusa’s long life in print, the

date of printing given in the colophon was never

changed. It is thus not possible to determine which

printing any given copy represents.

In a letter of April 27, 1957 Cahn told R. Tucker Ab-

bott: “Incidentally, all the original woodblock plates are

still intact; I have seen them. They probably number

well into the thousands, there being often as many as a

dozen or more for a single shell print: they occupy an

entire room from the floor to the ceiling. A represen-

tative of Unsodo has recently confirmed that the blocks

continue to exist, though reassembling the plates for

printing would now involve prohibitive expense.

LITERATURE CITED

Callomon, P. and A. Tada. Yoichiro Hirase and his Role in

Japanese Malacology. In prep.

Higo, S., P. Callomon and Y. Goto. 1999. Catalogue and bib-

liography of the marine shell-bearing mollusca of Japan.

Elle Scientific Publications, Yao, 749 pp.

Hirase, Y. 1914-1922. Kai Chigusa/The Illustrations of a

Thousand Shells. 4 vols. Unsodo, Kyoto.

THE NAUTILUS

Volume 118

2004

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AUTHOR INDEX

ANRIDITUAL, "IN|g 1th aoodaapabeoncoussdeusuasosaetteoaradanesnoocE eee nse 131 lbruyNistiiah: (Cus sneangoseaauboounouvabeesbcomsasedsedsuognarmecauaoanosy 167

ISTaRINIEIIE, IDs. No: oscoscossososconoo ove oosooobasosoadasonaveaboononaye 127 IAIN TOR es Veee eee eet ere cece cree tan cere one one re 1

(GALINDO: (Ga ScoenchoadadesbononeuaeecunddcGee MeEnnte mmnnaeeeeermrnoe 129 IMINNorios: (Ges segonta deauenanswoeadcoateease inant aaeeatonesenn ocean 121

(GINTEIUOIMION, 1B, .c00ansasosncnsaesnnnssosson9sq000g9aneeseceecsscoane: 175 INES EIN SRS Viagaec qantas unartuce nea moana game nan iaubaeauacinsaeiaee 88

CAZZANICA, N. Joes cceeceeeeseetee tsetse eee tes a ORTUZ Ate inte et Pha ens (ili eee Bac seiner tetanus ae 103

COLES, Boies 55 IPD AIDA IKE JOM ana asepanaetovereeaneraean aaa conte onet esos Sean oa 127

(CHonigiiia), R16 nednaddoneanoqadeee nou aneseassescenE Gea sierenae aaAmEe!” A iC PERCE RUA rine rete de wit he ire a ein te 103

ANTS, 1B Co cacsoasceareosseeazessvesosgcosbees0ceeasa020sbs2c00030% Ian FRORGIUDS INIE Tieassecassaacde enemel tel Pecan te matt 103

IDNGIBIRUR:, IE a ddbaseasosnesuneebee eros Genk occ oe aBem US RAC On Sine ae aeeee 152

TPunLiGney; WULieoe a eusceedhechers ae aan ne Oe nen TD] —-SEMIONIB, Hr Ne Ihe wcveoscocososvssasozo csanszeysesescucusesaocnsees Let

GALEATIDO, (Cr ShicddadoneneeeseonenGonsensen tne neBBeE eae RERe noses 121 SLAPCINSKY, Jo voes cece ee ceeeer ese reese ner serene erreeess ee

Garnett, IR [ished cbieemes ncn ane eee) enn 139, 144 TIRONCOSO), Je Sooo eco e saeco e eae eee sansa: 139

Giammaicars IML Se ssacdasscossakoseatocs foneasons Hon bescaaeemencnne Fall WRGORRT Wore oases se so elsnyessee esse seer 144

GonALEZWAN@UIaMIDI, IMI, .oocoosoosoosoopconenodenanpecnnos 103 (VATED) EEA ee RE NOE, Aa eho en in MOU me ea A De NE 131

(GROVE SSE Ma RE eee ee es EN Re cc die atin! 43 WISI, Co Joocacvosses0000s000s0cspsnccussssea000sc0000ss0a0000006 157

TRLARVAG TEN AYGIEI, IMIG Goa enon oe bonnie dsekabacosssudeannanateecenans 1, 93 ZAENANB IS: a Esl Cis goauanonucdounsoneorcunvere mane docem Tom ae rOnetnDS 144

ipliomipminin, (GL 'Sy coopncoadasdaceb boaoeedeuaucdacddennsooraancanacsas 157 TATUNG DR Cue bu snae hoebe lontsanees easenane cobansooH na en ne 112, 160

NEW TAXA PROPOSED IN VOLUME 118 (2004)

GASTROPODA

Anetarca brasiliana Garcia and Troncoso, 2004, new species (Facelinidae) .................0 cece cece eeceeee ce eeet ee eeseeeeeceeneseee ees 139

Armina elongata Ardila and Valdés, 2004, new species (Arminidae) ................c. ccc eceeee crete eet ee ee eeee eee eneneeseeeenneeeeeeenes 136

Bernaya (Bernaya) beardi Groves, 2004, new species (Cypracidae) ........0....0....ceeeeeeeeeeeseeeeeeeseeeeeseceueseseusssetneseteuseseees 47

Bennayay(Bernaya) yeanae Groves, 2004, mew, species (G@ypracidae)) -pecyasseeeasseneneeee eee rescence cece ee eneeeeeee ae: 46

Bernaya (Protocypraea) popenoei Groves, 2004, new species (Cypraeidae) ............ 500... c cece ee eee cee eeeceeeesteneeeteeeeeeeeesene ees 47

Cimon: cirrikum Dem, AUOL, mewy Gaeeies (BOO ANCE) ,...00056000000000000000000509000000000000040000a0000d8d00000eNABB0a99000009040" 154

Coluzea kallistropha Harasewych, 2004, new species (Turbinellidae) ..............0..ccee cece eee eee eee e cet eeeeete ree eeeeeeeteeeeeeees 93

Coluzea madagascarensis Harasewych, 2004, new species (Turbinellidae) ...........0....0. 0c cece eeeeeee cee ee cee eect eeeeeeeeeeeeeeenes 95

Coluzea naxa Harasewych, HOA, MSw FoSCies (MwrorinllWGlAS)) nc.sooscccacscaceoscaconosaaoaoacoosaoagoabacoonoacoacasossoscncoqscooondcoonee 100

Columbarium quadrativaricosum Harasewych, 2004, new species (Turbinellidae) .............0.....0ccceceee teeter este eeee te eeee eee 98

Drepanodontus Harasewych and Kantor, 2004, new genus (Buccinulidae) .........00......0000ee cece eee eect esses eee seen neeeeeee ees 7

Drepanodontus tatyanae Harasewych and Kantor, 2004, new species (Buccinulidae) ..................0ccce cc cee cette eeee tees te eeeeee ees 7

Germonea Harasewych and Kantor, 2004, new senus\(Buccimulidas)) ish sesesaehae caeeermac asst kates eninseortnanad sasels aaspeam esi 20

Germonea rachelae Harasewych and Kantor, 2004, new species (Buccinulidae) .............. 0.00... 0 cee cence ete ee ee eee eee ester eee tees 90

Lusitromina Harasewych and Kantor, 2004, new genus (Buccinulidae) ..............0.....00cesseeeseeeee tees ecteeeeteneeeeteeeettneseeenes 205

Muffinbuccinum Harasewych and Kantor, 2004, new genus (Buccinulidae) ......0.0.....0.0..ee cece eee e ett eee ete eeee eter neeeeee ees 13

Muffibuccinum catherinae Harasewych and Kantor, 2004, new species (Buccinulidae) .....................cccee tee eee eeeee et eeee ees 14

Ohvancillariamatanzana Nielsen, 2004, new species) (Olividae)) Pov. 3.yssceecos.s sores esse eeeo ess sees sess seen ssseeneenee seen 90

Orbitestella patagonica Simone and Zelaya, 2004, new species (Orbitestellidae) ...........0......0 cece eee eect eeee teste teeeee eee eeeees 161

Palaeocypraea (Palaeocypraea) wilfredi Groves, 2004, new species (Cypraeidae) ........... 5.60. c cece eee ee eee eeee en eeeeeeeeeeeeeeeen es 45

Pilsbryna nodopalma Slapcinsky and Coles, 2004, new species (Gastrodontidae) ..........0......00 0 cece cette eee eter eeee tern eeeeeeees 61

Pilsbryna quadrilamellata Slapcinsky and Coles, 2004, new species (Gastrodontidae) ..............0..2...0cceeeee eet settee eeeeeeeeees 63

Spikebuccinum Harasewych and Kantor, 2004, new genus (Buccinulidae) ............00.... ccc bce e eee eter e eee e een eens 2

Spikebuccinum stephaniae Harasewych andl Kantor, 3004, MEW FOSGISS (BUETMUIUCHS)) ...coorsancanceooasseecaovosssecooenssesacse0080000 2

Stramonita penelaevis Vermeij and Herbert, 2004, new species (Muricidae) ...............00...cccee cee eee ee eeeee sees eeeeeeeeeeeeseen ees 157

POLYPLACOPHORA

Leptochiton (Leptochiton) troncosoi Zalvide et al., 2004, new species (Leptochitomidae) ..................0. ccc eee ceeeee etter 144

Leptochiton (Leptochiton) pepezamorai Zalvide et al., 2004, new species (Leptochitonidae) .................0.0e cc eeeeteeeeeeeeee es 147

BIVALVIA

Pista connie Newent, A004, mew GaeCles (GDMAETNGES)) ..000000200000000200000000000040600000200000000000000090080005000000000008900030000000 167

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CONTENTS

Newel 1. US

Volume 119, Number 1

Kevin J. Roe

Paul D. Hartfield

Russell L. Minton

Steven P. Savarese, Jr.

Frank Kéhler

Matthias Glaubrecht

John Slapcinsky

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Claude Vilvens

March 28, 2005

ISSN 0028-1344

Hamiota, a new genus of freshwater mussel (Bivalvia:

Unionidae) from the Gulf of Mexico drainages of the

Goutneasienn Write] States. oo ccccccccccdcoaccnvdvavc:

Consideration of genetic relationships in management

decisions for the endangered Anthonys riversnail, Leptoxis

crassa anthonyi (Redfield, 1854) (Gastropoda:

PLSUTOCETIG AC) Beery ees eee aoe nee aa ta aap re

Fallen into oblivion—the systematic affinities of the

enigmatic Sulcospira Troschel, 1858 (Cerithioidea:

Pachychilidae), a genus of viviparous freshwater gastropods

ITO MM) AVA pis ahah eal alas Mrathacet mys Gern alae nt di Spertecaat ms sas 8

Six new species of Paryphantopsis (Gastropoda: Pulmonata:

Charopidae) from the Papuan Peninsula of New Guinea...

First record of Akera Miiller, 1776, from the eastern

Pacific, with the description of a new species ............

Description of Calliotropis pulvinaris new species

(Gastropoda: Trochidae: Eucyclinae: Calliotropini) from

WestiMadapascartvicnisnsist os waar tomes bcnmatascetls

Seen eat pen eee I

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IN OU CO RE Sera ee eee Re ee erro nr or ae re eR pes isso ease MOR Log tg ets owt orig at tam pe coe eae 04

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THE NAUTILUS 119(1):1-10, 2005

Page 1

Hamiota, a new genus of freshwater mussel

(Bivalvia: Unionidae) from the Gulf of Mexico drainages

of the southeastern United States

Kevin J. Roe

Delaware Museum of Natural History

4840 Kennett Pike

Wilmington, DE 19807 USA

[email protected]

Paul D. Hartfield

U.S. Fish and Wildlife Service

6578 Dogwood View Parkway, Suite A

Jackson, MS 39213 USA

[email protected]

ABSTRACT

Hamiota, a new genus of freshwater mussel containing four

species formerly assigned to the genus Lampsilis Rafinesque,

1820, is described. In addition to the genus Lampsilis, mem-

bers of Hamiota had previously been placed in the genera Vil-

losa Frierson, 1927, and Ligumia Swainson, 1840. Several char-

acters including the packaging of their larvae in a superconglu-

tinate lure to attract host fishes, placement and shape of the

marsupia, and release of glochidia through the excurrent si-

phon, support the recognition of these species as a distinct

genus.

INTRODUCTION

Following the discovery in 1988 by Robert Butler of the

first superconglutinate lure ensnared on a snag in a trib-

utary of the Choctawhatchee River, a number of publi-

cations (e.g., Haag et al., 1995; Hartfield and Butler,

1997; O’Brien and Brim Box, 1999; Blalock-Herod et al.,

2002) have confirmed through direct observation the

supposition that these unique lures are produced by four

species of freshwater mussels endemic to the Gulf of

Mexico drainages of the southeastern United States.

Herein, we confirm earlier published suggestions that

these four species represent a distinct genus of fresh-

water mussels (Fuller and Bereza, 1973; O’Brien and

Brim Box, 1999). The recognition of this genus is based

on several characters including the production of the su-

perconglutinate lure, and the unique shape and place-

ment of the marsupia (the region of the demibranchs

where female unionoid mussels brood developing lar-

vae), and is supported by molecular evidence (Roe et al.,

2001). Use of marsupial features is consistent with pre-

vious designations of unionid “divisions” initiated by

Simpson (1900a) and continued by Heard and Guckert

(1970). The designation of Hamiota increases the num-

ber of North American unionids genera to 50 (Turgeon

et al., 1998).

MATERIALS AND METHODS

A list of specimens examined is included in Appendix 1.

Acronyms used in the text are: Academy of Natural Sci-

ences of Philadelphia (ANSP), Delaware Museum of

Natural History (DMNH), Florida Museum of Natural

History (UF), United States National Museum (USNM),

University of Alabama Unionid Collection (UAUC), and

Mississippi Museum of Natural Science (MMNS). Ab-

breviated synonymies are presented for each taxon and

include novel combinations and publications with illus-

tration. Measurements were taken to the nearest 0.05

mm using dial calipers.

SYSTEMATICS

Family Unionidae Rafinesque, 1820

Tribe Lampsilini von Ihering, 1901

Hamiota new genus. Type species: Hamiota subangulata (Lea,

1840) by original designation

Diagnosis: A monophyletic group of freshwater bi-

valves (Roe et al., 2001) in which all of the glochidia are

released simultaneously encased in mucous packages

that are referred to as superconglutinates (Haag et al.,

1995, fig. 1). The superconglutinate lure exits the mantle

cavity via the excurrent opening and is encased within a

transparent mucous tube (Hartfield and Butler, 1997;

O’Brien and Brim Box, 1999). When acted upon by wa-

ter currents the superconglutinate mimics the move-

ments of a swimming fish, and has been shown to elicit

attacks from fishes (Haag and Warren, 1999). The mar-

supium is restricted to the ventral portion of the outer

demibranchs of female mussels (Figure 1). The precise

shape and pigmentation of the marsupia, as well as the

degree of posterior mantle margin development, varies

across species.

Description: Members of this genus are small- to me-

dium-sized freshwater bivalves, and adult valves gener-

ally are between 45-100 mm in length. Shells range

from ovate to elliptical in outline, and are somewhat

Page 2

THE NAUTILUS, Vol. 119, No. 1

Figures 1, 2.

Inner mantle and outer demibranchs of gravid superconglutinate-producing mussels in the genus Hamiota and

inner mantle and outer demibranchs typical of non-superconglutinate-producing mussels of the genera Lampsilis, Ligumia, and

Villosa. 1. Female Hamiota australis. 2. Female Villosa vibex.

compressed to moderately inflated. Shell thickness rang-

es from heavy to thin. Sexes display some degree of di-

morphism in ‘shell shape. Shells of male marcel are typ-

ically more acutely pointed posteriorly, whereas shells of

female mussels display an expanded posterior margin.

Periostracum is typically smooth, but can be very glossy

in some species. Background color ranges from dark-

brown and black through chestnut- brown to straw- yel-

low. Black to bright green rays of variable width are of-

ten present and may be limited to the posterior slope,

or cover the entire disk. Nacre color is typically white

although other colors such as salmon or blue may be

seen as well, particularly in the beak cavities or the pos-

terior margins. The marsupia are often asymmetrical in

shape; the anterior portion is typically broadest, tapering

toward the posterior end. The ventral margin of the mar-

supium is darkly pigmented in gravid females. Pigmen-

tation of marsupia varies across species and populations

and colors may include purple, red, black, or white. In

females, the mantle margins anterior to the branchial

opening are elaborated to varying degrees. The mor-

phology of the glochidial valves of members of Hamiota

is similar to that of members of Lampsilis or Villosa

(O’Brien and Brim Box, 1999).

Etymology: Hamiota = angler. Derived from the

Greek word hamus, meaning hook. This name refers to

the means by which members of this genus attract host

fishes by packaging their parasitic larvae in a lure that

mimics a small fish.

Remarks: Species of Hamiota generally have been

treated as Lampsilis due to the similarity in shape and

coloration of their shells. Some authors also have in-

cluded these species in the genus Villosa or Ligumia,

due to shell shape, thickness, and/or ornamentation of

the mantle flap. The shells of Hamiota species are in-

K. J. Roe and P. D. Hartfield, 2005

Page 3

deed similar to the shells of these genera, and the man-

tle flap may exhibit characters of both Lampsilis and

Villosa. However, in Hamiota, the marsupia is restricted

to the ventral half of the posterior portion of the outer

demibranchs, while in Lampsilis, Liguwmia, and Villosa

the marsupium fully occupies the water tubes of the pos-

terior portion of the outer demibranchs (Figure 1). Un-

like members of Lampsilis, which release larvae through

pores in the ventral edge of the demibranchs, members

of Hamiota release the superconglutinate lure through

the excurrent opening. The placement and shape of the

marsupium and the extraordinary method of glochidial

release and host fish attraction, the superconglutinate,

are uniquely derived characters in the Lampsilini that

warrant genus-level recognition.

Hamiota altilis (Conrad, 1834) new combination

Unio altilis Conrad, 1834; Conrad, 1834: 43, pl. 2, fig. 1;

Chenu, 1845: 21, pl. 1, fig. 1; Reeve, 1865: pl. 23, fig. 109.

Margarita (Unio) altilis (Conrad, 1834); Lea, 1836: 24.

Margaron (Unio) altilis (Conrad, 1834); Lea, 1852a: 27.

Lampsilis altilis (Conrad, 1834); Simpson, 1900a: 529; Par-

malee and Bogan, 1998: 125, pl. 47.

Unio clarkianus Lea, 1852: Lea, 1852b: 251: Lea, 1852c: 273,

pl. 21, fig. 30; Lea, 1852d: 29, pl. 21, fig. 30.

Margaron (Gnio) clarkianus (Lea, 1852); Lea, 1852a: 27.

Lampsilis clarkianus (Lea, 1852); Simpson, 1900a: 532.

Unio gerhardtii Lea, 1862; Lea, 1862a: 168; Lea, 1862b: 208,

pl. 31, fig. 277; Lea, 1862c: 30, pl. 31, fig. ie

Margaron (Unio) gerhardtii (Lea, 1869). Lea, 1870: 35.

Lampsilis (Lampsilis) g gerhardtii (Lea, 1862); Simpson, 1900a:

532.

Unio doliaris Lea, 1865; Lea, 1865: 88; Lea, 1868: 260, pl. 32,

fig. 75; Lea, 1869: 20, pl. 32, fig. 75.

Margaron (Unio) doliaris (Lea, 1865); Lea, 1870: 42.

Lampsilis (Lampsilis) doliaris (Lea, 1865); Simpson, 1900a:

533.

Description: Described by Conrad (1834) as sub-oval,

thin, and inflated. The periostracum was described as

“rugose” and “blackish” with rays and “numerous short

vermicular lines on the posterior slope,” and the nacre

as whitish and iridescent. The periostracum of speci-

mens of H. altilis is typically brown to chestnut-brown

in color with a variable number of dark green rays. The

left valve has two heavy, spatulate pseudocardinal teeth,

the smaller above the larger. The right valve has two

nearly equally sized triangular teeth, the larger anterior

to the smaller. The lateral teeth are short but blade-like,

two in the right valve, and one in the left.

The posterior mantle margin of the female is expand-

ed into a well-developed flap with papillae along the bor-

der. Coloration, number, and size of papillae vary some-

what between populations. In general, however, the in-

terior mantle flap is colored red to dark red or brown

with darker spots, while the exterior of the flap is brown

to black, often with vertical lighter bars, and with a

small, but prominent dark “eye spot” on the posterior

end. Small papillae are present along the mantle flap,

usually becoming more robust anteriorly. In males, the

mantle flap is typically not expanded and is reddish in

color. Marsupia of H. altilis are finely tapered at each

end when immature, becoming broadly rounded on the

ends in most populations, tapering anteriorly in others.

Marsupia color is a dark reddish-brown or black along

the margin and white above. The anus is usually pig-

mented el and the incurrent and excurrent siphons are

reddish-brown to black. The glochidia of H. altilis are

described by Haag et al. (1999).

Type Material: Unio altilis Conrad, 1834, Lectotype

ANSP 56419 (Figures 3, 4) here designated. Type lo-

cality: Alabama River, near Claibome [Monroe Co., Al-

abama].

Unio clarkianus Lea, 1852, Type not found. Type local-

ity: Williamsport, [Maury Co.], Tennessee; Georgia or

Alabama.

Unio doliaris Lea, 1865, Lectotype USNM 84936, here

designated. Type locality: Etowah River, Georgia.

Unio gerhardtii Lea, 1862, Holotype USNM 25711 by

monotypy. Type locality: Chattanooga, Georgia.

Remarks: The most variable species included in Ham-

iota, H. altilis, is endemic to the Mobile River Basin.

Some of the conchological variation is undoubtedly eco-

phenotypic in nature, although the extent and nature of

the variation in shell shape. nd pigmentation has not

been adequately explored.

Conrad (1834) in his original description did not iden-

tify a primary type. Johnson and Baker (1973) identified

ANSP 56419 as the figured holotype, although the spec-

imen label indicates the locality as “Ogeechee R., Ga.”

Johnson and Baker (1973) state that the label is in error,

and “probably was mixed with ANSP 46418, which is

labeled ‘Claiborne, Alabama’, by error.” Conrad (1834)

clearly indicated that the specimen(s (s) of U. altilis de-

scribed were collected from the “Alabama River, near

Claiborne.” Conrad (1834) did not specifically designate

a holotype and according to the ICZN recommendation

73F and Article 74.5 dhe holotype designation of John-

son and Baker (1973) is deemed relic In ordér to

preserve stability of nomenclature, we herein designate

specimen ANSP 56419 as the lectotype of U. altilis. In

accordance with ICZN Article 74.7, we herein designate

the specimen USNM 84936 as the lectotype of U. do-

liaris Lea in order to maintain taxonomic stability and

because this specimen appears to be that figured by Lea

(1868). Hamiota altilis is considered threatened by the

United States Fish and Wildlife Service (USFWS, 1994).

Life History: Mature gravid females have been re-

ported from March through June. Hamiota altilis have

also been observed releasing g glochidia in a superconglu-

timate (Haag et al., 1999). ihe ge centrarchid fishes, in-

cluding Micropterus coosae Hubbs and Bailey, 1940, M.

punctulatus (Rafinesque, 1819), M. salmoides (Lace-

péde, 1802), and Lepomis cyanellus Rafinesque, 1819,

have been confirmed as suitable hosts (Haag et al.,

1999).

Range: Hamiota altilis was historically reported

Page 4 THE NAUTILUS, Vol. 119, No.

Figures 3-10. Type material of species of Hamiota. Photographs are of the interior of left valve and exterior of right valve. 3,

4. Lectotype of H. altilis ANSP 56419. 5, 6. Holotype of H. australis USNM 150473. 7, 8. Lectotype of H. perovalis ANSP 56416.

9, 10. Lectotype of H. swbangulata USNM 85801.

K. J. Roe and P. D. Hartfield, 2005

throughout the Mobile River Basin, including the Tom-

bigbee, Black Warrior, Cahaba, Alabama, Tallapoosa, and

Coosa River drainages in Alabama, Georgia, Mississippi,

and Tennessee. The species is currently restricted to lo-

calized portions of the Cahaba, Coosa, and Tallapoosa

rivers and some of their tributaries (USFWS, 2003).

Hamiota perovalis (Conrad, 1834) new combination

Unio perovalis Conrad, 1834; Conrad, 1834: 43, pl. 2, fig. 2;

Chenu, 1845: 21, pl. 1, fig. 2; Kiister, 1861: 257, pl. 87,

fig. 2; Reeve, 1866: pl. 38, fig. 209.

Margarita (Unio) perovalis (Conrad, 1834); Lea, 1836: 24.

Margaron (Unio) perovalis (Conrad, 1834); Lea, 1852a: 27.

Lampsilis perovalis (Conrad, 1834); Simpson, 1900a: 531.

Unio spillmanii Lea, 1861; Lea, 1861: 39, Lea, 1862d: 98, pl.

15, fig. 246; Lea, 1862e: 102, pl. 15, fig. 246, Reeve, 1868:

pl. 82, fig. 435.

Margaron (Unio) spillmanii (Lea, 1861); Lea, 1870: 42.

Lampsilis (Lampsilis) spillmani (Lea, 1861); Frierson, 1927: 69

[misspelling].

Description: Conrad (1834) described this species as

oval and inflated with a moderately thick shell. He noted

two color varieties, one in which the periostracum was

olivaceous and obscurely rayed with white nacre and an-

other i in which the periostracum was reddish-brown with

“rose colored” nacre. The periostracum of specimens of

H. perovalis is generally lighter in color than H. altilis

and range from straw-yellow to light brown. The number

of rays is variable and can cover the entire disk. The left

valve contains two robust equal sized pseudocardinal

teeth. The right valve has two pseudocardinals and the

anterior tooth is smaller than the posterior tooth. The

lateral teeth are elongate, two in the left, one in the

right.

The mantle margins of female H. perovalis are ex-

panded into well-developed flaps, pigmented red on the

interior and darker red to brown or black on the exterior.

No eyespot is present and short papillae are present

along the mantle edge. Males possess a rudimentary

mantle margin with weak pigmentation and few papillae.

The marsupia of H. perovalis are pisciform in shape,

broader anteriorly and narrowly tapering posteriorly.

The marsupium is reddish or darker along the margin,

often with a darker spot of pigment on the broader an-

terior end that resembles an eyespot in the supercon-

glutinate lure. The anus can be pigmented red and

black, and the incurrent and excurrent siphons are usu-

ally reddish or brown in color.

Type Material: Unio perovalis Conrad, 1834, Lecto-

type ANSP 56416 (Figures 7, 8), here designated. Type

locality: Alabama River, at Claiborne [Monroe Co., Al-

abama].

Unio spillmanii Lea, 1861, Lectotype USNM 84925,

here designated. Type locality: Luxpalila Creek, near Co-

lumbus, Mississippi.

Remarks: There has been some question as to the dis-

tinctiveness of H. perovalis from H. altilis, perhaps be-

Page 5

cause both were described from practically the same lo-

cality. An examination of mitochondrial DNA sequences

by Roe et al. (2001) recovered these two taxa as a clade,

but failed to resolve them into reciprocally monophyletic

groups. Hurd (1974) considered perovalis a junior syn-

onym of altilis, as did Burch (1975). Frierson (1927)

considered U. doliaris (Lea, 1865) a synonym of perov-

alis, although Parmalee and Bogan (1998) include U. do-

liaris as a synonym of altilis. Based on the collection

locality and the appearance of the type specimen, we

place U. spillmanii Lea, 1861 as a synonym of H. per-

ovalis. As with U. altilis, Conrad (1834) did not specifi-

cally designate a holotype for U. perovalis, therefore,

according to the ICZN Article 74.5 and recommendation

73F the holotype designation of Johnson and Baker

(1973) is deemed to be in error. In an effort to maintain

nomenclatural stability we herein designate ANSP 56419

the lectotype for U. perovalis. According to ICZN Arti-

cle 74.7, the specimen USNM 84925 is here designated

as the lectotype of U. spilmanii in order to fix the name

and maintain nomenclatural stability. This species is list-

ed as threatened by the United States Fish and Wildlife

Service (USFWS, 1994).

Life History: Discharge of superconglutinates was

first observed in H. perovalis (Haag et al., 1995). Glo-

chidia mature and are discharged Bemveen March and

June, with releases concentrated in early April (Hartfield

and Butler, 1997). Micropterus coosae, M. punctulatus,

and M. salmoides have been identified as suitable host

fishes for the orange-nacre mucket (Haag and Warren,

1999).

Range: Hamiota perovalis was historically known from

the Mobile Basin’s Alabama, Tombigbee, Black Warrior,

and Cahaba rivers and their tributaries in Alabama and

Mississippi. The species has apparently become extir-

pated from the main channels of the larger rivers, but

continues to survive in some tributaries of all four drain-

ages (USFWS, 2003).

Hamiota subangulata (Lea, 1840) new combination

Unio subangulatus Lea, 1840; Lea, 1840: 287; Lea, 1842a: 209,

pl. 13, ‘fig. 23; Lea, 1842b: 47, pl. 8, fig. 23; Kiister, 1861:

578, pl. 94, fig. 2; Simpson, 1892: 415, pl. 58, fig. 1; Reeve,

1868: pl. 65, fig. 327.

Margaron (Unio) subangulatus (Lea, 1840); Lea, 1852a: 29.

Lampsilis subangulatus (Lea, 1840); Simpson, 1900a: 556;

Clench and Turner, 1956: 196, pl. 2, fig. 2.

Ligumia subangulata (Lea, 1840); Haas, 1969: 443.

Villosa subangulata (Lea, 1840); Heard, 1979: 44.

Unio kirklandianus S. H. Wright, 1897; S. H. Wright, 1897:

136.

Lampsilis kirklandianus (S. H. Wright, 1897); Simpson, 1900a:

557; Simpson, 1900b: 76, pl. 1, fig. 7

Description: A medium-sized mussel that reaches ap-

proximately 85 mm in length (Brim Box and Williams,

2000). Specimens are generally elongate; the posterior

ridge is rounded and the posterior slope is usually con-

Page 6

cave. Periostracum color is variable in this species. In-

dividuals range from straw-yellow to chestnut-brown in

color with a variable number of black to bright emerald

green rays of variable width. These rays emanate from

the umbo and radiate across the disk. Most shells are

shiny, although some populations exhibit an extremely

glossy periostracum. This species has been described as

one of the most beautiful of all North American fresh-

water mussels (S. H. Wright, 1897; Clench and Turner,

1956).

Wright (1897) described U. kirklandianus from the

Ochlockonee River in Leon County, Florida and re-

marked that his specimens were more polished and had

broader rays than typical of U. subangulatus, and that

the shells were “deeper and broader.” The right valve

has two somewhat spatulate pseudocardinal teeth, the

smaller nearly directly above the other. The left valve

has two pseudocardinal teeth, the anterior one much

larger than the other. Lateral teeth are thin, but not

delicate, two in the left and one in the right valve. Ex-

amination of specimens from across the range of this

species reveals substantial variation in shell color and

size. The mantle margins in females are only slightly ex-

panded into a modest flap. The flaps are light brown in

coloration on the interior and freckled-brown on the ex-

terior, and no eyespot is present. Short papillae are pres-

ent along the margin of the flap, becoming larger ante-

riorly. In males, the mantle is only slightly expanded,

without pigment, and with very short papillae. The mar-

supia are pisciform in shape, broader anteriorly and ta-

pered behind, and darkly pigmented along the margin

often with a darker spot of pigment anteriorly. The anus

is unpigmented and the siphons are brownish. Glochidia

were figured and described by O’Brien and Brim Box

(1999).

Type Material: Unio subangulatus Lea, 1840, Lecto-

type USNM 85801 (Figures 9, 10) designated by Clench

and Turner (1956). Type locality: Chattahoochee River,

Columbus, [Muscogee Co.,| Georgia.

Unio kirklandianus S. H. Wright, 1897, Paratype USNM

149648. Type locality: Oclocknee River, Leon Co., Flor-

ida.

Remarks: Hamiota subangulata is listed as an endan-

gered species (USFWS, 1998). The analysis of Roe et al.

(2001) resolved this taxon as monophyletic and in a clade

with H. australis.

Life History: Discharge of superconglutinate lures

has been documented by O’Brien and Brim Box (1999)

from late May through early June. Micropterus punc-

tulatus and M. salmoides appear to be primary hosts for

the species (O’Brien and Brim Box, 1999).

Range: Hamiota subangulata was found throughout

the Apalachicola River Basin and the Ochlockonee River

drainage (Brim Box and Williams, 2000). Currently the

species continues to survive in some small streams and

headwater rivers (USFWS, 1998).

THE NAUTILUS, Vol. 119, No. 1

Hamiota australis (Simpson, 1900) new combination

Lampsilis australis Simpson, 1900; Simpson, 1900a: 544; Simp-

son, 1900b: 75, pl. 2, fig. 2; Clench and Tumer, 1956: 199,

. 2, fig, 3.

Ligumia australis (Simpson, 1900); Haas, 1969: 432.

Villosa australis (Simpson, 1900); Heard, 1979: 44.

Description: Simpson (1900b) described the shells of

this species as “long and elliptical . .. moderately inflat-

ed.” The periostracum was described as “smooth, shin-

ning, greenish yellow, rayed with green” and the nacre

as “bright bluish white.” Specimens examined were el-

liptical to elongate oval and often terminate in a blunt

point. Specimens are almost always stained black, with

some green rays visible on the posterior slope. The per-

iostracum is often glossy. The right valve contains two

pseudocardinal teeth, the smaller above the larger. The

left valve has two compressed pseudocardinal teeth. Lat-

eral teeth are elongate and slightly curved, two in the

left valve and one in the right. Posterior mantle flaps are

poorly developed with a streak of red stain along the

margins. There is no eyespot on the mantle flap, and

only a few very small papillae. The marsupia are pisci-

form, broadly rounded anteriorly and tapering behind

(Figure 1). Color of the marsupium is white along the

margin and black above. The anus is unpigmented and

the incurrent and excurrent siphons are reddish in color.

Glochidia were described and figured by Blalock-Herod

et al. (2002).

Type Material: Lampsilis australis Simpson, 1900,

Holotype USNM 150473 by original designation (Fig-

ures 5, 6). Type locality: Little Patsiliga Creek, south-

eastern Alabama.

Remarks: Fuller and Bereza (1973) stated that this

species represented an “undescribed lampsiline genus”

and that its marsupium allied it closely with Ptychobran-

chus. This species was incorrectly synonymized with Pty-

chobranchus jonesi (van der Schalie, 1934) by Clench

and Tumer (1956). The phylogenetic analysis of Roe et

al. (2001) clearly placed this species with the other su-

perconglutinate producers.

Life History: Superconglutinate releases have been

documented in this species by Blalock-Herod et al.

(2002). Micropterus spp. are likely hosts (Blalock-Herod

et al., 2002).

Range: Hamiota australis was known historically from

the Escambia, Yellow, and Choctawhatchee River sys-

tems. It continues to survive in some river and stream

segments within these systems. This species is not cur-

rently protected under the Endangered Species Act.

ACKNOWLEDGMENTS

We express thanks to the many individuals who have

worked with this unique group of mussels and directly

or indirectly assisted in the production of this work. Sev-

eral museum curators provided access to the specimens

K. J. Roe and P. D. Hartfield, 2005

in their care including the National Museum of Natural

History (Dr. Robert Hershler), The Florida Museum of

Natural History (Dr. Fred Thompson), The Academy of

Natural Sciences of Philadelphia (Dr. Dan Graf), The

University of Alabama Malacology Collection (Dr.

~ Charles Lydeard), and the Mississippi Museum of Nat-

ural Science (Dr. Robert Jones and Leann Staton). The

comments of several individuals greatly improved this

manuscript: We are grateful to Art Bogan, Jeffrey Gar-

ner, Dan Graf, Jeanne Serb, and Jim Williams. Gary

Bloomer (DMNH) assisted with figures.

LITERATURE CITED

Blalock-Herod, H. N., J. J. Herod and J. D. Williams. 2002.

Evaluation of conservation status, distribution, and repro-

ductive characteristics of an endemic Gulf Coast fresh-

water mussel, Lampsilis australis (Bivalvia: Unionidae).

Biodiversity and Conservation 11: 1877-1887.

Brim Box, J. and J. D. Williams. 2000. Unionid mollusks of the

Apalachicola basin in Alabama, Florida, and Georgia. Bul-

letin of the Alabama Museum of Natural History 21: 1—

143.

Burch, J. B. 1975. Freshwater Unionacean clams (Mollusca:

Pelecypoda) of North America. Malacological Publica-

tions, Hamburg [Michigan], 204 pp.

Chenu, J. C. 1845. Bibliothéque Conchyliologique. 1** série,

vol. 3. A. Franck, Paris, 153 pp., 34 pls.

Clench, W. J. and R. D. Tumer. 1956. Freshwater mollusks of

Alabama, Georgia, and Florida from the Escambia to the

Suwannee River. Bulletin of the Florida State Museum 1:

97-239.

Conrad, T. A. 1834. New fresh water shells of the United

States, with coloured illustrations; and a monograph of the

genus Anculotus of Say; also a synopsis of the American

wetadk, Judah Dobson, Philadelphia, 76 pp. + § plates.

Frierson, L. S. 1927. A classified and annotated check list of

the North American naiads. Baylor University Press,

Waco, 111 pp.

Fuller, S. L. H. and D. J. Bereza. 1973. Recent additions to

the naiad fauna of the eastern Gulf Drainage (Bivalvia:

Unionoida: Unionidae). Association of Southeastern Biol-

ogists Bulletin 20: 53.

Haag, W. R., R. S. Butler and P. D. Hartfield. 1995. An ex-

traordinary reproductive strategy in freshwater bivalves:

prey mimicry to facilitate larv. al. : dispersal. Freshwater Bi-

ology 34: 471476.

Haag, W. RB. and M. L. Warren. 1999. Mantle displays of fresh-

water mussels elicit attacks from fish. Freshwater Biology

42. 35-40.

Haag, W. R., M. L. Warren and M. Shillingsford. 1999. Host

fishes and host attracting behavior of Lampsilis altilis and

Villosa vibex (Bivalvia: Unionidae). American Midland

Naturalist 141: 149-157.

Haas, F. 1969. Superfamilia Unionacea. Das Tierreich (Berlin)

88: x + 663 pp.

Hartfield, P. and R. Butler. 1997. Observations on the release

of superconglutinates by Lampsilis perovalis (Conrad

1834). In: Cummings, K.S., A. C. Buchanan, C. A. Mayer,

and T. J. Naimo (eds.) Conservation and management of

freshwater mussels II: Initiatives for the future. Proceed-

ings of a UMRCC symposium, 16-18 October 1995, St.

Louis, Missouri. Upper Mississippi Conservation Com-

mittee, Rock Island, Illinois, pp. 11-14.

Heard, W. H. 1979. Identification manual of the freshwater

clams of Florida. State of Florida Department of Envi-

ronmental Regulation, Technical Series, vol.4, no. 2, 83

Dp.

Heart W. H. and R. H. Guckert. 1970. A re-evaluation of the

recent Unionacea (Pelycepoda) of North America. Mala-

cologia 10: 333-355.

Hubbs, C. L. and R. M. Bailey. 1940. A revision of the black

basses (Micropterus and Huro) with descriptions of four

new forms. Miscellaneous Publications Museum of Zool-

ogy, University of Michigan 48: 1-51.

Hurd, J. C. 1974. Systematics and zoogeography of the Union-

acean mollusks of the Coosa River Drainage of Alabama,

Georgia and Tennessee. Unpublished Ph.D. dissertation,

Auburm University.

Johnson, R. I. and H. B. Baker. 1973. The types of Unionacea

(Mollusca: Bivalvia) in the Academy of Natural Sciences

of Philadelphia. Proceedings of the Academy of Natural

Sciences of Philadelphia 125: 145-186, pls. 1-10.

Kiister, H. C. 1861. In: Systematisches Conchylien Cabinet von

Martini und Chemnitz. 2"* ed.

Lacepéde, B. G. E. 1802. Histoire naturelle des Poissons,

vol. 4.

Lea, I. 1836. A Synopsis of the family of Naiades. Carey, Lea,

and Blanchard, Philadelphia, 59 pp., 1 pl.

Lea, I. 1840. Descriptions of new freshwater and land shells.

Proceedings of the American Philosophical Society of

Philadelphia 1: 284-289.

Lea, I. 1842a. Description of new freshwater and land shells.

Transactions of the American Philosophical Society 8[new

series]: 163-250, pls. 5-27.

Lea, I. 1842b. Observations on the genus Unio, together with

descriptions of new species in the families Naiades, Col-

imacea, Lymnaeana, Melaniana and Peristomiana. 3: 1—

88, pls. 5-27.

Lea, I. 1852a. A synopsis of the family of Naiades. Philadelphia.

3rd edition, Blanchard and Lea, Philadelphia, 88 pp.

Lea, I. 1852b. Descriptions of new species of the family Union-

idae. Proceedings of the American Philosophical Society

of Philadelphia 5: 251-252.

Lea, I. 1852c. Descriptions of new species of the family Union-

idae. Transactions of the American Philosophical Society

10[New Series]: 253-294, pls. 12-29.

Lea, I. 1852d. Observations on the genus Unio, together with

descriptions of new species in the families Unionidae, Col-

imacea and Melaniana 5: 9-61, pls. 30.

Lea, I. 1861. Descriptions of twenty-five new species of Union-

idae from Georgia, Alabama, Mississippi, Tennessee and

Florida. Proceedings of the Academy of Natural Sciences

of Philadelphia 13: 38-41.

Lea, I. 1862a. Descriptions of ten new species of Unionidae

from the United States. Proceedings of the Academy of

Natural Sciences of Philadelphia 14: 168-169.

Lea, I. 1862b. New Unionidae of the United States and arctic

America. Journal of the Academy of Natural Sciences of

Philadelphia 5[New Series]: 187-216, pls. 24-33.

Lea, I. 1862c. Observations on the genus Unio, together with

descriptions of new species, their soft parts, and embry-

onic forms, in the family Unionidae, and descriptions of

new genera and species of the Melanidae. 9: 9-178, pls.

24-38.

Lea, I. 1862d. New Unionidae of the United States. Journal of

Page §

THE NAUTILUS, Vol. 119, No. 1

the Academy of Natural Sciences of Philadelphia 5 [New

Series]: 14-109, pls. 1-18.

Lea, I. 1862e. Observations on the genus Unio, together with

descriptions of new species, their soft parts, and embry-

onic forms, in the family Unionidae. 8: 9-114, 34 pls.

Lea, I. 1865. Descriptions of eight new species of Unio from

the United States. Proceedings of the Academy of Natural

Sciences of Philadelphia 17: 88-89.

Lea, I. 1868. New Unionidae, Melanidae, etc., chiefly of the

United States. Journal of the Academy of Natural Sciences

of Philadelphia 6 [New Series]: 249-302, pls. 29-45.

Lea, I. 1869. Observations on the genus Unio, together with

descriptions of new species in the family Unionidae, and

descriptions of new species of the Melanidae and Palu-

dinae. 12: 9-105, 26 pls.

Lea, I. 1870. A synopsis of the family of Naiades. Philadelphia,

4th edition, H. C. Lea, Philadelphia, 184 pp.

OBrien, C. and J. Brim Box. 1999. Reproductive biology and

juvenile recruitment of the Shinyrayed Pocketbook,

Lampsilis subangulata (Bivalvia: Unionidae) in the Gulf

Coastal Plain. American Midland Naturalist 142: 129-140.

Parmalee, P. W. and A. E. Bogan. 1998. The Freshwater Mus-

sels of Tennessee. The University of Tennessee Press,

Knoxville, 328 pp.

Rafinesque, C. S. 1819. Prodrome de 70 nouveaux generes

d’animaux découverts dans lintérieur des Etats-Unis

dAmérique, Durant l'année 1818. Journal de Physique,

de Chimie, d'Histoire Naturelle, et des Arts, Paris 88:

417-429.

Rafinesque, C. S. 1820. Monographie des coquilles bivalves

fluviatiles de la Riviere Ohio, contenant douze generes et

soixante-huit espéces. Annales générales des sciences Phy-

siques, a Bruxelles 5: 287-322.

Reeve, L. 1865-1868. Monograph of the genus Unio. Con-

chologica Iconica. L. Reeve and Co., London.

Roe, K. J., P. D. Hartfield and C. Lydeard. 2001. Phylogeo-

graphic analysis of the threatened and endangered super-

conglutinate-producing mussels of the genus Lampsilis

(Bivalvia: Unionidae). Molecular Ecology 10: 2225-2234.

Simpson, C. T. 1892. Notes on the Unionidae of Florida and

the southeastern United States. Proceedings of the United

States National Museum 15: 405-436, pls. 49-74.

Simpson, C. T. 1900a. Synopsis of the naiads or pearly fresh-

water mussels. Proceedings of U.S. National Museum 22:

501-1044.

Simpson, C. T. 1900b. New and unfigured Unionidae. Pro-

ceedings of the Academy of Natural Sciences of Philadel-

phia 52: 74-86.

Swainson, W. 1840. A treatise on malacology or the natural

classification of shells and shell-fish. London, 419 pp.

Turgeon, D. D., J. F. Quinn, Jr, A. E. Bogan, E. V. Coan, F.

G. Hochberg, W. G. Lyons, P. M. Mikkelsen, C. F. E.

Roper, G. Rosenberg, B. Roth, A. Scheltema, M. J. Swee-

ney, F. G. Thompson, M. Vecchione and J. D. Williams.

1988. Common and scientific names of aquatic inverte-

brates from the United States and Canada: Mollusks, 274

edition. American Fisheries Society, Special Publication

26, Bethesda, Maryland, 526 pp.

U.S. Fish and Wildlife Service. 1994. Endangered and threat-

ened wildlife and plants, 50 CFR 17.11 and 17.12. Divi-

sion of Endangered Species, U. S. Fish and Wildlife Ser-

vice, Washington, D.C.

U. S. Fish and Wildlife Service. 1998. Endangered and threat-

ened wildlife and plants, Determination of endangered

status for five freshwater mussels and threatened status

for two freshwater mussels from eastern Gulf slope drain-

ages of Alabama, Florida, and Georgia. Federal Register

63: 12664-12687.

U.S. Fish and Wildlife Service. 2003. Endangered and threat-

ened wildlife and plants; proposed designation of critical

habitat for three threatened mussels and eight endangered

mussels in the Mobile River Basin; proposed rule. Federal

van der Schalie, H. 1934. Lampsilis jonesi, a new naiad from

southeastern Alabama. The Nautilus 47: 125-127.

von Ihering, H. 1901. The Unionidae of North America. The

Nautilus 15: 37-39, 50-53.

Wright, S. H. 1897. Contribution to the knowledge of the Unit-

ed States Unionidae. The Nautilus 10: 136-139.

APPENDIX 1

SPECIMENS EXAMINED

Hamiota altilis

ALABAMA

Alabama River: USNM 25948: Alabama River, near Clai-

borne: ANSP 56419 [Lectotype U. altilis Conrad, 1834],

ANSP 56418; Big Swamp Creek, Macon Co.: USNM

361723; Jackson Shoals, Choccolocco Creek: ANSP

103834, ANSP 103871; Beaver Creek: ANSP 103863;

Coosa River: ANSP 41120, DMNH 130623; Coosa Riv-

er, Coosa Valley: ANSP 103771; Higgin’s Ferry, Coosa

River, Chilton Co.: USNM 218118; McGowen’s Ferry,

near Wilsonville, Coosa River: USNM 521359: Weduska

Shoals, Coosa River: ANSP 48001, DMNH 075252,

DMNH 150037, DMNH 150038, USNM 348970,

USNM 452169; Coosa River, [incomprehensible hand-

writing] Shoals, Shelby Co.: ANSP 341399; Shoal Creek,

Pine Glen Recreational Area, Cleburne Co.: MMNS

7743, MMNS 8084, MMNS 8085, UAUC 120, UAUC

121, UAUC 125; Yellowleaf Creek, Jumbo, Chilton Co.:

ANSP 89031; Cane Creek, West of CR 65, 2 mi. West

CR 78 Jet., T15S, RIE, sec 3: UAUC 3292, MMNS

8081; Little Cane Creek, at CR 78, East of Edwardsville:

UAUC 3293; Chewacla Creek at CR 22, ~4 mi. East of

Tuskeegee, Macon Co.: UAUC 246, UAUC 247, UAUC

248; Uphappee Creek, 0.5 mi. upstream of Hwy. 29,

Macon Co.: MMNS 8082; Cahaba River: USNM

152026; Cahaba River: ANSP 126054; Little Cahaba

River, 0.5 mi. below Cahaba Beach Rd. bridge, Jefferson

Co.: UAUC 149; Coosa River, Weduska Shoals, Shelby

Co.: UF 3255; Coosa River at Fort William Shoals, Tal-

ladega Co.: UF 65420; Hurricane Creek, Cherokee Co.:

UF 175098; Chewacla Creek, 8 mi. ESE of Tuskeegee

Co. Rd. 22, Macon Co.: UF 202249: Shoal Creek, St.

Clair Co.: UF 245989; Tuskeegee National Forest, Ma-

con Co.; UF 266048: Cahaba River: UF 269576; Shoal

Creek, St. Clair Co.. MMNS 8083.

GEORGIA

Etowah River: USNM 84936 [Lectotype, U. doliaris

Lea, 1865]; Chattooga River: ANSP 89102; Chattooga

K. J. Roe and P. D. Hartfield, 2005

River, Chattooga Co.: USNM 59527; Chattanooga:

USNM 25711 [Holotype, U. gerhardtii, Lea, 1862]; Fish

Creek at Highway 278/ GA State Rt. 6, ~3.9 air mi.

West of Rockmart, Polk Co.: UAUC 538, UAUC 539;

Conasauga River at Tibbs Bridge Murray CR 109 (Whit-

field CR100), Murray/Whitfield Co’s.: UAUC 515,

MMNS 8092; Conasauga River, Muskrat Midden, Tri-

togonia Shoals (CRM 46.70), Whitfield/Murray Co’:

UAUC 376; Conasauga River, south of state line, Murray

Co.: MMNS 8090; Conasauga River: DMNH 150124,

USNM 84937, USNM 348969: Etowah River: USNM

123202.

TENNESSEE

Conasauga River: DMNH 014683; Conasauga River,

Conasauga: ANSP 341305, ANSP 347949; Conasauga

River, Polk Co.: MMNS 8091.

Hamiota perovalis

ALABAMA

Alabama River: ANSP 56416 [Lectotype U. perovalis

Conrad, 1834], ANSP 333496: Alabama River, Clai-

borne: USNM 84938: Coosa River: ANSP 56415; Coosa

River, Gadsen: ANSP 126051; Coosa River, Talladega

Co.: ANSP 126048; Black Warrior River: ANSP 88483:

Mulberry River [Fork, Black Warrior River]: ANSP

88485: Rush Creek [Black Warrior River Dr.], FS Rt.

245, Winston Co.: MMNS 7745, MMNS 8088, UAUC

426; Flannigan Creek at FS Rd. 229, Lawrence Co.:

MMNS 7744, UAUC 423, UAUC 424, UAUC 425:

Toadvine, Valley Creek, Black Warrior River Dr., Jeffer-

son Co.: UF 65302, UF 65304; Forks of the Warrior

River, Walker Co.: UF 65305; North River, near Hagler’s

Mill, Black Warrior Dr., Tuscaloosa Co.: UF 65306; Sip-

sey Fork at N.F. 234, Bankhead National Forest, Win-

ston Co.: UF 79069, UF 79072, UF 79082, UF 79085,

UF 79136, UF 79137; Brushy Creek above Brushy Lake

Recreational Area, Bankhead National Forest, Winston

Co.: UF 79094; Capsey Creek, 50 mi. from Jct. with

Brushy Creek, Bankhead N.F., Winston Co.: UF 79115;

Sipsey Fork at mouth of Hurricane Creek, Bankhead

National Forest, Winston Co.: UAUC 95; Brown Creek,

Bankhead National Forest, Winston Co.: UAUC 1774:

Alabama: ANSP 126049: North River, near Samantha

[Black Warrior River], Tuscaloosa Co.: UAUC 107; But-

tahatchee River, Hamilton: ANSP 100657, DMNH

075231: Tuscaloosa Co. Alabama: DMNH 146496; Black

Warrior River, Jefferson Co.: UF 269609; Squaw Shoals,

Black Warrior River, Jefferson Co.: UF 65298, UF

65299, UF 65300, UF 65303, UF 65307, UF 65429, UF

269518; Garden City, Mulberry Fork: UF 69207, UF

244558; Banks of Brushy Creek, at N.F. Rd. 255, Bank-

head N.F, Winston Co.: UF 69269, UF 79177, UF

79178, MMNS 7748, MMNS 8089; Sipsey Fork, 1 mile

N. of AL Hwy. 33 crossing, Winston Co.: UF 69279;

Brushy Creek at N.F. Rd., Bankhead N.F., Winston Co.:

Page 9

UF 79076; Sipsey River at Sipsey Recreational Area,

Bankhead N.F., Winston Co.: UF 79089: Borden’s

Creek, 1 mi., upstream of from Jct. Sipsey Fork, Sipsey

Wildemess, Bankhead National Forest, Winston Co.::

UF 79092, UF 79181; Sipsey Fork at N.F. Rd., 234,

Bankhead National Forest, Winston Co.: UF 79139:

Limestone Creek, 6.3 mi WNW of Monroeville, Monroe

Co.: UF 197636; Blackwater Creek upstream from Har-

ris bridge, Walker Co.: UF 266369; Sipsey River, 1.6 mi

N. of Pleasant Ridge, Greene Co.: UF 197671; North

River at Co. Hwy. 30, Fayette Co.: UF 197686; Sipsey

River, 200 m. below Co. Hwy 23, Greene Co.: UF

197552; Sipsey River, 4—6 mi. below Co. Hwy. 2, Greene

Co.: UF 197566; Sipsey River at CR 2, downstream of

boat ramp, Pickens Co.: UAUC 156; Tombigbee River:

ANSP 126053; Elrod, Sipsey River, Tombigbee R.:

DMNH 146493: Lubbub Creek, at CR 24, 3.25 mi.

Northeast of Aliceville, Pickens Co.: UAUC 67; Tombig-

bee River: USNM 159989; Lubbub Creek, 1.8 mi SSE

of Aliceville above Hwy 14: UF 197619, UF 197632;

Sipsey River, 3.6 mi. W. of Jena downstream of CR 2,

Greene Co.: UF 197697; Sipsey River near confluence

with Carpenter's Creek, Greene Co.: UF 197801; Sipsey

River, 5.7 mi. NNE of Mantua, Greene Co.: UF 197857,

UF 197862; Sipsey River, near Elrod, Tuscaloosa Co.:

UF 269559, UF 65301; Trussels Creek at CR 19 bridge,

Greene Co.: MMNS 8087.

MIssIsSIPPI

Luxpalila Creek, near Columbus: USNM 84925 [Lec-

totype, U. spillmanii Lea, 1861], USNM 123279; Colum-

bus, Lowndes Co.: UF 269560.

Hamiota subangulata

ALABAMA

Uchee Creek, Russell Co.: UAUC 116.

FLORIDA

Chipola River, Look-Tremble Falls near Alpha, Calhoun

Co.: ANSP 175750; Chipola River, near Pole Bluff land-

ing, Calhoun Co.: ANSP 175751; Chipola River: ANSP

84324: Ochlockonee River: DMNH 150098; Ochlocko-

nee River, Leon Co.: USNM 149648 [Paratype, U. kirk-

landianus S. H. Wright, 1897]; Ochlockonee River, Tal-

lahassee, Leon Co.: ANSP 156892, ANSP 341307; Och-

lockonee River, 7 mi. west of Tallahassee: ANSP 157553;

Ochlockonee River, 10 mi. west of Tallahassee, Leon

Co.: ANSP 159126: Ochlockonee River, 11 mi. north-

west of Tallahassee: DMNH 119506; Ochlockonee Riv-

er, 2 mi. west of Bloxham, Liberty Co.: ANSP 360553;

Spring Creek, Marianna: ANSP 160210: State Rt. 167,

1 mi. north of Marianna, Chipola River, Jackson Co.:

ANSP 349631; Spring Creek, Reynoldsville, Seminole

Co.: UF 177; 1 mi. north Marianna, Chipola River: UF

390; 3.5 mi. east of Quincy, Little River: UF 415; ca. 2

Page 10

mi. east of Clarksville, Chipola River, Calhoun Co.: UF

418; Chipola River, 9.2 km ENE Kinard, 12.5 km NW

Lewis, 16.4 km N. Ida, Calhoun Co.: MMNS 8099.

GEORGIA

Chattahoochie River: ANSP 56477; Chattahoochie Riv-

er, Columbus: USNM 85081 [Lectotype, U. subangula-

tus Lea, 1840]; ANSP 126272; Cooleewahee Crests 0.9

mi. NE of Newton, Baker Co.: USNM 853746; Coolee-

wahee Creek at GA Rt. 91, Baker Co.: MMNS 8095;

Abram’s Inlet, Flint River: ANSP 190294: Mill Creek,

Flint River, several mi. north of Albany: ANSP 267572;

Kinchafoonee Creek at GA Rt. 41 crossing, just south of

Preston, Webster Co.: UAUC591; Kinchafoonee Creek

at GA Rt. 49 bridge ~9 air mi. northeast of Dawson,

Terrell/Sumter Co’s.. UAUC 602, UAUC 603, UAUC

604; Kinchafoonee Creek at GA Rt. 32, Lee Co., GA:

MMNS 8096: Chickasawhatchee Creek at CR 130

bridge ~4 air mi. SW of Chickasawhatchee, Terrell Co.:

UAUC 1753; Muckalee Creek at GA Rt. 195 bridge

~3.5 air mi. Northeast of Leesburg, Lee Co.: UAUC

312; Whitewater Creek on Morgan Mill Rd., Fayette

Cos UAUC 645; Ochlockonee River: DMNH 173390;

Ochlockonee River, 7 mi. S. of Cairo: ANSP 194640,

DMNH 048538, UF 412; Mimsville: ANSP 47892,

DMNH 075151; Georgia: ANSP 126273; Calvary: ANSP

47891; Ochlockonee River, Thomas/Grady Co's. Georgia:

MMNS 8101; Spring Creek at GA Rt. 84, Decatur Co.:

MMNS 8094, MMNS 8100: Line Creek at GA Rt. 85/

THE NAUTILUS, Vol. 119, No. 1

74 bridge, Coweta/Fayette Co.: MMNS 8097; Ichaway-

nochaway Creek at GA Rt. 216, Baker Co.: MMNS

8098.

Hamiota australis

ALABAMA

Andrews fish trap, Pea River, Barbour Co.: UF 65309;

7 mi. east of Brundidge, Pea River, Pike Co.: UF

123284; Bozemans landing, Conecuh River, near Cren-

shaw Co. line, Covington Co.: UF 65313; Lightwood

Knot Creek, 1.6 mi. west of Opp, Covington Co.:

ANSP#; Little Patsaliga Creek: USNM 150473 [Holo-

type, Lampsilis australis, Simpson, 1900]; West Fork

Choctawhatchee River at Blue Spring State Park, Bar-

bour Co.: UAUC 134, UAUC 511, UAUC 512, UAUC

513, UAUC 514; Little Choctawhatchie Creek, near

Drian bridge, Houston Co.: UF 229532; Conecuh River

on CR 28 ~1 mi. east of Goshen, Pike Co.: UAUC 510;

Flat Creek at AL Hwy 153, near Flat Creek Church,

Geneva Co.: UAUC 547; Five Runs Creek, Conecuh N

F., Covington Co.: MMNS 8086.

FLORIDA

Shoal River at Hwy 85, 1 mi. south of I-10 Jct. in Crest-

view, Okaloosa Co.: UAUC 550, UAUC 551, UAUC 552,

UAUC 643, UAUC 644; Shoal Creek, ca. 1 mi. above

U.S. Highway 90, Okaloosa Co.: UF 261852; Limestone

Creek, Walton: MMNS 7746, MMNS 7747.

THE NAUTILUS 119(1):11-14, 2005

Page 11

Consideration of genetic relationships in management decisions

for the endangered Anthony’ riversnail, Leptoxis crassa

anthonyi (Redfield, 1854) (Gastropoda: Pleuroceridae)

Russell L. Minton

Museum of Natural History

University of Louisiana at Monroe

Monroe, LA 71209-0504 USA

[email protected]

Box 870345

Steven P. Savarese, Jr.

University of Alabama

Tuscaloosa, AL 35487 USA

ABSTRACT

Anthony’s riversnail, Leptoxis crassa anthony, is a federally en-

dangered pleurocerid restricted to three natural populations i in

the Tennessee River drainage. Recovery plans organized the

three populations into two management units, and called for

specific numbers of populations for downlisting or delisting.

Given that nothing was known about the genetic structure of

these populations and that individuals were being randomly

transplanted, we examined each population using COI mtDNA

sequences. All three populations possessed unique sequence

haplotypes, and the two units identified in the recovery plan

did not group the populations accurately in a phylogenetic con-

text. Potential management decisions in light of our findings

are discussed.

INTRODUCTION

Anthony’ riversnail, Leptoxis crassa anthonyi (Redfield,

1854), is a pleurocerid gastropod currently listed as en-

dangered by the United States Fish and Wildlife Service

(USFWS) under the Endangered Species Act of 1973 as

amended (USFWS, 1994). Adult shells of L. crassa an-

thonyi are large compared to sympatric pleurocerids,

globose to ovate, greenish to greenish-brown in color,

often with purple bands. The body whorl may be sculp-

tured with low, indistinct tubercles, giving the shell a

bumpy appearance. The aperture is ovate with a thin

outer lip, and the columellar lip is reflected so that it

covers a prominent umbilicus (Tryon, 1873). Historically,

L. crassa anthonyi had been documented from the Ten-

nessee River drainage in Alabama, Georgia and Tennes-

see, including the lower reaches of some of the larger

tributaries (Burch and Tottenham, 1980; Bogan and Par-

malee, 1983). Most populations of the species were ex-

tirpated when much of the Tennessee River and its trib-

utaries were impounded by the Tennessee Valley Au-

thority. Natural populations of L. crassa anthonyi persist

at three localities: a small, scattered population in the

main channel of the Tennessee River near the Alabama

and Tennessee state line; a large population in Lime-

stone Creek, Limestone County, Alabama; and a small

population limited to a single stretch of the Sequatchie

River, Marion County, Tennessee (Gamer, 1994; Jenkin-

son, 1994; USFWS, 1997; Figure Il),

Two morphologically similar snails have occurred

within the historic range of Leptoxis crassa anthonyji.

The species often occurs sympatrically with L. praerosa

(Say, 1821), and adults of L. praerosa are easily confused

with juveniles of L. crassa anthonyi (Dillon and Ahls-

tedt, 1997). The other taxon, L. crassa crassa (Halde-

man, 1841), was described as being similar to L. crassa

anthonyi but with larger, more prominent tubercles on

the body whorl, is presumed extinct (Turgeon et al.,

1998). Leptoxis crassa anthonyi shells are most easily

recognized as juveniles, given their saucer shape and the

presence of a heavy carina that disappears with age (Dil-

lon and Ahlstedt, 1997). The distinctness of L. crassa

anthonyi has been supported by both allozyme (Dillon

and Ahlstedt, 1997) and mitochondrial sequence data

(Holznagel and Lydeard, 2000).

In 1997, USFWS published their recovery plan

(USFWS, 1997) for L. crassa anthonyi. The plan iden-

tified two “populations” of L. crassa anthonyi: individ-

uals from the Sequatchie and Tennessee Rivers as one

population; and individuals from Limestone Creek as the

other. Recovery criteria for L. crassa anthonyi were to

protect the extant populations and to successfully re-es-

tablish other populations. If a total of four “populations”

could be established, the species could be downlisted to

threatened status; if six “populations” could be estab-

lished, the species would be delisted entirely. Though

the plan treated the three localities as two “populations,”

no effort had been made to determine if they were ge-

netically homogeneous (i.e., a single genetic population)

or genetically variable (i.e., two or three separate pop-

ulations). In order to reach the stated goals of the re-

covery plan, the number of genetically distinct popula-

tions within the species needed to be determined. A

working baseline of genetic information about each pop-

ulation was even more important due to reports of snails

Page 12

Figure I. J

ural populations of Leptoxis crassa anthonyi. L. = Limestone

pe Me

Map iat ee the locations of the three extant nat-

Creek, S = Sequatchie River; T = Tennessee River.

being translocated into existing populations and moved

to new localities in the Tonnies River drainage (J. Gar-

ner and D. Hubbs, pers. comms.). In this study, we used

mitochondrial DNA sequence data to conduct a com-

parative g genetic analysis of extant L. crassa anthon yi

populations to examine the degree of genetic differen-

tiation among the three populations. Knowledge of the

genetic suaneune would help direct management efforts

in determining which populations could and should

serve as sources for augmentations and reintroductions

if desired.

MATERIALS AND METHODS

Adult specimens of L. crassa anthonyi from the three

natural populations were collected under an endan-

gered/threatened species subpermit (SA99-13). The

maximum allowed number of specimens were collected,

twenty from Limestone Creek, and five each from the

Sequatchie and Tennessee Rivers (see Appendix 1). For

Figure 2. Strict consensus of two most parsimonious trees (TL =

THE NAUTILUS, Vol. 119, No. 1

comparative purposes, a reduced data set from previous

analyses (Minton and Lydeard, 2003) was used to place

L. crassa anthonyi in the proper systematic context. Fif-

teen individuals of L. crassa anthonyi from Limestone

Creek and five each from the other two populations

were included in the genetic study, along with four L.

praerosa, each from different river drainages, and one

each of the other taxa used in the previous study (Ap-

pendix 1).

Mitochondrial DNA sequences for a 1 kb portion of

the cytochrome oxidase subunit I gene (COI) were gen-

erated using published methods (Minton and Lydeard,

2003) for genomic isolation, PCR amplification, and ge-

netic analysis that followed. Sequences were aligned by

eye (Hall, 1999) and phylogenetic hypotheses g generated

by PAUP® 4.0b10 (Swofford, 2002) under maximum par-

simony with the following options: 50 replicates of heu-

ristic search with emda addition, uninformative char-

acters were ignored, branches with minimum zero

length collapsed, and minimal length trees kept. This

gene fragment showed significant — “phylogenetic signal

and no base composition bias or sequence saturation in

a more inclusive pleurocerid dataset (Minton and Ly-

deard, 2003). Internal branch stability was assessed by

jackknife (Farris et al., 1996) and Bremer support (Bre-

mer, 1994).

RESULTS

Aligned sequences resulted in a data matrix of 890 char-

acters, of which 222 were parsimony-informative. Each

population of Leptoxis crassa anthonyi possessed a

unique haplotype, and all individuals from a population

shared the same haplotype. Maximum parsimony anal-

ysis yielded two trees (Figure 2, strict consensus). All

specimens of L. crassa anthonyi constituted a monophy-

letic group. Uncorrected p-distances were 1.46% be-

tween the Limestone Creek and Sequatchie River spec-

Pleurocera canaliculatum

Pleurocera walkeri

Pleurocera prasinatum

Elimia sp. 2

Elimia sp. 1

Elimia hydei

Lithasia

lo fluvialis

Leptoxis praerosa

Leptoxis c. anthonyi T (n=5)

Leptoxis c. anthonyi L (n=15)

Leptoxis c. anthonyi S (n=5)

718, CI = 0.54) based on mitochondrial COI sequences.

Leptoxis crassa anthony? specimens identified as in Figure 1. Jackknife values >50% above branches, Bremer values below. Com-

position of Leptoxis and Lithasia clades are given in Table 1; both clades were monophyletic with jackknife support >50% and

Bremer support >1.

R. L. Minton and S. P. Savarese, Jr., 2005

imens, 2.02% between the Limestone Creek and Ten-

nessee River specimens, and 3.03% between the Se-

quatchie and Tennessee River specimens. The L. crassa

anthonyi clade was sister to a clade of L. praerosa spec-

imens. Most clades were well supported by jackknife and

Bremer values, as were the relationships within each

clade.

DISCUSSION

Our results further support the validity of Leptoxis cras-

sa anthonyi and its distinctiveness from L. praerosa, and

show each of the three populations of L. crassa anthonyji

to be genetically unique based on COI haplotypes. Un-

corrected pairwise genetic differences between the three

populations of L. crassa anthonyi were consistent with

published intraspecific differences seen in other pleu-

rocerids using COI sequences (Minton and Lydeard,

2003). The USFWS recovery plan calls for the presence

of no fewer than four viable populations of L. crassa

anthonyi before any change in listing status can occur.

For establishment of new populations, introductions of

each haplotype to new areas within their current river

system would be preferred in order to avoid mixing of

unique evolutionary entities and possible elimination of

the current genetic diversity.

The USFWS has designated Leptoxis crassa anthonyi

from the Sequatchie and Tennessee Rviers as a single

population. Based on that assumption, their manage-

ment efforts might target one of the populations as a

source for augmenting the other. If translocations are to

be used, our analysis suggests that the Limestone Creek

population would be a better option for augmenting the

Sequatchie River population due to lower genetic dif-

ference. Additionally, if a single population is to serve as

a source of new introductions and augmentations, recent

survey work (J. Garner, pers. comm.) indicates that

Limestone Creek would be preferred because of its

large population size. Regardless, controlled experimen-

tal populations should be established, either in the field

or laboratory, that could be monitored genetically and

for population growth and be compared with non-aug-

mented populations to determine whether fitness has

been enhanced or diminished from the introduction of

unique haplotypes from other populations. This measure

would be especially prudent given that haplotypes are

being randomly introduced through human activity. Any

recovery plan that involves translocation of L. crassa an-

thonyi should use juveniles, as they are more easily iden-

tified by their strong keel. This will help ensure the

movement of L. crassa anthonyi and potentially prevent

the introduction of adult, non-endemic L. praerosa.

ACKNOWLEDGMENTS

This project was completed in the lab of C. Lydeard at

the University of Alabama. We thank S. Ahlstedt and J.

Garner for specimens, and two anonymous reviewers for

Page 13

their comments. Funding was provided by a USFWS

grant to C. Lydeard (1448-40181-97-G-033).

LITERATURE CITED

Bogan, A. E. and P. W. Parmalee. 1983. Tennessee’s Rare Wild-

life Volume II: The Mollusks. Tennessee Wildlife Re-

sources Agency, Nashville, 123 pp.

Bremer, K. 1994. Branch support and tree stability. Cladistics

10: 295-304.

Burch, J. B. and J. L. Tottenham. 1980. North American fresh-

water snails. Species list, ranges, and illustrations. Walk-

erana 1: 81-215.

Dillon, R. T. and S. A. Ahlstedt. 1997. Verification of the spe-

cific status of the endangered Anthony’s riversnail, Athear-

nia anthonyi, using allozyme electrophoresis. Nautilus 10:

97-101.

Farris, J. S., V. A. Albert, M. Kallersjo, D. Lipscomb and A.

G. Kluge. 1996. Parsimony jackknifing outperforms neigh-

bor-joining. Cladistics 12: 99-124.

Gamer, J. 1994. Survey of mollusks: Tennessee River mile

412.1. Aquatic Resources Center, Franklin, 2 pp.

Jenkinson, J. 1994. Freshwater mollusk survey at CSX railroad

bridge, near Bridgeport, Alabama, Tennessee River Mile

414.5. Tennessee Valley Authority, Chattanooga, 15 pp.

Haldeman, S. S. 1841. A monograph of the Limnaides and

other freshwater univalve snails of North America. No. 3,

[Limnea]. J. Dobson, Philadelphia, 16 pp.

Hall, T. A. 1999. BioEdit: a user-friendly biological sequence

alignment editor and analysis program for Windows 95/

98/NT. Nucleic Acids Symposium Series 41: 95-98.

Holznagel, W. E. and C. Lydeard. 2000. A molecular phylogeny

of North American Pleuroceridae (Gastropoda: Cerithioi-

dea) based on mitochondrial 16S rDNA sequences. Jour-

nal of Molluscan Studies 66: 233-257.

Minton, R. L. and C. Lydeard. 2003. Phylogeny, taxonomy,

genetics, and global heritage ranks of an imperiled, fresh-

water snail genus Lithasia (Pleuroceridae). Molecular

Ecology 12: 75-87.

Redfield, J. H. 1854. Descriptions of new species of shells.

Annual Report of the Lyceum of Natural History of New

York 6: 130-132.

Say, T. 1821. Descriptions of the univalve shells of the United

States. Journal of the Academy of Natural Sciences of

Philadelphia 2: 149-179.

Swofford, D. 2002. PAUP®: Phylogenetics Analysis Using Par-

simony, version 4.0b10. Sinauer Associates, Sunderland,

Massachusetts.

Tryon, G. W. 1873. Land and freshwater shells of North Amer-

ica. Part IV. Strepomatidae. Smithsonian Miscellaneous

Collections 253: i-iv, 1-435.

Turgeon, D. D., J. F. Quinn, A. E. Bogan, E. V. Coan, F. G.

Hochberg, W. G. Lyons, P. M. Mikkelsen, R. J. Neves, C.

F. E. Roper, G. Rosenberg, B. Roth, A. Schletema, F. G.

Thompson, M. Vecchione and G. D. Williams. 1998.

Common and scientific names of aquatic invertebrates

from the United States and Canada: Mollusks (second edi-

tion). American Fisheries Society Special Publication 26,

Bethesda, 526 pp.

United States Fish and Wildlife Service. 1994. Endangered and

threatened wildlife and plants; determination of endan-

gered status for the royal snail and Anthony’s riversnail.

Federal Register 59: 17994-17998.

United States Fish and Wildlife Service. 1997. Recovery plan

for Anthony’s riversnail. Atlanta, Georgia. 21 pp.

Page 14

Appendix 1. Systematic list of taxa and specimens used in this study. Complete locality information is available from the authors.

UAG = University of Alabama gastropod collection.

Taxon

Locality

Genus Elimia

E. hydei

Elimia sp. 1

Elimia sp. 2

Genus Io

Io fluvialis

Genus Leptoxis

L. crassa anthonyji

L. praerosa

Genus Lithasia

L. armigera

L. geniculata fuliginosa

L. lima

L. verrucosa

Genus Pleurocera

P. canaliculatum

P. prasinatum

P. walkeri

UAG voucher

Black Warrior River, Tuscaloosa Co., AL

Green River, Hart Co., KY

Collins River, Warren Co., TN

Clinch River, Hancock Co., TN

Limestone Creek, Limestone Co., AL

Sequatchie River, Marion Co., TN

Tennessee River, Jackson Co., AL

Harpeth River, Davidson Co., TN

Shoal Creek, Lawrence Co., AL

Sequatchie River, Marion Co., TN

Tennessee River, Jackson Co., AL

Harpeth River, Cheatham Co., TN

Red River, Robertson Co., TN

Duck River, Maury Co., TN

Buffolo River, Humphreys Co., TN

Bear Creek, Colbert Co., AL

French Broad River, Knox Co., TN

White River, Woodruff Co., AR

Duck River, Maury Co., TN

Yellowleaf Creek, Shelby Co., AL

Shoal Creek, Lauderdale Co., AL

UAG 584

UAG 574

UAG 407

UAG 585

UAG 581

UAG 582

UAG 583

UAG 404

UAG 560

UAG 588

UAG 589

UAG 555

UAG 398

UAG 403

UAG 406

UAG 570

UAG 576

UAG 577

UAG 590

UAG 591

UAG 592

THE NAUTILUS, Vol. 119, No. 1

Genbank accession

AF435775

AF435759

AF435761

AF43577|

AF435772

AF435773

AF435774

AF435779

AF435780

AF435781

AF435782

AF435739

AF435754

AF435749

AF435747

AF435757

AF435767

AF435771

AF435783

AF435784

AF435785

THE NAUTILUS 119(1):15-26, 2005

Page 15

Fallen into oblivion—the systematic affinities of the enigmatic

Sulcospira Troschel, 1858 (Cerithioidea: Pachychilidae), a genus

of viviparous freshwater gastropods from Java

Frank K6éhler!

Matthias Glaubrecht®

Museum of Natural History

Humboldt University

Institute of Systematic Zoology

Department of Malacozoology

Invalidenstrale 43, D-10115 Berlin

GERMANY

1 frank. [email protected]

> [email protected]

ABSTRACT

Sulcospira Troschel, 1858, is not only the taxonomically oldest

but also one of the most poorly known genera of Southeast

Asian Pachychilidae. It serves as an instructive case study as to

how the puzzling systematics of freshwater Cerithioidea has

hampered a deeper understanding of their phylogeny and evo-

lution. The genus has been established for the Javan freshwater

gastropod Melania sulcospira Mousson, 1849, on the grounds

of its round, multispiral operculum and an elongated main cusp

in the central radula teeth. Although of great systematic sig-

nificance, this taxon has been widely ignored by subsequent

authors. We here recapitulate the taxonomic history of the ge-

nus and describe and evaluate the morphological properties to;

the type species S. sulcospira on basis of the limited existing

material, in order to facilitate a better understanding of pachy-

chilid systematics. In addition, in an attempt to clarify its sys-

tematic affinity, we compare the properties of another allegedly

related species from Java, S. martini (Schepmann, 1898). We

show that these two viviparous species exhibit different pro-

toconch morphologies, which are indicative of reproductive

strategies distinct from other pachychilids. Finally, we outline

preliminary suggestions as to the systematics of Sulcospira

within the family Pachychilidae.

MATERIALS AND METHODS

MATERIALS

This study is based on the examination of material from

various museum collections worldwide (see repositories).

All of these samples comprise dry shells only, which as

a rule were empty. Only few shells contained fragmen-

tary soft parts; some of them were re-hydrated for ex-

aminations. However, these bodies generally did not fa-

cilitate morphological examinations except for the ex-

traction of small radula fragments. We have not tried to

extract DNA from dried tissues because earlier attempts

with comparable material of Brotia failed. In order to

acquire fresh material, collecting trips have been under-

taken in 2000 and 2002. We have searched rivers and

creeks in different sectors of there course (i.e. upstream,

midstream, downstream) for a period of altogether 7

days in West Java (along the roads between Jakarta al

Serang, Bogor and Sukabumi, Sukabumi and Pelabuhan

Ratu, Bogor and Cipanas, Bogor and Cianjur, in the Bo-

tanical Garden Bogor) and for 3 days in East Java (be-

tween Taksimalaya, Cipatujah, and Pangadaran). During

these trips we were not able to find any material of Sul-

cospira, though.

Because freshwater biotopes on Java are facing dra-

matic devastation by a multitude of causes related to the

dense population on this island such as pollution, flow

regulation, drainage, impoundment and a general deg-

radation of collecting areas by agriculture, industry and

settlements (own observations: see also Dudgeon, 2000,

for SE Asia in general), we believe that S. sulcospira has

become extinct in vast areas on Java. It remains unclear

whether and at which localities populations of this spe-

cies still exist.

REPOSITORY INSTITUTIONS

Voucher material is housed with the following museums:

Natural History Museum, London (BMNH), Museum

of Comparative Zoology, Cambridge, Mass. (MCZ), Mu-

séum d Histoire Naturelle, Genéve (MHNG), Muséum

National d Histoire Naturelle, Paris (MNHN), Natural

History Museum Naturalis, Leiden (RMNH), Sencken-

bergmuseum, Frankfurt/Main (SMF), Zodlogisch Mu-

seum, Amsterdam (ZMA), Museum fiir Naturkunde,

Berlin (ZMB).

We were not able to locate material in the following

museum collections: Academy of Natural Sciences, Phil-

Page 16

adelphia (ANSP), United States National Museum,

Washington (UNSM), Zoologisches Institut und Muse-

um, Universitit Hamburg (ZMH), Zoologische Staats-

sammlung, Miinchen (ZSM).

MORPHOLOGICAL EXAMINATIONS

Dimensions of all shells were measured to 0.1 mm pre-

cision. The shell height (H) is the maximum dimension

parallel to the axis of coiling, breadth (B) the maximum

dimension perpendicular to H, including the aperture.

The length of the aperture (LA) is the greatest length

from the junction of the outer lip with the penultimate

whorl to the anterior lip, the width (WA) the greatest

length perpendicular to LA. The height of the body

whorl (BW) is the distance from the base of the shell to

the upper suture of the first whorl exactly above the

junction of the outer lip with the penultimate whorl.

Morphometrical parameters used in the analyses, beside

the shell dimensions, were: H/B, H/LA, H/BW, H/LA

and B/BW. These shell parameters were statistically an-

alyzed by performing t-tests, one-way ANOVA, and a

discriminate analysis.

Protoconchs removed from dried adults were cleaned

by soaking in 10% KOH solution, flushed in distilled

water, and sonicated to remove residual contaminations

prior to scanning electron microscopy. Radulae were tak-

en from dried shells or from historic preparations. Rad-

ulae from dried shells were enzymatically cleaned as de-

scribed by Holznagel (1998); an old radula embedded in

Canada balsam was cleaned with xylene followed by son-

ication. Radulae and juvenile shells were mounted on

aluminum specimen stubs using adhesive carbon tabs or

double-sided tape, respectively, and coated with gold-

palladium for 120 s at 20 mA for examination under a

scanning electron microscope (LEO 1450 VP) at 10 keV.

NOMENCLATORIAL REMARKS

Some species-group names introduced by Troschel

(1857-1858) are open to discussion. Bouchet (pers.

comm.) argued that the usage of the names Bithyniae,

Lithoglyphi, Hydrobiae, Ancyloti, Thiarae, and Pachy-

chili by Troschel (op. cit.) contrasts with the rest of his

work (Troschel, 1856-1863), in which he stated the

ranks of the categories he used and formed names with

endings -idea, -ina, or -acea. Because Troschel stated

explicitly that he refrained from allocating these group-

ings at family rank given the somewhat ambiguous mor-

phological data he was faced with, it was suggested that

one should ignore these names (Bouchet, pers. comm.).

However, some of these names, such as Bithyniidae,

Thiaridae, or Hydrobiidae, have been usually published

with Troschel (1857-1858) as author. Unless otherwise

stipulated, we prefer to refer to Troschel (1857-1858) as

author of these names not only because we regard them

as available and valid but also in order to maintain sta-

bility in their usage; for a statement to the contrary see

Bouchet and Rocroi (submitted).

THE NAUTILUS, Vol. 119, No. 1

i a

Figure 1. Original drawing of the radula of “Sulcospira typ-

ica (Melania sulcospira Mousson)” by Troschel (1858: pl. 9, fig.

6). In the upper part of the figure a row of teeth is shown

comprising a rachidian flanked on each side by a lateral tooth

and an inner and outer marginal tooth; a magnified represen-

tation of the rachidian is shown below. Characteristics that are

typical for Pachychilidae are, e.g., the enlarged main cusp of

the rachidian and lateral teeth, respectively, which is flanked

by three (or two) smaller cusps on each side that taper in size;

the presence and shape of the glabella (or ramp); and marginal

teeth possessing two cusps.

Nomenclatorial aspects raised in this paper refer to

the stipulations of the 4" edition of the International

Code of Zoological Nomenclature (“ICZN”) issued by

the International Commission of Zoological Nomencla-

ture (1999).

RESULTS

Sulcospira Troschel, 1858

Sulcospira Troschel, 1858: 117-118; Brot, 1874: 56; Thiele,

1929: 190; Morrison, 1954: 381.

Diagnosis: Sulcospira possesses a rather conical shell

sculptured by spiral lirae; axial sculptural elements are

lacking. Protoconchs possess a smooth sculpture with a

fine granular texture or faint growth lines.

Type Species:

monotypy.

Nomenclature and Systematics: The genus Sulcos-

pira was described by Troschel (1858) for the Javan spe-

cies M. sulcospira exhibiting certain characteristics that

were held to be peculiar of this species, namely a round,

multispiral operculum with four regular whorls and the

radula with an enlarged main cusp of the rachidian (Fig-

ure 1, 17-18). Troschel (1858: 114) based his description

on material received from August Brot in Geneve, i.e.,

Melania sulcospira Mousson, 1849, by

F. Kohler and M. Glaubrecht, 2005

Figure 2. Shell dimensions. B: breadth; BW: weight of the

body whorl; H: height; LA: length of the aperture; WA: width

of the aperture.

on material that is likely housed at MHNG today (see

Material Examined).

Sulcospira represents the oldest available generic

name established for representatives of the Southeast

Asian Pachychilidae and is here considered valid. How-

ever, the diagnosis of Troschel (1858) is not sufficient to

characterize the taxon unambiguously. Neither a round

and multispiral operculum with four whorls nor the pos-

session of a pronounced main cusp of the rachidian are

considered as diagnostic features of Sulcospira alone

(see discussion). Nonetheless, these features character-

ize Sulcospira as a member of the Pachychilidae (Glau-

brecht, 1996, 1999: KGhler and Glaubrecht, 2001, 2002,

2003).

In more recent literature Sulcospira has been widely

ignored. Thiele (1929) suggested subdividing Sulcospira

into two subgenera, Sulcospira and Tylomelania F. and

P. Sarasin, 1898. According to Thiele, Sulcospira would

include Paracrostoma, Acrostoma Brot, 1870, and Bro-

tella Rovereto, 1899, as junior synonyms, consequently

comprising two species: S. sulcospira from Java and S.

Page 17

huegeli (Philippi, 1843) from South India. Tylomelania,

however, was considered to encompass a small number

of species restricted to Sulawesi.

This concept of Thiele (1929) led Subba Rao (1989:

107) to wrongly assume that M. huegeli would be the

type species of Sulcospira, which it is not (see Troschel’s

original designation).

Later authors had different taxonomic views. Morri-

son (1954) followed Abbott (1948) but not Thiele (1929)

and treated Acrostoma, Brotella, and Paracrostoma as

synonyms of Brotia, while considering Tylomelania as a

genus on its own. He also claimed that our understand-

ing of Sulcospira is not satisfactory. Based on some su-

perficial similarities with Brotia, but also with Tylome-

lania and Balanocochilis (a thiarid), Morrison (1954) pro-

posed the allocation of Sulcospira “tentatively to the Me-

lanoides complex’. Since Melanoides is a thiarid, this

allocation is rejected here.

All these classification schemes were suggested in ab-

sence of phylogenetic analyses of morphological char-

acters. In addition to the type species, other taxa have

been assigned to the genus by previous authors, al-

though this has not hecn done consistently (Table 1).

The various opinions led also to different assumptions

on the species circumscription and diversity of this tax-

on. For example, Brot (1874) subsumed a number of

taxa under Sulcospira, of which we currently only con-

sider two to be actually pachychilids: Melania spadicea

Reeve, 1860, and M. hainanensis Brot, 1872. Yen (1939)

added two more taxa, M. ebenina Brot, 1883, and M.

biconica Brot, 1886. Boettger (1890), Oostingh (1932),

and Adam and Leloup (1838) treated M. testudinaria

von dem Busch, 1842, as member of Sulcospira but did

not mention the former taxa. In contrast, other authors

assigned those taxa to Brotia instead (Rensch, 1934;

Benthem-Jutting, 1956, 1959; Knipper, 1958; Dudgeon,

1982, 1989: Kohler and Glaubrecht, 2001, 2002). Even-

tually, Benthem-Jutting (1956) assumed that S. sulcos-

pira is the only representative of the genus.

In order to clarify the puzzling taxonomy and system-

atics, a revision of Sulcospira with an evaluation of its

anatomical characters is needed. A sound classification

has to be based on autapomorphic features, which is

lacking to date. Unfortunately, a comprehensive descrip-

tion of the morphology of Sulcospira suffers from the

Table 1. Comparison of previous views on the systematics and circumscription of Sulcospira.

Brot (1874)

Taxonomy at generic Melania (Sulcospira)

Sulcospira (Sulcospira),

Authors

Morrison (1954):

Benthem-Jutting K6hler and Glaubrecht

Thiele (1929) (1956) (2002)

Sulcospira Sulcospira

level Sulcospira (Tylomelania)

Included taxa M. sulcospira, M. spadicea,

M. hainanensis, and other

non-pachychilid taxa

S. sulcospira, S. huegeli,

T. neritiformis, T. carbo,

T. porcellanica

S. sulcospira S. sulcospira, S. spadicea

Page 18

THE NAUTILUS, Vol. 119, No. 1

Table 2. Shell parameters [mm] of S. sulcospira and S. martini. Abbreviations: B: breadth; BW: weight of the body whorl; H:

height; LA: length of the aperture; m: median; N: number of whorls; No: number of shells; sd: standard deviation; WA: width of

the aperture.

Lot No H B LA WA BW N

S. sulcospira (total) m 26 19.3 10.1 9.5 4.9 14.3 4.7

sd 2.5 1.3 1.0 0.6 1.9 0.5

Holotype M. sulcospira 1 23.2 12.0 9.7 5.0 15.2 4.0

S. sulcospira (ZMA) mm = SL 19.5 10.2 9.5 4.9 14.4 4.6

sd 2.2 1.2 0.9 0.5 0.6 0.6

S. sulcospira (MNHN) m 4 18.7 9.9 9.3 49 14.1 48

sd 3.6 1.8 1.5 0.9 2.9 0.4

S. martini (total) m 058 27.6 12.0 11.0 5.9 17.7 6.0

sd 4.5 1.5 1.3 0.8 2.1 1.0

Syntypes M. spadicea (BMNH 19990497) m 3 25.5 11.6 10.2 5.2 16.1 6.0

sd 0.7 0.2 0.9 0.6 0.9 0.5

Syntypes M. junghuhni (RMNH, ZMA) m 292 29.9 12.4 11.6 6.0 18.4 6.3

sd 4.6 1.8 1.4 0.9 2.4 0.7

Syntypes var. flammulata, var. fasciata mm 1G 28.7 12.3 11.7 6.1 179 6.6

(RMNH) sd 2.9 1.2 Ll 0.7 1.7 0.6

S. martini (ZMB 4.074) m 17 23.8 1 11.0 5.7 16.9 49

sd 2.9 1.3 1.0 0.5 Wy 0.7

lack of well-preserved material of the type species. We

compile below the anatomical data based on the material

of S. sulcospira available.

Sulcospira sulcospira (Mousson, 1849)

Melania sulcospira Mousson, 1849a [1848]: 269; Mousson,

1849b; 68, pl. 9, fig. 3; Martens, 1897: 245 (partim); Les-

chke, 1914: 251.

Sulcospira typica Troschel, 1858: 117, 118, pl. 9, fig. 6 [intro-

duced as replacement name for M. sulcospira Mousson,

1849].

Pachycheilus sulcospira [sic]—H. and A. Adams, 1858: 299.

Melania (Sulcospira) sulcospira.—Brot, 1870: 277; Brot, 1874:

56-57, pl. 6, fig. 11; Boettger, 1890: 245.

Sulcospira sulcospira.—Morrison, 1954: 381; Kéhler and Glau-

brecht, 2002: 149, fig. 3 L.

Diagnosis: Shell relatively small (Table 2), conical

with spiral lirae, and a subsutural depression; aperture

elongately ovate, abapically flared. Radula with squarish

rachidian exhibiting a straight upper and lower rim, the

cutting edge of all teeth with one very pronounced main

cusp of triangular shape.

Description: Shell (Figures 3-5): small, ovate to

conical, solid, spire with eroded apex and up to six flat-

tened whorls, separated by a narrow suture; sculpture

consisting of fine, regular spiral lirae that are most prom-

inent at the base and may almost lack on upper whorls,

and faint growth lines; whorls with a subsutural depres-

sion; color from yellowish to olive or dark brown. Ap-

erture elongately ovate, abapically flared, peristome

sharp; columella slightly bent and thickened.

Protoconch (Figures 19-20): Relatively large, com-

pared to the adult; with height of about 1.2 mm com-

prising one and a half whorl; generally smooth. Apical

whorl inflated, dome-shaped, with a granular surface

sculpture, transition in sculpture visible on first whorl

from granular to faint growth lines. One sample of four

specimens housed at MNHN (ex coll. Staat) has a label

stating that “about 140 embryos were obtained from the

large specimen”. However, the fate of these protoconchs

is unknown.

Operculum: With four whorls regularly increasing in

diameter and a sub-central nucleus.

Radula (Figures 16-17): Rachidian tooth squarish with

a straight upper and lower rim, cutting edge with one

heavily enlarged main cusp of triangular shape, flanked

by two, much smaller accessory cusps on each side; gla-

bella well developed, rather rectangular with a rounded

basal margin not exceeding the lowes rim of the rachi-

dian tooth, lateral margins straight and not well defined.

Lateral teeth with one heavily enlarged main cusp, and

considerably smaller accessory cusps, one at the outer

side, two or three at the inner side. Inner and outer

marginal teeth with two cusps, the outer one being very

large, triangular in shape, and the inner one being point-

ed and small. Inner marginal teeth broader than outer

marginals. Outer lateral flange inconspicuous.

Anatomy: Unknown.

Type Material: Indonesia: Holotype ZMZ 522306, In-

donesia: Java, leg. Zollinger (Figure 3) [Mousson, 1849b

refers to “das einzige Exemplar dieser Art ...” = the

only specimen of this species . . .].

Type Locality: “Java”, Indonesia.

Other Material Examined: Indonesia: Java (MHNG;

MNHN: MNHN, ex coll. Staat; ZMA (2 lots); ZMB

200.101) (if not stated otherwise, a single lot from each

collection, mostly without reference number, was ex-

amined): museums without material: ANSP, BMNH,

MCZ, RMNH, SMF, USNM, ZMH, ZSM.

F. Kéhler and M. Glaubrecht, 2005

Page 19

Figures 3-16. Shell morphology of Sulcospira sulcospira (3-5) and S. martini (6-16) (apertural and abapertural, respectively).

3. S. sulcospira. Holotype (ZMZ 522306). 4-5. S. martini. Two shells from Java (ZMA). 6. Lectotype of M. spadicea (BMNH

19990497/A). 7-8. Two paralectotypes of M. spadicea (BMNH 19990497/B). 9. Lectotype of M. junghuhni (RMNH 71326). 10-

11. Two syntypes of M. junghuhni var. flammulata (RMNH 71327).

12-13. Two syntypes of M. junghuhni var. fasciata (RMNH

71328). 14-16. Three shells from Java, Malangbong (ZMB 4.074). Scale bar = 10 mm.

Page 20 THE NAUTILUS, Vol. 119, No. 1

Figures 17-24. SEM images of the radula and juvenile shells removed from dry shells of S. sulcospira (17-20) and S. martini

(21-24). S. sulcospira: 17. Radula (ZMA). 18. Radula (ZMB 200.101). 19. Juvenile shell, apertural view (ZMA). S. martini: 20.

Juvenile shell, apical view (ZMA). 21. Juvenile shell, apertural view (ZMB 4.074). 22-23. Apical view. 24. Detail of the apical

portion. Scale bars = 100 pm.

F. Kéhler and M. Glaubrecht, 2005

oo os

m 12 ange,

528

4 a See OS. sulcospira

12} ®S. sulcospira holotype

a @S. spadicea types

OS. martini types

XS. martini (ZMB 4074)

10 15 20 25 30 35 40 45

Figure 25. Comparison of the shells of S. sulcospira and S.

martini based on shell parameters height (H) and breadth (B).

Distribution: Indonesia: Java. More precise localities

were given by Boettger (1890: Bogor Botanical Garden)

and Martens (1997: Jakarta, Malangbong, Cipanas).

However, the oe of Boettger could not be traced

(SMF? ), and Martens misidentified material of Melan-

oides tuberculata (Cipanas) and S. martini (Malangbong;

ZMB 4.076); no voucher material was found from Ja-

karta.

Nomenclature and Systematics: Melania sulcospira

is the type species of the genus Sulcospira by original

designation. When describing the new genus, Troschel

(1858), mentioned M. sulcospira as the “typical species”,

and introduced the name S. typica for the same taxon

as a new, unnecessary substitute name meant to replace

an older available name (nomen novum). Thus, S. typica

is a junior synonym of S. sulcospira (ICZN Art. 72.7.).

Sulcospira typica is invalidated as potential type name

(ICZN Art. 68.4.) making M. sulcospira the type by ab-

solute autonomy. Meriane (1897) assumed that M. spad-

icea is a synonym of this species, which was rejected by

Leschke (1914) and Benthem-Jutting (1956: 373), stat-

ing that Martens (1897) had misidentified specimens of

Melanoides tuberculata for M. spadicea.

Remarks: Compared to Sulcospira martini, S. sulcos-

pira is more conical in shape; the former lacks a con-

spicuous subsutural depression. Most conspicuously,

both species differ in their protoconch morphology.

Shells of B. testudinaria are more elongated in shape,

attain a larger size (between 25 to 40 mm in shell

height), lack a subsutural depression, and exhibit a dif-

ferent radular morphology: e.g., rachidian with inflated

and rounded upper corners, a smaller main cusp, mar-

ginal teeth possess two equally shaped cusps.

Given the scarcity of material and imprecise earlier

locality data for this material, we tried to restrict the type

locality from historical accounts. The Swiss malacologist

Albert Mousson (1805-1890) based his descriptions on

material collected by the Swiss botanist Heinrich Zollin-

ger (1818-1859), who traveled in Indonesia between

1842 and 1848. However, as Zollinger collected not only

H/B

N= 25 59

S. sulcospira S. martini

Figure 26. Comparison of S. sulcospira and S. martini based

on shell parameters ratio H/B. Box plot diagram showing the

median, the 25%- and 75%-percentile and largest non-ex-

tremes (less than 1.5 times of box height).

in West Java, but later also in East Java, and on some

other islands (Wanner, 1984), it turned out that a re-

striction of the type locality is not possible and “Java”

remains as the only known reference.

Sulcospira(?) martini (Schepmann, 1898)

Melania spadicea Reeve, 1860: pl. 19, species 132 (not Melania

spadicea Philippi, 1849); Brot, 1870: 277.

Melania (Sulcospira) spadicea.—Brot, 1874: 57-58, pl. 6, fig.

12.

Brotia spadicea.—Benthem-Jutting, 1956: 372-373, fig. 75.

Sulcospira spadicea.—Kohler and Glaubrecht, 2002: 148, fig.

3G.

Melania junghuhni Schepman, 1896: 135-136, pl. 2, fig. 1.

(“Java’; lectotype and 41 paralectotypes RMNH 71326; 24

paralectotypes ZMA; two paralectotypes SMF 292406; var.

flammulata: 16 syntypes RMNH 71327; 3 syntypes MCZ

96926, var. fasciata: 18 syntypes RMNH 71328; 24 syn-

types ZMA; 8 syntypes MCZ 96898) (not M. junghuhni

Martin, 1879); Leschke, 1914: 251; Benthem-Jutting,

1929: 84.

Melania martini Schepmann, 1898: 84.

Brotia testudinaria —K6ohler and Glaubrecht, 2001: 301-304

(partim); Kéhler and Glaubrecht, 2002: 140, 141, 150

(partim).

Diagnosis: Conical shell with convex to flattened

whorls sculptured with fine spiral ridges. Protoconch

with about 2.5 regular whorls; apical whorl not inflated,

no transition in shell structure is visible in the first two

whorls. Axial ribs may be present in the juvenile shell

from the second whorl on. Most conspicuously distin-

guished from all other pachychilids by its protoconch

morphology (Figures 27-34); distinguishable from S.

sulcospira by its more elongated and larger shell (Figure

26); though adult shell not distinguishable from B. tes-

tudinaria.

Page 22

THE NAUTILUS, Vol. 119, No. 1

Figures 27-34. Comparison of protoconchs of different southeastern Asian Pachychilidae (apertural and apical view, respectively).

27-28. Brotia costula. 29-30. Brotia hainanensis. 31-32. Tylomelania patriarchalis (with kind permission of Thomas von Rintelen).

33-34. Pseudopotamis semoni. Scale bars = 100 jm.

Description: Shell (Figures 6-16): Small to medium

sized (Table 2), ovate to conical, spire with eroded apex

and eight to ten flattened whorls; sculpture consisting of

fine, closely spaced regular spiral lirae, may lack almost

completely, and faint growth lines; color yellowish brown

to olive, brown spiral band or patches may be present.

Aperture elongated ovate, produced below, peristome

sharp.

Protoconch (Figures 21-24): Height of about 1.2 mm

comprising 2% whorls; apical whorl not inflated, corre-

sponding to the regular diameter of the subsequent

whorls; first two whorls smooth, only faint growth lines

visible, without transition in sculpture, from the second

whorl on smooth axial ribs may be present.

Operculum: Consisting of three whorls and a sub-basal

nucleus Brot (1874).

Anatomy and Radula: Unknown.

Type Material: Indonesia, Java: Lectotype and 41

paralectotypes of M. junghuhni, RMNH 71326, leg. Jun-

ghuhn (Figure 9), designated by Kéhler and Glaubrecht

(2002); 24 paralectotypes, ZMA; two paralectotypes,

SMF 292406; 16 syntypes of M. junghuhni var. flam-

mulata, RMNH 71327; three syntypes, MCZ 96926, 18

syntypes of M. junghuhni var. fasciata, RMNH 71328;

eight syntypes, MCZ 96898. Without locality: Lectotype

of M. spadicea, BMNH 19990497/A, Cuming collection,

designated by Kohler and Glaubrecht (2002); two para-

lectotypes, BMNH 19990497/2.

Type Locality: “Java”, Indonesia.

Other Material Examined: Without locality (ZMA;

MHNG); Indonesia: Java (MHNG), Malangbon (ZMB

4.074) (a single lot from each collection, mostly without

reference number). No material was found in the fol-

lowing museums: ANSP, MNHN, SMF, USNM, ZMH,

ZSM.

Nomenclature and Systematics: Melania spadicea

Reeve, 1860, is a primary homonym of M. spadicea Phi-

lippi, 1849 (ICZN Art. 53.3) and, as such, is permanently

invalid (ICZN Art. 57.2). The next available names are

M. junghuhni Schepmann, 1896, M. junghuhni var. flam-

F. Kohler and M. Glaubrecht, 2005

mulata Schepmann, 1896, and M. junghuhni var. fasciata

Schepmann, 1896. However, the first is a primary hom-

onym of Melania junghuhni Martin, 1879, a fossil spe-

cies from Java, as stated by Schepmann (1898). The oth-

er two are junior primary synonyms of Melania flam-

mulata von dem Busch in Philippi, 1843, and Melania

fasciata Menke, 1828, respectively.

Schepmann (1898) suggested Melania martini as a re-

placement name for M. junghuhni, which is the valid

name for this taxon. The two color morphs described by

Schepmann (1896), fasciata and flammulata, are not

considered here to represent extant evolutionary entities

and therefore are treated as junior synonyms of M. mar-

tint.

The classification of this species by former authors is

inconsistent. Benthem-Jutting (1956) considered it to be

a member of Brotia; but unaware of the fact that M.

spadicea Reeve, 1860, is not valid she assumed that this

name had priority over M. martini. KGhler and Glau-

brecht (2002) assumed that M. martini and M. spadicea

are distinct and treated the former as a synonym of B.

testudinaria and the latter as a species closely related to

S. sulcospira as was suggested earlier by Brot (1874).

Distribution: Java, as the only known locality. Ma-

langbon in Central Java, east of Bandung, is the only

eau exact locality (ZMB 4.074, catalogued in 1859).

Analyses of Shell Morphometry: Sulcospira martini

can be distinguished from S. sulcospira by its higher

shell and more slender shape (see analyses of shall pa-

rameters below). However, to differentiate between

shells of “M. sulcospira” and “M. martini” is no easy task;

contradictory statements on their taxonomy abound in

earlier accounts (Brot, 1874; Benthem-Jutting, 1956:

KGhler and Glaubrecht, 2002). In fact, shells exhibit a

very similar shape, sculpture, and coloration. However,

the two taxa can be distinguished by statistical analyses

of shell morphometry. We Teed one-way ANOVA and t-

test for two independent groups of variables to discrim-

mee specimens that were assigned beforehand either to

S. sulcospira or to S. martini according to their shell

morphology. The t-test showed that both taxa vary sig-

nificantly by the following parameters (P<5%): H, N,

H/B, H/LA, and H/BW; the one-way-ANOVA yielded

corresponding results.

The shells of the only lot from Java with precise lo-

cality data (ZMB 4.074) is identified here as S. martini

given its elongated shell (while the original label states

“M. sulcospira” instead). The statistical test has been

employed to explore whether these shells can signifi-

cantly be discriminated either from shells of S. sulcos-

pira or S. martini in regard to shell morphometry. Com-

parison of shells of the lot ZMB 4.074 with shells of S.

sulcospira (MNHN, ZMA, ZMB 200.101, ZMZ 522306)

by t-test reveals that both groups differ significantly in

the parameters H/LA, H/BW, and with a weak support

(P = 0.51) for H/B, whereas no significant differences

were found when comparing the lot ZMB 4.074 with

Page 23

Table 3. Results of the discriminate analysis of shell param-

etensy

Predicted group

membership S. sulcospira S. martini

S. sulcospira 25 (100.0%) 0 (0.0%)

S. martini 1 (1.7%) 59 (98.3%)

other shells of S. martini (BMNH 19990497, RMNH

71326-8).

A graphic comparison of the two taxa by means of

selected shell parameters is shown in Figures 25—26.

Shells assigned to each of the two taxa according to their

morphology were found to be correctly classified by a

discriminate analysis of morphometric data with good

statistical support (Table 3). It is concluded that shells

of S. sulcospira are smaller and more conical in shape

than shells of S. martini (Table 1, Figure 26).

DISCUSSION

I. EVALUATION OF MORPHOLOGICAL CHARACTERS OF

SULCOSPIRA

The operculum and radula of S. sulcospira led Troschel

(1858) to describe a new genus for this species. How-

ever, among the Pachychilidae the operculum is known

to be relatively conservative in its general organization

(that is, to be multispiral, rounded or ovate) “but quite

variable in relation to their number of whorls and in-

crease in diameter, even within a single genus (Kéhler

and Glaubrecht, 2001, for Brotia; Kohler and Glau-

brecht, 2003, for Jagora; Glaubrecht and Rintelen, 2003,

for Pseudopotamis; Rintelen, 2003, for Tylomelania;

Kohler, 2003). Consequently, an operculum possessing

four regular whorls might be typical for S. sulcospira,

but only at the species level. The possession of a round

to oval, multispiral operculum led Sarasin and Sarasin

(1898) to group several taxa within the so-called “pa-

laeomelanians”, as contrasted with the so-called “neo-

melanians”, which exhibit a paucispiral operculum. In

fact, this grouping coincides well with the modern con-

cept of the Pachychilidae and Thiaridae, respectively

(Glaubrecht, 1996, 1999: Kéhler and Glaubrecht, 2001,

2002, 2003). Therefore, operculum morphology in S.

sulcospira corroborates the placement of the taxon with-

in the Pachychilidae. This, however, represents a ple-

siomorphic character state among the representatives of

this family, and is not a suitable character to establish

generic distinction.

By and large, the same can be stated for the radula.

The molluscan radula is generally considered a conser-

vative character (Fretter ‘and Graham, 1994). The pat-

tern described and depicted by Troschel (1858) (Figure

1) is commonly found among pachychilids (Kéhler and

Glaubrecht, 2001, 2002, 2003; Glaubrecht and Rintelen,

2003: Rintelen and Glaubrecht, 1999, 2003). This had

been already noticed by Troschel (1858), when allocating

taxa such as Pachychilus and Sulcospira (but also Me-

Page 24

THE NAUTILUS, Vol. 119, No. 1

lanopsis) under the “Pachychili*. However, we have

been unable to identify a single radular character pe-

culiar to Sulcospira, based on the limited anatomical fea-

tures discussed here. An enlarged main cusp is also

found in other pachychilid species, such as B. pagodula

(Kohler and Glaubrecht, 2001). Moreover, it is evident

that radular characters may especially be prone to ad-

aptation, parallelism, and convergence and that intraspe-

cific variability and plasticity in general may be consid-

erable, as described, e.g., for littorinid gastropods (Pa-

dilla, 1998; Reid and Mak, 1999; Reid, 2000). Thus, pos-

session of an elongated cusp alone is not considered a

characteristic suitable for the diagnosis of Sulcospira.

Gross anatomy of S. sulcospira andl S. martini remains

unknown, due to the lack of ethanol-preserved speci-

mens. It has been shown for other pachychilids, though,

that particularly characters of the reproductive organs

(pallial oviduct, gonads, brooding structures) and the

protoconch may bear essential systematic information

(Kohler and Glaubrecht, 2001, 2003).

Juvenile shells extracted from dried adults (Troschel,

1858; and own observations) deliver circumstantial evi-

dence that Sulcospira is viviparous. Furthermore, as

made evident from the following comparison, the pro-

toconchs of S. sulcospira are similar to those of some

species we have primarily assigned to the “Brotia-tes-

tudinaria-group” (Kéhler and Glaubrecht, 2001, e.g.,

Brotia testudinaria and B. hainanensis: Figures 29- 30).

In contrast, other taxa constituting the genus Brotia sen-

su stricto (denominated the “Brotia pagodula group” by

Kohler and Glaubrecht, 2001) have juveniles with an ir-

regularly wrinkled sculpture of the apical whorl of the

protoconch (Figures 25-26). Protoconchs of Tylomelania

and Pseudopotamis exhibit yet another fine morphology

(Figures 27-34). They attain a relative large size and

exthalit a relatively small apical whorl with a smooth shell

as well as regularly increasing whorls (Rintelen and

Glaubrecht, 1999, 2003; Glaubrecht and Rintelen,

2003).

As discussed in some detail by Kéhler and Glaubrecht

(2001), distinct protoconch morphologies of several pa-

chychilid genera are correlated with different reproduc-

tive strategies. For example, Brotia possesses a subhae-

mocoelic brood pouch while representatives of Tylome-

lania and Pseudopotamis are characterized by an eu-vi-

viparous mode of reproduction utilizing a modified

oviduct as brood pouch (= uterine proc pouch; Rin-

telen and Glaubrecht, 1999: Glaubrecht and Rintelen,

2003). Females of the latter two taxa retain a small num-

ber of embryos in the uterus that are nourished by se-

cretions produced by the albumen gland.

The protoconchs of Sulcospira correspond to those of

the so-called “Brotia testudinaria group” (Kéhler and

Glaubrecht, 2001), which indicates to us that S. swlcos-

pira possesses a subhaemocoelic brood pouch as well.

II. COMPARISON OF S. SULCOSPIRA AND S. MARTINI

Sulcospira martini, which has tentatively been allocated

to the genus by Kohler and Glaubrecht (2002), can be

distinguished by its shell, but much more conspicuously

by its ‘different ‘protoconch morphology, which is unique

among the Pachychilidae.

Because the protoconch morphology is related to the

mode of reproduction, it is assumed that S. martini ex-

hibits reproductive features (e.g. incubatory structure,

reproductive strategy) that may be distinct from those

known from any other pachychilid taxon.

III. SysteEMATIC CONCLUSIONS

Shell, operculum, and radula of S. sulcospira (and also

of S. martini) are typically pachychilid; a basally well

rounded and flared aperture, a round to oval, multispiral

operculum, and a rachidian tooth with an enlarged main

cusp flanked by up to three accessory cusps that taper

in size are diagnostic characters of this family. However,

these characters are symplesiomorphic and, thus, unin-

formative at the generic level. Soft body morphology,

which could bear crucial information, is not known. Still,

some systematic conclusions can be drawn based on the

evaluation of protoconch morphology, which is consid-

ered to be more or less constant at the generic level:

First, species of Brotia (Figures 27-28), Tylomelania

(Figures 31—32), and Pseudopotamis (Figures 33-34) are

not congeneric with S. sulcospira (see Figures 19-20)

since they exhibit each a distinct protoconch, which is

testimony to a different reproductive strategy as dis-

cussed above. The same holds true for Jagora (Kéhler

and Glaubrecht, 2003). All these pachychilid genera

have been shown to represent independent monophy-

letic lineages characterized by morphological features,

such as peculiar reproductive morphologies. Second, the

protoconch of S. sulcospira is very similar to that known

from species of the “Brotia-testudinaria-group” denom-

inated by Kéhler and Glaubrecht (2001) (Figures 29, 30

for B. hainanensis). Hence, Sulcospira Troschel, 1858,

being available and valid, might be an appropriate ge-

neric name for this species group under the precondition

that it can be shown that a protoconch with a smooth

and dome-shaped apical whorl is a character possessed

by the members of this group and derived by shared

ancestry. However, the alterative explanation that a

similar protoconch represents an ancestral state that is

present in two different lineages has to be ruled out.

Otherwise, inferring generic relationship in absence of a

phylogenetic evaluation could lead to the erection of

paraphyletic taxa when the characters considered are

plesiomorphic. Before we cannot show by phylogenetic

analyses of morphological or molecular data that S. sul-

cospira and species of the “Brotia testudinaria group”

indeed belong to the same taxon, we therefore refrain

from a respective taxonomic suggestion.

Third, we conclude that S$. martini can be recognized

as a species and that it does not represent a synonym of

S. sulcospira. Furthermore, it is clear that the morphol-

ogy of the protoconch of S. martini does resemble nei-

ther of the known pachychilid genera very closely (Fig-

ures 27-34) including that of cn sulcospira (Figures 19-

F. Kohler and M. Glaubrecht, 2005

20). Just based on this single feature, it could be de-

duced that S. martini might be a representative of a yet

undescribed genus. However, as has been stated for S.

sulcospira, a sound decision on its systematics should

rely on a more comprehensive data set.

For the time being, we suggest to maintain Sulcospira

as a monotypic genus endemic to Java. Furthermore, we

refrain from a taxonomic decision on the generic rela-

tionship of “Sulcospira” martini. It likely is not a mem-

ber of one of the described pachychilid ¢ genera, but its

true relationships remain unknown.

Irrespective of our anticipation that both species dealt

with in this paper may have already become extinct in

large parts of their original distribution area, we still

hope that suitable material will turn up, eventually al-

lowing to find an answer to the remaining questions and

to solve another of the many puzzling aspects of pachy-

chilid phylogeny and systematics that long hampered a

deeper understanding of the evolution of this intriguing

and instructive case study among the lymnic Cerithioi-

dea.

ACKNOWLEDGMENTS

We are most grateful to the curators Philippe Bouchet

(MNHN), Yves Finet (MNHG), Edmund Gittenberger

and Jeroen Goud (RMNH), Ronald Janssen (SMF), Tru-

di Meier (ZMZ), Robert Moolenbeek (ZMA) and Kathie

Way (BMNH) for making material of their collections

available to us. Thomas von Rintelen kindly provided

SEM images of a juvenile shell of Tylomelania patriar-

chalis. We thank Philippe Bouchet and an anonymous

reviewer for most helpful comments on the manuscript,

in particular as related to nomenclatorial aspects. Their

comments helped much to improve the quality of this

paper. The work of F. K. was funded through a post-

graduate scholarship of the Konrad-Adenauer-Stiftung,

Sankt Augustin (Germany). This paper is part of a pro-

ject on Southeast Asian pachychilids supported through

grant GL 297/4 to M.G. by the Deutsche Forschungs-

gemeinschaft.

LITERATURE CITED

Abbott, R. T. 1948. Handbook of medically important molluscs

of the Orient and the western Pacific. Bulletin of the Mu-

seum of Comparative Zoology 100: 285-299.

Adam, W. and E. Leloup. 1938. Prosobranchia et Ophisto-

branchia. In: Straelen, V. van (ed.). Resultats scientifiques

du voyage aux Indes Orientales Néerlandaises de LL. AA.

RR. le Prince et la Princesse Léopold de Belgique. Mé-

moires du Musée Royal d'Histoire Naturelle de Belgique

2(19): 1-209.

Adams, H. and A. Adams. 1858. Genera of recent molluscs

arranged according to their organization, 1. John v. Voorst,

London, 661 pp.

Benthem-Jutting, W. S. S. v. 1956. Systematic studies on the

non-marine Mollusca of the Indo-Australian archipelago.

5. Critical revision of the Javanese freshwater gastropods.

Treubia 23(2): 259-477.

Page 25

Benthem-Jutting, W. S. S. v. 1959. Catalogue of the non-marine

Mollusca of Sumatra and of its satellite islands. Beaufortia

TAI

poueeee O. 1890. Ad. Strubell’s Konchylien aus Java I. Bericht

dex senckenbergischen naturforschenden Gesellschaft. pp:

SINS,

Bouchet, P. and J. P. Rocroi. Classification and nomenclator of

gastropod families. Malacologia (submitted).

Brandt, R. A. M. 1968. Description of new non-marine mol-

lusks from Asia. Archiv fiir Molluskenkunde 98(5/6): 213—

289.

Brandt, R. A. M. 1974. The non-marine aquatic Mollusca of

Thailand. Archiv fiir Molluskenkunde 105(1/4): 1-423.

Brot, A. 1870. Catalogue of the recent species of the family

Melanidae. American Journal of Conchology 6: 271-325.

Brot, A. 1874. Die Melaniaceen (Melanidae) in Abbildungen

nach der Natur mit Beschreibungen. In: Martini, F. H.

W. and Chemnitz, J. H. (ed.). Systematisches Conchylien-

Cabinet, 1(24). Bauer & Raspe, Niirnberg, 488 pp.

Dudgeon, D. 1982. The life history of Brotia hainanensis

(Brot, 1872) (Gastropoda: Prosobranchia: Thiaridae) in a

tropical forest stream. Zoological Journal of the Linnean

Society 76: 141-154.

Dudgeon, D. 1989. Ecological strategies of Hong Kong Thiar-

idae (Gastropoda: Prosobranchia). Malacological Review

Me BAB.

Dudgeon, D. 2000. Conservation of freshwater biodiversity in

oriental Asia: constraints, conflicts, and challenges to sci-

ences and sustainability. Limnology 1(3): 237-243.

Fretter, V. and A. Graham. 1994. [Siatslh Prosobranch Molluscs.

24 ed. The Ray Society, Andover, 819 pp.

Glaubrecht, M. 1996. Evolutionsékologie und Systematik am

Beispiel von Siif- und Binelawasterechineelen (Mollusca:

Caenogastropoda: Cerithioidea): Ontogenese-Strategien,

paliiontologische Befunde und Historische Zoogeogra-

phie. Backhuys Publishers, Leiden, 499 pp.

Glaubrecht, M. 1999. Systematics and the evolution of vivi-

parity in tropical freshwater gastropods (Cerithioidea:

Thiaridae sensu lato)—an overview. Courier Forschung-

sinstitut Senckenberg 125: 91-96.

Glaubrecht, M. and T. v. Rintelen. 2003. Systematics, molec-

ular genetics and historical zoogeography of the viviparous

freshwater gastropod Pseudopotamis (Cerithioidea, Pachy-

chilidae): a relic on the Torres Strait Islands, Australia.

Zoologica Scripta 32: 415-435.

Holznagel, W. E. 1998. A nondestructive method for cleaning

gastropod radulae from frozen, alcohol-fixed, or dried ma-

terial. American Malacological Bulletin 14: 181-183.

International Commission on Zoological Nomenclature. 1999.

International Code of Zoological Nomenclature, Fourth

edition. The International Trust for Zoological Nomencla-

ture, London, 306 pp.

Knipper, H. 1958. Die Typen und Typoide des Uberseemu-

seums Bremen, 5: Mollusca (Gastrop. Prosobranch.): Ner-

itidae und Thiaridae. Veréffentlichungen des Ubersee-

museums Bremen, Reihe A 3(1): 39-74.

Kohler, F. 2003. Brotia in space and time. Phylogeny and evo-

lution of Southeast Asian freshwater gastropods of the

family Pachychilidae (Caenogastropoda, Cerithioidea).

Unpublished Ph.D. Thesis, Humboldt University Berlin.

363 pp

K6hler, F. and M. Glaubrecht. 2001. Toward a systematic re-

vision of the Southeast Asian freshwater gastropod Brotia

H. Adams, 1866 (Cerithioidea: Pachychilidae): an account

Page 26

of species from around the South China Sea. Journal of

Molluscan Studies 67: 281-318.

Kohler, F. and M. Glaubrecht. 2002. Annotated catalogue of

the nominal taxa of Southeast Asian freshwater gastro-

pods, family Pachychilidae Troschel, 1858 (Mollusca:

Caenogastropoda: Cerithioidea), with an evaluation of the

types. Mitteilungen aus dem Zoologischen Museum, Ber-

lin, 78: 121-156.

Kohler, F. and M. Glaubrecht. 2003. Morphology, reproductive

biology and molecular genetics of ovoviviparous freshwa-

ter gastropods (Cerithioidea: Pachychilidae) from the Phil-

ippines, with description of the new genus Jagora. Zool-

ogica Scripta 32(1): 35-59.

Leschke, M. 1914. Zur Molluskenfauna von Java und Celebes.

Mitteilung des naturhistorischen Museums Hamburg 21:

205-284.

Lydeard, C., W. E. Holznagel, M. Glaubrecht and W. F. Pon-

der. 2002. Molecular phylogeny of a circum-global, diverse

gastropod superfamily (Cerithioidea: Mollusca: Caenogas-

tropoda): Pushing the deepest phylogenetic limits of mi-

tochondrial LSU rDNA Sequences. Molecular Phyloge-

netics and Evolution 22: 399-406.

Martens, E. v. 1897. Sii8- und Brackwasser-Mollusken des In-

dischen Archipels. In: Weber, M. (ed.). Zoologische Er-

gebnisse einer Reise in Niederlaindisch Indien (4). Brill,

Leiden, 331 pp.

Morrison, J. P. E. 1954. The relationship of old and new world

melanians. Proceedings of the United States National Mu-

seum 103: 357-394.

Mousson, A. 1849a [1848]. Uber die Land- und

Siibwassermollusken von Java. Mittheilungen der Natur-

forschenden Gesellschaft, Ziirich 1: 264-273.

Mousson, A. 1849b. Die Land- und Siisswasser-Mollusken von

Java. Schulthess, Ziirich, 126 pp.

Oostingh, C. H. 1935. Die Mollusken des Plioziins von Boe-

miajoe (Java). Wetenschappelijke Mededeelingen 26: 1-

Sie

Padilla, D. K. 1998. Inducible phenotypic plasticity of the rad-

ula in Lacuna (Gastropoda: Littorinidae). The Veliger 41:

201-204.

Ponder, W. F. and A. Warén. 1988. A systematic list of the

family-group names and higher taxa in the Caenogastro-

poda and Heterostropha. In: Ponder, W. F. (ed.) Proso-

branch Phylogeny. Proceedings of a Symposium held at

the 9th International Malacological Congress, Edinburgh,

1986. Malacological Review, Supplement 4; 288-328.

Reid, D. G. 2000. The use of the radula in the taxonomy and

phylogeny of gastropods: cautionary cases of convergence,

THE NAUTILUS, Vol. 119, No. 1

intraspecific variation and plasticity, Phuket Marine Bio-

logical Center Special Publication 21(2): 329-345.

Reid, D. G. and Y.-M. Mak. 1999. Indirect evidence for eco-

phenotypic plasticity in radular dentition of Littoraria spe-

cies (Gastropoda: Littorinidae). Journal of Molluscan

Studies 65: 355-370.

Rensch, B. 1934. Siifwassermollusken der deutschen limnol-

ogischen Sunda-Expedition. Archiv fiir Hydrobiologie

Supplement 8: 203-254.

Rintelen, T. v. 2003. Phylogenetic analysis and systematic re-

vision of a species flock of viviparous freshwater gastro-

pods in the ancient lakes on Sulawesi (Indonesia)—a mod-

el case of adaptive radiation?. Unpublished Ph.D. Thesis,

Humboldt University Berlin. 259 pp.

Rintelen, T. v. and M. Glaubrecht. 1999. On the reproductive

anatomy of freshwater gastropods of the genera Brotia H.

Adams, 1866 and Tylomelania Sarasin and Sarasin, 1897

in the central lakes on Sulawesi, Indonesia (Cerithioidea:

Melanatriidae). Courier Forschungsinstitut Senckenberg

125: 163-170.

Rintelen, T. v. and M. Glaubrecht. 2003. New discoveries in

old lakes: three new species of Tylomelania Sarasin and

Sarasin, 1897 (Gastropoda: Cerithioidea: Pachychilidae)

from the Malili lake system on Sulawesi, Indonesia. Jour-

nal of Molluscan Studies 69: 3-18.

Robertson, R. 1957. Publication dates of Troschel’s “Das Ge-

biss der Schnecken”. The Nautilus 70: 136-138.

Schepmann, M. M. 1896. Descriptions of new Melaniidae.

Notes from the Leyden Museum 18: 135-139.

Schepmann, M. M. 1898. Conchological corrections and ad-

ditions. Notes from the Leyden Museum 20: 84-86.

Subba Rao, N. V. 1989. Handbook, Freshwater Mollusks of

India. Zoological Survey of India, Calcutta, 289 pp.

Thiele, J. 1929. Handbuch der Systematischen Weichtierkun-

de. Gustav Fischer, Jena, 376 pp.

Troschel, F. H. 1856-1863. Das Gebiss der Schnecken zur Be-

griindung einer natiirlichen Classification. Nicolaische

Verlagsbuchhandlung, Berlin, 252 pp.

Vaught, K. C. 1989. A classification of the living Mollusca.

American Malacologists Inc., Melbourne, Florida, 189 pp.

Wanner, H. 1984. Heinrich Zollinger, 1818-1859. Ein Ziiricher

Schulmann als Naturforscher und Pflanzen in Indonesien.

Sein Leben und seine Zeit. Vierteljahrsschrift der Natur-

forschenden Gesellschaft in Ziirich 128(5): 1-32.

Yen, T.-C. 1939. Die chinesischen Land- und SiifSwasser-Gas-

tropoden des Natur-Museums Senckenberg. Abhandlun-

gen der senckenbergisch-naturforschenden Gesellschaft

444. |—233.

THE NAUTILUS 119(1):27-42, 2005

Page 27

Six new species of Paryphantopsis (Gastropoda: Pulmonata:

Charopidae) from the Papuan Peninsula of New Guinea

John Slapcinsky

Florida Museum of Natural History

University of Florida

Gainesville, FL 32611 USA

[email protected]

ABSTRACT

Six new species of Paryphantopsis, a genus of charopid snails

endemic to New Guinea, are described from the geologically

complex eastern terminus of the Papuan Peninsula. All descrip-

tions are based on material collected in 2002-2003 and include

information on shell, genital, and radular anatomy. Of the 14

previously described species, genital anatomy was known for

three and radular morphology for only one species. Examina-

tion of these under-utilized character sets has uncovered tax-

onomically useful interspecific variation. The six new species

appear to have distributions limited to particular mountain

groups on the extreme eastern terminus of the Papuan Pen-

insula. Species in close proximity or sympatry share unique

shell, genital and radular characters suggesting local speciation

and poor dispersal ability. Diversity in Paryphantopsis has been

underestimated and it is likely that there are many species yet

to be discovered with narrow ‘geographic and ecological ranges

in the under-explored mountains of New Guinea.

INTRODUCTION

Paryphantopsis, a genus of charopid snails endemic to

New Guinea, are found at moderate and high altitudes

(600-4000 m) and are distributed from Western Papua

(Irian Jaya) to the Louisiade Archipelago. The genus

contains fourteen previously described species: P. arcu-

ata Jutting, 1964, P. dualoensis Solem, 1970, P. elegans

(Fulton, 1902), P. filosa Jutting, 1964, P. fultoni ( (Coen,

1922), P. P. globosa (Hedley, 1890), P. lamelligera (Thiele,

1928), P. ieesor Jutting, 1964, P. looistaderine (Mollen-

dorff, 1899), P. platycephala Jutting, 1964, P. pygmaea

(Bavay, 1908), P. sculpturata Jutting, 1964, P. similis

(Thiele, 1928), and P. striata (Fulton, 1902). Solem

(1970) reviewed the genus, redescribing all species ex-

cept those then recently described or reviewed by Jut-

ting (1964). Most Paryphantopsis species are known

from small samples of shells, often only from their type

localities, and data on genital morphology are limited to

three species, and on radular morphology to only one

species (Solem, 1970, Wiktor, 2003). Paryphantopsis has

not been reported previously east of longitude 148° E

on the relatively poorly sampled eastern part of New

Guinea, which is known as the Papuan Peninsula. The

eastern end of the peninsula includes the Cloudy Moun-

tains to the south, and the disjunct terminus of the

Owen Stanley Range to the north, separated from the

main Owen Stanley uplands by extensive lowlands west

of Mount Suckling. The geologically complex Papuan

Peninsula is rammed largely. by the East Papua Compos-

ite Terrane (EPCT), a tectonic province composed of at

least 4 separate geological units with differing ages, or-

igins, and histories. These units appear to have assem-

bled northeast of modern New Guinea during the Pa-

leocene, 62-57 Myr ago, and fused to the main body of

be island in the Late Oligocene to Early Miocene, 28-

2 Myr ago (Pigram and Davies, 1987). Because of its

fale “anita offshore amalgamation, the EPCT may have

developed a distinct and largely endemic biota. This is

the first in a series of papers describing the results of

ten weeks of field surveys that took place during April—

May, 2002 and January—M March, 2003. These surveys ex-

plored the extreme eastern terminus of the Papuan Pen-

insula where two geological units lie in close proximity,

the Cloudy Mountains of the Port Moresby Terrane and

eastern terminus of the Owen Stanley Range of the Kutu

Terrane (Figure 1).

MATERIALS AND METHODS

Specimens were hand-collected or sifted from samples

of leaf-litter. Live collected animals were drowned over-

night and then preserved in 75% ethanol. Gross anatom-

ical dissections were made under 75% ethanol using a

dissecting microscope. Radulae were isolated from dis-

seated! Ibmewll messes using a saturated KOH solution.

Scanning electron micrographs of radulae were made us-

ing a field emission SEM. Drawings of the genital anat-

omy were made with the assistance of a camera lucida,

and measurements were taken using an ocular microm-

eter. Shell measurements were made as follows. Whorl

count (W) was measured from the suture of the first

whorl to the body whorl and fractions of a whorl were

determined with the aid of a cardboard circle divided

into 10 equal parts of 36° (Figure 2, line 1—-2.9). Spire

Page 28

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S 92

S Loe

S die

E 149°

E 150°

0-100m

100-1000m

1000-2000m

2000-3000m

= Se

Figure 1. Distribution of Paryphantopsis on the eastern terminus of the Papuan Peninsula, Papua New Guinea between 9° S,

149° E and 11° S, 151° E. A = P. abstrusa, K = P. koragae, L = P. lebasii, M = P. matawanensis, U = P. ubwamensis, Y = P.

yawii, O = other sites sampled.

diameter (SD) was the length of a straight line passing

from the apertural edge of the suture through the mid-

dle of the apex to the opposite suture (Figure 2, line A—

B). Diameter (D) was the greatest width of the shell

perpendicular to the shell axis (Figure 3, line C—D).

Height (H) was the greatest distance between the apex

and the base of the aperture measured parallel to the

shell axis (Figure 3, line D-E). Spire height (SH) was

measured from the top of the body whorl to the apex of

the shell (Figure 3, line F-G). Aperture width (AW) was

the greatest distance from the columellar edge to the

outer edge of the aperture (Figure 3, line E-H). Aper-

ture height (AH) was measured from the suture to the

base of the aperture, parallel to the shell axis (Figure 3,

line H-I). The lengths of radular teeth were measured

from the top of the mesocone to the posterior edge of

the basal plate. The widths of radular teeth were mea-

sured as the greatest width of the cusps, not the basal

plate. The following abbreviations are used in figures of

genital anatomy: AT = atrium, DI = diverticulum, EP

= epiphallus, OV = free oviduct, PE = penis, PG =

prostate gland, PP = penial pilasters, PR = penial re-

tractor muscle, SD = spermathecal duct, SP = sper-

matheca, VA = vagina, VD = vas deferens, VP = vergic

J. Slapcinsky, 2005

Periostracal

Extensions

Periostracal

Processes

Page 29

Figures 2-3. Diagram of shell measurements. 2. Whorl count (line 1-2.9), spire width (line A-B). 3. Diameter (line C-D),

height (line D-E), spire height (line F—G), aperture width (line E—H), aperture height (line H—I).

papillae. Specimens are deposited in the following insti-

tutions: Bernice P. Bishop Museum, Honolulu (BPBM),

Florida Museum of Natural History, Gainesville (UF),

Papua New Guinea National Museum, Port Moresby

(PNGNM), Wroclaw University Museum of Natural

History (MNHW).

SYSTEMATICS

Family Charopidae Hutton, 1884

Genus Paryphantopsis Thiele, 1928 (Type species:

Flammulina (Paryphantopsis) lamelligera Thiele, 1928,

by original designation.)

Description: Moderate to large-sized charopid snails

with loosely coiled shells of approximately three whorls.

Shell shape varies among species from globose to de-

pressed with an elevated to flat spire and a rounded to

keeled margin. The umbilicus, usually covered by an ex-

pansion of the peristome, is sometimes perforate. Nu-

clear whorls (protoconch) are sculptured with spiral rows

of small pits that become weaker and less regular on

later whorls. Postnuclear whorls (teleoconch) are usually

sculptured with growth lines accentuated with short per-

iostracal extensions punctuated with occasional longer

extensions at regular intervals. These longer extensions

can bear additional processes at the shell margin (Figure

2). A few species do not have longer periostracal exten-

sions and some species do not bear any extensions. Body

color is usually yellow in life. The epiphallus is apically

inflated, often with an apical diverticulum. The penis is

textured with convoluted pilasters of varying complexity.

Central teeth of the radula are tricuspid and of similar

size to the tricuspid and slightly assymetrical lateral

teeth. The many lateral teeth become shorter and less

symmetrical, grading in shape with the marginal teeth.

Marginal teeth are usually assymetric, their endocones

longer than their ectocones. Endocones and ectocones

and less often mesocones can bear accessory cusps.

Paryphantopsis abstrusa new species

(Figures 4-10, Table 1)

Description: The adult shell is small for the genus,

4.4-4.8 mm (mean = 4.6) in diameter and 3.5-3.8 mm

(mean = 3.6) in height, with 2.9-3.0 (mean = 3.0) rap-

idly expanding whorls (Figures 4-6, Table 1). The spire

is elevated 0.3 mm. Postnuclear whorls descend regu-

larly and the shell height/diameter ratio is 0.76—-0.81

(mean = 0.79). The shell has 1.3 evenly rounded nuclear

whorls, sculptured with 12 spiral rows of small pits. The

postnuclear whorls bear weak and irregular malleations,

most readily visible at the base, and indistinct and irreg-

ular striae on the apical surface. The postnuclear whorls

are also sculptured with weak growth lines. Approxi-

mately every fourth growth line is accentuated by a per-

iostracal extension. The periostracal extensions bear

small (0.15 mm) trigonal processes at the periphery.

These processes occur on every periostracal extension

for the first two whorls, then become less regular and

finally absent from the final % whorl. The fragile pro-

cesses are often partially worn from the earlier whorls

of adult shells. The nuclear whorls are white, the post-

nuclear whorls brown. The umbilicus is closed by a re-

flection of the peristome. The aperture is large, ovate to

nearly circular, with an aperture-diameter to aperture-

height ratio of 0.54-0.67 (mean = 0.63).

The body color is uniform bright yellow-white in life,

fading to cream in specimens preserved in ethanol. The

vas deferens narrows rapidly from the prostate gland and

remains narrow to the swollen head of the epiphallus

(Figure 7). A long coiled diverticulum inserts laterally

on the epiphallus soon after the junction with the vas

deferens. The epiphallus narrows slightly after the di-

verticulum and widens towards the junction with the pe-

nis. The penis is half the length and three times the

width of the epiphallus, robust, widest centrally, and nar-

rowing slightly at each end. Apically there are several

small convoluted pilasters and one much larger conyvo-

luted pilaster that extends to near the base (Figure 8).

The penial retractor muscle is robust, originating from

Page 30 THE NAUTILUS, Vol. 119, No. 1

35, diameter 4.6 mm. 7—8. Camera lucida

drawing of genitalia, UF 299677, maximum width 7.2 mm. 9-10. Scanning electron micrograph of radula, UF 299677, field width

of central and lateral teeth 41 jzm, marginal teeth 42 xm.

J. Slapeinsky, 2005

Page 31

Table 1. Measurements in mm of undamaged adult shells of six species of Paryphantopsis, N = count, H = height, D = diameter,

SH = spire height, SD = spire diameter, AH = aperture height, AD = aperture diameter, W = number of whorls.

Species N H D SH SD AH AD W

P. abstrusa 3 mean=SD 36+02 46+02 O03+£00 21+00 26+02 28+01 30+01

range 3.5-3.8 44-48 0.3-0.3 2.0-2.1 9) ALS) 7/ 2.7-2.9 2.9-3.0

P. koragae o men 2 SID 2Osa Ol 422200 OF = 00 IGS Ol Sls O BOs Ol Ass 2 Op

range 2.83.0 4-42 0.1-0.2 1.5-1.7 2.0-2.1 D5 9.7-2.8

P. lebasii AML mage = SID) BOs O82 COfjseOQo Ola Oi Bsa O2 Ws ae Oe 2he) se Ozh bys} se Ohl

range 3.3-4.0 6.1-7.5 0.0-0.2 2.32.8 2.43.0 3.9-4.9 2.7-2.9

P. matawanensis >) mean = SDE Aa = Os 70204 01 =O 2603" Boz 0s 452 04 2:9) = 0:2

range 4.0-4.9 6.6-7.3 0.0-0.1 29-28 3.24 | 41-48 2.7-3.0

P. ubwamensis 3 mg@ms GD S0=02 BLE O7 OB=Ol BOSOS B8=203 43205 30) = Ol

range 4.85.1 6.4-7.7 0.2-0.4 DTBS 3.64.1 3.9-4.9 2.9-3.1

P. yawii 3 means SD Sl=O3 S9=O4 O2=O01 B®=OS B=O08 B7=O4 87 =O

range 2.6-3.6 5.0-6.5 0.1-0.3 1.9-2.6 YO—7/ 3.24.1 2.6-2.8

the diaphragm and inserting at approximately mid-point

on the epiphallus. The spermathecal duct is robust, nar-

rowing abruptly at mid-point and remaining narrow until

joining the relatively small, ovate spermatheca. The free

oviduct joins the moderate length vagina above the atri-

um.

The central teeth of the radula (second row from left)

are tricuspid, 8-9 wm wide and 11-12 pm long, roughly

the same shape as, but smaller than, the fost lateral

teeth, which are 9-10 zm wide and 12-13 wm long (Fig-

ure 9). The mesocones of both the central and fase he

eral teeth are tall, slender, and blade-shaped, joining the

rectangular basal plates close to, but not on, their pos-

terior edge. The mesocones of the central teeth barely

project beyond the anterior edge of the basal plates,

those of the lateral teeth project well beyond the edge.

The ectocones are trigonal and short only one third of

the height of the mesocones, joining the posterior edge

of the basal plates. The lateral teeth are asymmetrical,

their endocones are slightly taller then their ectocones.

The marginal teeth are dorsoventrally compressed and

tricuspid to multicuspid, 8-9 wm wide and 11-12 wm

long (Figure 10). The endocones of the marginal teeth

are half the height of the mesocones and only slightly

taller than the ectocones. The mesocones of the margin-

al teeth often bear small cusps near the mid-point.

Type Material: Holotype: UF 308235, J. Slapcinsky,

16 April 2002; Paratypes: UF 299667 (2 specimens), UF

303588 (4 specimens), type locality, J. Slapcinsky, 16

April 2002.

Type Locality: Papua New Guinea, Milne Bay Prov-

ince, Cloudy Mountains, Ubwam Mountain, headwaters

of the Watuti River, 10° 29.8’ S, 150° 14.02’ E, 670 m

altitude.

Habitat: Observed crawling near the base of trees

with smooth bark, in native forest, during wet weather

at 670 meters altitude.

Etymology: From the Latin abstrusa, a feminine ad-

jective meaning hidden, concealed and reserved, allud-

ing to the difficulty finding this species and to its subtle

almost concealed periostracal processes.

Remarks: Paryphantopsis abstrusa differs from most

other Paryphantopsis smaller than 5 mm in diameter by

having periostracal extensions with processes at their

margins. Paryphantopsis similis is the only other small

species with periostracal extensions. It is more de-

pressed, with a height/diameter ratio of 0.70 compared

to a height/diameter ratio of 0.79 in P. abstrusa.

Paryphantopsis koragae new species

(Figures 11-17, Table 1)

Description: The adult shell is small for the genus,

4.14.2 mm (mean = 4.2) in diameter and 2.8-3.0 mm

(mean = 2.9) in height, with 2.7-2.8 (mean = 2.8) rap-

idly expanding iho (Figures 11-13, Table 1). The

spire is slightly elevated, 0.1-0.2 mm (mean = 0.2),

postnuclear. whorls descend slowly and regularly. Shell

height/diameter ratio is 0.67-0.71 (mean = 0: 69). There

are 1.1 evenly rounded nuclear whorls, sculptured with

12 spiral rows of small pits that are not continued on

the postnuclear whorls. The postnuclear whorls have

regular growth wrinkles with irregular, short, periostracal

exiensions that are slightly coal basally. None of the

periostracal extensions extend further than the others.

The nuclear whorls are white, the postnuclear whorls are

dark brown. The umbilicus is closed by a reflection of

the peristome. The aperture is large, flattened apically

and slightly angled at the periphery and base of the col-

umella. The aperture-diameter to aperture-height ratio

is 0.78—0.80 (mean = 0.79).

The body color is uniform yellow im life, fading to

cream in specimens preserved in ethanol. The vas def-

erens narrows rapidly after the prostate gland and re-

mains narrow until entering the swollen, ovate head of

the epiphallus (Figure 14). The interior of the head of

the epiphallus bears two strong pilasters that enter the

short broad diverticulum, which is roughly one quarter

of the length of the epiphallus. The epiphallus is two to

three times longer than the penis and one third narrower

Page 32 THE NAUTILUS, Vol. 119, No. 1

Figures 11-17. Paryphantopsis koragae. 11-13. Photographs of shell, Holotype UF 308237, diameter 4.2 mm. 14-15. Camera

lucida drawing of genitalia, UF 303586, maximum width 6.3 mm. 16-17. Scanning electron micrograph of radula, UF 303586,

field width of central and lateral teeth 55 zm, marginal teeth 41 wm.

J. Slapeinsky, 2005

at the junction with the penis. The very short penial

retractor muscle originates on the diaphragm and inserts

near the mid-point of the epiphallus. The interior of the

penis bears several smooth, low, regular pilasters and

one much larger convoluted pilaster (Figure 15). The

spermatheca is oblong-ovate, its duct is apically narrow

and widens abruptly at the mid-point. The free oviduct

is relatively robust, joining the long vagina well above

the atrium.

The central teeth of the radula (center row) are tri-

cuspid, 9-10 zm wide and 12-13 wm long, slightly

smaller than the first lateral teeth, which are 10-11 wm

wide, 13-14 ym long (Figure 16). The mesocones of

both central and lateral teeth barely project beyond the

basal plate. The ectocones of the central and lateral

teeth are about one half the height of the mesocones.

The lateral teeth are tricuspid and very slightly assy-

metric with the endocone of each lateral slightly taller

than the ectocone. The marginal teeth are dorsoventrally

compressed and tricuspid or weakly and irregularly mul-

ticuspid, 8-13 jzm wide and 8-10 «wm long (Figure 17).

The endocones of the marginal teeth are nearly the

height of the mesocones and can have very weak sec-

ondary cusps; the ectocones are shorter and unicuspid

to irregularly multicuspid.

Type Material: Holotype: UF 308237, J. Slapcinsky,

20 February 2003; Paratypes: Papua eas Guinea, Milne

Bay Province, Mount Matawan (Mount eee UF

303586 (2 specimens), UF 303587 (2 specimens), type

locality; UF 303584 (1 specimen), plateau ENE of sum-

mit, 10°2.1’ S, 149°34.6’ E, 2567 m altitude, J. Slapcin-

sky, 17 February 2003; UF303585 (1 specimen), Bunisi

Village, 10°1.1' S, 149°36.2 E, 1450 m altitude, J. Slap-

cinsky, 16 February 2003.

Type Locality: Papua New Guinea, Milne Bay Prov-

ince, Mount Nie (Mount Simpson): NE of summit,

10°1.7' S, 149°34.7' E, 2100 m altitude.

Habitat: Active on plants and leaf litter usually near

the ground. Observed aestivating in suspended leaf litter

within 1.5 m of the ground in tropical hardwood forest

and cloud forest from 1450 m to 2600 m altitude.

Etymology: This matronym honors Ms. Helen Kor-

age, Councelor for the Village of Ikara, who facilitated

our access to the Mount Matawan area.

Remarks: Paryphantopsis koragae differs from other

species of Paryphantopsis in being sculptured with

growth lines accentuated with very short periostracal ex-

tensions. All other small species (= 5 mm diameter),

including P. filosa, P. pygmaea, P. arcuata, P. sculpturata,

P. similis, P. platycephala, and P. abstrusa have occasion-

al longer periostracal extensions. The anatomy of only

one Paryphantopsis species of similar size has been fig-

ured previously: P filosa from Karkar Island near Ma-

dang (Wiktor, 2003, fig. 9). This species differs from P.

koragae in not having a diverticulum on the epiphallus.

Page 33

Paryphantopsis lebasii new species

(Figures 18-24, Table 1)

Description: The adult shell is slightly larger than av-

erage for the genus, 6.1-7.5 mm (mean = 6.8) in di-

ameter and 3.3-4.0 mm (mean = 3.6) in height, with

2.7-2.9 (mean = 2.8) rapidly expanding whorls (Figures

18-20, Table 1). The spire is flat to slightly elevated, 0.0-

0.2 mm (mean = 0.1). Postnuclear whorls descend slow-

ly and regularly and shell height/diameter ratio is 0.45—

0.61 (mean = = 0) 53). There are 1.3 nuclear whorls, with

weak, peripheral and supraperipheral angles, and sculp-

tured with about 12 spiral rows of small pits. These pits

become larger and less regular on the postnuclear

whorls, where they are visible through the periostracum

as weak malleations. The sculpture of spiral rows of ob-

long pits is clearer where the periostracum is removed.

The shell periphery is weakly keeled. Apical surface of

the whorls is broadly rounded or with a very weak su-

praperipheral angle. Regular growth wrinkles accentu-

ated with short periostracal extensions are present on

the postnuclear whorls. On the apical surface, these ex-

tensions are folded along their length towards the ap-

erture and are weakly appressed to the shell. Approxi-

mately every fifth periostracal extension protrudes about

0.5 mm beyond the shell margin forming large rectan-

gular processes. These processes overlap "each other on

the penultimate whorl, but not on the body whorl. They

are approximately equally spaced, rectangular, distally

rounded, and of approximately equal length. Periostracal

extensions of the growth lines are showter erect, and less

prominent basally. Nuclear whorls are white; postnuclear

whorls pale } yellow brown. The umbilicus is perforate or,

less often, closed, covered to varying degrees by a re-

flection of the peristome. The aperture is large, de-

pressed-ovate with an aperture-diameter to aperture-

height ratio of 0.51-0.75 (mean = 0.65).

The body color is uniform bright creamy-yellow in

life, fading to creamy-white in specimens preserved in

ethanol. The vas deferens narrows to the junction with

the ovate head of the epiphallus (Figure 21). The epi-

phallus is approximately one quarter the diameter of the

penis and does not bear a diverticulum. The penial re-

tractor muscle is a little less than half the length of the

epiphallus, originating from the diaphragm and inserting

at the base of the epiphallus. The robust penis is a little

shorter than the epiphallus, with three strong pilasters

that run its entire length (Figure 22). The atrium is

short, expanding slightly toward the junction with the

penis and vagina. The spermathecal duct is massive at

the base, tapering rapidly at mid point, the remainder is

relatively narrow until its junction with the spherical

spermatheca. The free oviduct is slightly coiled and nar-

row, joining the very short vagina just above the atrium.

The central teeth of the radula (fifth row from left)

are symmetrically tricuspid, 8-9 jzm wide and 13-14 wm

long, and are similar in shape and length to the slightly

niger (9-10 um), and slightly asymmetrical lateral teeth

(Figure 23). The bluntly conical and erect mesocones of

Page 34 THE NAUTILUS, Vol. 119, No. 1

D fron ee AD 22

21-22. Camera

lucida drawing of genitalia, UF 299671, maximum width 6.7 mm. 23-24. Scanning electron micrograph of radula, UF 299671,

field width of central and lateral teeth 67 jzm, marginal teeth 38 jxm.

J. Slapcinsky, 2005

Page 35

the central and lateral rows join their basal plates cen-

trally and barely project beyond the anterior of their bas-

al plates. The ectocones of both the central and lateral

rows are trigonal and short, about half the height of the

mesocones; they join the posterior edge of their basal

plates at a low buttress. The endocones of the lateral

teeth are slightly larger but otherwise of similar shape

to their ectocones. The marginal teeth are dorsoventrally

compressed, multicuspid, about 9-10 jum wide and 10-

11 wm long (Figure 24). The endocones are nearly the

same height as the mesocones while the ectocones are

much shorter, about one half to one third the height of

the mesocones and divided into three cusps.

Type Material: Holotype: UF 308233, J. Slapcinsky,

9 April 2002; Paratypes: Papua New Guinea, Milne Bay

Province: UF 299676 (1 specimen); UF 299699 (5 spec-

imens), Cloudy Mountains, Ubwam Mountain, headwa-

ter of the Watuti River, 10°29.8’ S, 150°14.0’ E, 675

meters altitude, J. Slapcinsky, 16 April 2002; UF 299674

(8 specimens); UF 303593 (4 specimens), Pini Range,

Duabo Mission Station, 10°25’ 05” S, 150°18’ 24 E, 325

meters altitude, J. Slapcinsky, 9 April 2002; BPBM (2

specimens); MNHW 978 (2 specimens); PNGNM 004-

105 (2 specimens); UF 299671 (19 specimens); UF

303591 (1 specimen), 30 April 2002; UF 303590 (7 spec-

imens), 2 March 2003, Pini Range, E of Duabo Mission

Station, 10°25.0’ S, 150°18.6’ E, 325 meters altitude, J.

Slapcinsky; UF 299677 (1 specimen), 30 April 2002; UF

303592 (1 specimen), 1 May 2002, Pini Range, aban-

doned logging road W of Duabo Mission Station, 10°

24.9’ S, 150° 18.3’ E, 325 meters altitude, J. Slapcinsky.

Type Locality: Papua New Guinea, Milne Bay Proy-

ince, Pini Range, Duabo Mission Station, 10°25’ 04.7”

S, 150°18’ 24.4” E, 325 meters altitude.

Habitat: Found on logs, mossy rocks and wet ground,

usually near streams in disturbed and undisturbed

broadleaf forest in hilly terrain from 325 to 700 meters

altitude. Observed active during the day.

Etymology: This patronym honors Mr. Biga Lebasi,

our host and guide at Duabo Mission Station, the type

locality.

Remarks: Only four other Paryphantopsis species, P.

elegans, P. fultoni, P. yawii, and P. lamelligera have weak

to strong peripheral keels. Paryphantopsis lebasii is un-

like all species except P. yawii and P. fultoni, in having

large, rectangular periostracal extensions oriented par-

allel to the keeled shell margin. These peripheral exten-

sions are all of equal length, unlike in P. fultoni, and do

not overlap on the body whorl, unlike in P. yawit.

Paryphantopsis matawanensis new species

(Figures 25-31, Table 1)

Description: The adult shell is larger than average

size for the genus, 6.6—7.3 mm (mean = 7.0) in diameter

and 4.0-4.9 mm (mean = 4.5) in height, with 2.7—3.0

(mean = 2.9) rapidly expanding whorls (Figures 25-27,

Table 1). The spire is flat or very slightly elevated, 0.0-

0.1 mm (mean = 0.1). Postnuclear whorls descend reg-

ularly and shell height/diameter ratio is 0.61-0.69 (mean

= 0.64). There are 1.3 rounded nuclear whorls, sculp-

tured with 6 spiral rows of small pits that grade into

weak, spiral striae on the penultimate whorl; shell sculp-

ture is obscured by periostracum on the body whorl. The

shell is wider and slightly angular below the mid- -point.

The postnuclear whorls have regular growth wrinkles ac-

centuated with low periostracal extensions that alternate

with several much longer extensions approximately every

10 growth-lines. The grouping of several longer exten-

sions appears like a single very thick periostracal exten-

sion to the naked eye. The nuclear whorls are white, the

postnuclear whorls are dark brown to red-brown. A re-

flection of the peristome closes the umbilicus. The ap-

erture is large, depressed-ovate, with an aperture-di-

ameter to aperture-height ratio of 0.54-0.67 (mean =

0.63).

In life the body color is bright-yellow with lateral

patches of dark purple- -brown, he yellow fades to cream

in specimens preserved in ethanol. The vas deferens nar-

rows toward the junction with the inflated spherical head

of the epiphallus. Immediately after, and perpendicular

to the head of the epiphallus, there is a finger-shaped

diverticulum that is roughly one quarter the length, and

slightly narrower than the diameter of the epiphallus

(Figure 28). The remainder of the epiphallus is some-

what twisted and approximately the same length as the

penis. The penial retractor muscle is short, originating

from the diaphragm and inserting on the basal third of

the epiphallus. The epiphallus is roughly half the di-

ameter of the penis. The penis expands for its apical

third and then tapers basally to its junction with the sim-

ilarly sized atrium. The penis apex bears several regular

pilasters oriented perpendicular to the length of the pe-

nis (Figure 29). At the penis mid-point there is one very

large and convoluted pilaster that extends basally. The

wall of the base of the penis is thin, bearing regular small

pustules. The atrium is short and narrow ‘expanding

slightly towards the junction with the penis and the long

vagina. The base of the spermathecal duct is relativ ely

narrow, about the same diameter as the base of the penis

and free oviduct; it triples in size to its mid-point then

narrows abruptly for the remaining third before joining

the spherical spermatheca. The free oviduct is narrow,

joining the long vagina well above the atrium.

The central teeth of the radula (center row) are tri-

cuspid, 11-12 zm wide and 18-19 ym long, roughly the

same size and shape as the first lateral teeth (Figure 30).

The mesocones of both the central and first lateral teeth

are tall and sharp, tapering apically and narrowing ba-

sally. Mesocones are attached to their basal plates along

their entire length, except for their apical quarter that

extend beyond the anterior margin of the basal plates.

The ectocones of the central teeth and the symmetric

ectocones and endocones of the lateral teeth are trigo-

nal, about half the height of the mesocones. The mar-

Page 36 THE NAUTILUS, Vol. 119, No. 1

{Gite —

os © an ue oes r a 3 y Stay

Figures 25-31. Paryphantopsis matawanensis. 25-27. Photographs of shell, Holotype UF 308236, diameter 7.3 mm. 28-29.

Camera lucida drawing of genitalia, UF 303581, maximum width 11.7 mm. 30-31. Scanning electron micrograph of radula, UF

303581, field width of central and lateral teeth 63 jm, marginal teeth 59 pm.

J. Slapeinsky, 2005

ginal teeth are dorsoventrally compressed and irregularly

multicuspid, 11-12 jm wide and 10-12 wm long (Figure

31). The ectocones of the marginal teeth are slightly

shorter than their endocones, which are slightly shorter

than their mesocones. Both the ectocones and endo-

cones are irregularly multicuspid, the mesocones are

broadly trigonal to broadly rounded.

Type Material: UF 308236, J. Slapcinsky, 19 Febru-

ary 2003; Paratypes: Papua New Guinea, Milne Bay

Province, Mount Matawan (Mount Simpson): UF

303581 (9 specimens), UF 303582 (3 specimens), type

locality, UF 306529 (1 specimen), NE of summit,

10°2.1’ S, 149°34.4’ E, 2700 m altitude, J. Slapcinsky, 18

February 2003; UF 303583 (1 specimen), NE of sum-

mit, 10°1.7’ S, 149°34.7’ E, 2100 m altitude, J. Slapcin-

sky, 20 February 2003.

Type Locality: Papua New Guinea, Milne Bay Proy-

ince, Mount Matawan (Mount Simpson), ridge top E of

summit, 10°2.5’ S, 149°34.6’ E, 2700 m ailttnadle.

Habitat: All specimens were collected in cloud forest

from 2100 to 2700 m altitude. Individuals were observed

from 1-2 m above ground, crawling on a variety of cloud

forest vegetation in wet or foggy weather. In drier weath-

er specimens were observed in leaf litter that was sus-

pended in trees, especially in the crowns of Pandanus

sp.

Etymology: Named for the type locality and known

range of this species: Mount Matawan (Mount Simpson).

Remarks: Of the other larger (>6 mm diameter) spe-

cies of Paryphantopsis, P. globosa, and P. louisiadarum

do not have periostracal extensions on the growth lines,

unlike P matawanensis. Of the species with periostracal

extensions, P. latior, P. lamelligera, P. fultoni, and P. stri-

ata do not have a repeating pattern of approximately 10

short periostracal extensions followed by several longer

extensions. P. matawanensis further differs from P. la-

melligera and P. fultoni in not having peripheral pro-

cesses on the periostracal extensions. The genital anat-

omy of P matawanensis differs from P. lamelligera in

having a diverticulum.

Paryphantopsis ubwamensis new species

(Figures 32-38, Table 1)

Description: The adult shell is large for the genus,

6.4—7.7 mm (mean = 7.1) in diameter and 4.8—5.1 mm

(mean = 5.0) in height, with 2.9-3.1 (mean = 3.0), rap-

idly expanding whorls (Figures 32-34, Table 1). The

spire is elevated, 0.2-0.4 mm (mean = 0.3), the post-

nuclear whorls descend relatively rapidly, especially near

the aperture. The shell height/diameter ratio is 0.66—

0.75 (mean = 0.70). The 1.2 nuclear whorls are evenly

rounded and sculptured with 12 spiral rows of small pits.

These pits do not continue on the postnuclear whorls,

which are sculptured only with weak growth lines that

do not bear periostracal extensions. The whorls are in-

flated, the sutures deeply impressed, and the periphery

evenly rounded. The nuclear whorls are white, the post-

nuclear whorls brown, with irregular lighter patches.

The umbilicus is perforate, narrowed by a reflection of

the peristome. The aperture is large, ovate, with an ap-

erture-diameter to aperture-height ratio of 0.79-0.93

(mean = 0.85).

The body color is uniform yellow in life, fading to

cream in specimens preserved in ethanol. The vas def-

erens is wide at the prostate gland, narrowing rapidly

and remaining narrow until the junction with the inflated

ovate tip of the epiphallus (Figure 35). The epiphallus

bears a long (approximately a quarter of the length of

the epiphallus), finger-shaped diverticulum just after the

junction with the vas deferens. The epiphallus is about

one third the diameter of the apex of the penis. The

penis is broad apically, narrowing abruptly to half its api-

cal diameter slightly before mid-point and remaining the

same diameter to the junction with the atrium. The pe-

nis is sculptur ed with several slightly convoluted pilasters

that extend in an arc from near basally to near apically,

and one much larger and more convoluted pilaster near

the penis mid-point (Figure 36). The penial retractor

muscle originates from the diaphragm and inserts at the

mid-point of the epiphallus. The spermathecal duct is

basally robust and narrows at mid-point, remaining nar-

row until the junction with the spherical spermatheca.

The free oviduct is narrow joining the moderate length

vagina above the atrium.

The central teeth of the radula (center row) are tri-

cuspid, 10-11 jm wide and 15-16 jm long, roughly the

same width and shape as, but a little shorter than, the

first lateral teeth, which are 18-19 wm long (Figure 37).

The mesocones of both the central and lateral teeth are

long, slender and blade shaped, projecting slightly be-

yond the basal plate. The ectocones and endocones of

the lateral teeth are symmetrical and half the height of

the mesocones. The marginal teeth are dorsoventrally

compressed and irregularly tricuspid-multicuspid, 12-15

yum wide and 13-15 ym long (Figure 38). The endo-

cones are tall, large to very large and sometimes irreg-

ularly multicuspid. The ectocones are unicuspid.

Type Material: Holotype: UF 303589, J. Slapcinsky,

22 April 2002; Paratypes: Papua New Guinea, Milne Bay

Province, Cloudy Mountains, Ubwam Mountain: UF

299666 (1 specimen), type locality; UF 299668 (2 spec-

imens), headwaters of the Watuti River, 10°29.8’ S,

150°14.02’ E, 670 m altitude, J. Slapcinsky, 16 April

2002.

Type Locality: Papua New Guinea, Milne Bay Prov-

ince, Cloudy Mountains, Ubwam Mountain, summit,

10°30.4’ S, 150°13.5’ E, 1000 m altitude.

Habitat: This species was found crawling on a moss-

covered rotting log in cloud forest at 1000 meters. Dead

Jnells were alli Found at G70 ameiters fn Tee lhigier att Une

base of a tree with smooth bark.

Page 38 THE NAUTILUS, Vol. 119, No. 1

Figures 32-38. Paryphantopsis ubwamensis. 32-34. Photographs of shell, Holotype UF 303589, diameter 7.7 mm. 35-36.

Camera lucida drawing of genitalia, UF 299666, maximum width 10.1 mm. 37-38. Scanning electron micrograph of radula, UF

299666, field width of ‘central and lateral teeth 66 2m, marginal teeth 45 pm.

J. Slapcinsky, 2005

Page 39

Etymology: This species is named for the type local-

ity, Ubwam Mountain, and indirectly for one of our

guides who is named after the mountain.

Type Material: Holotype: Papua New Guinea, Milne

Bay Province, Cloudy Mountains, Ubwam Mountain,

summit, 10°30.4’ S, 150°13.5’ E, 1000 m altitude, J.

Slapcinsky, 22 April 2002 (UF 303589); Paratypes: Papua

New Guinea, Milne Bay Province, Cloudy Mountains,

Ubwam Mountain: type locality (UF 299666, 1 speci-

men); headwaters of the Watuti River, 10°29.8’ S,

150°14.02’ E, 670 m altitude, J. Slapcinsky, 16 April

2002 (UF 299668, 2 specimens).

Paryphantopsis yawii new species

(Figures 39-45, Table 1)

Description: The adult shell is average in size for the

genus, 5.0-6.5 mm (mean = 5.9) diameter, and 2.6-3.6

mm (mean = 3.1) height with 2.6-2.8 (mean = 2.7)

rapidly expanding whorls (Figures 39-41, Table 1). The

spire is elevated, 0.1-0.3 mm (mean = 0.2). Postnuclear

whorls descend slowly and regularly. Shell height/diam-

eter ratio is 0.47-0.55 (mean = 0.53). The 1.2 nuclear

whorls bear rounded peripheral and supraperipheral

ridges; sculptured with about 15 spiral rows of small pits.

These pits become larger and less regular on postnuclear

whorls but usually are obscured by the periostracum.

However, pits are visible in areas where the periostra-

cum is removed, and within the aperture. The shell has

a peripheral keel and blunt supraperipheral ridge; the

surface between is flattened. Postnuclear whorls are

sculptured with regular growth wrinkles that are accen-

tuated with periostracal extensions. On the apical surface

of the shell, periostracal extensions are folded along their

length toward the aperture and appressed to the shell

except at the keeled margin where they form rectangular

processes that extend about 0.5 mm beyond the shell

margin. The large, distally rounded processes overlap,

forming a continuous periostracal fringe of uniform

length at the shell periphery. Periostracal extensions on

the growth lines extend basally but are short and erect.

Nuclear whorls are white, postnuclear whorls yellow

brown. The umbilicus is closed by a reflection of the

peristome. The aperture is large, depressed-ovate, with

an aperture-diameter/aperture-height ratio 0.53—0.66

(mean = 0.63).

The body color is uniform bright creamy-yellow in

life, fading to creamy-white in specimens preserved in

ethanol. The vas deferens narrows toward the junction

with the slightly inflated head of the epiphallus (Figure

42). The epiphallus is approximately three times longer

and one quarter of the diameter of the penis and does

not bear a diverticulum. The penial retractor muscle is

long, about two thirds the length of the epiphallus, orig-

inating from the diaphragm and inserting on the basal

third of the epiphallus. The penis is short and robust

with poorly defined pilasters in the apex (Figure 43).

The atrium is short and narrow, expanding slightly to-

wards the junction with the penis and the short vagina.

The base of the spermathecal duct is massive, tapering

slowly but remaining broad for more than one third of

its length; the remainder is relatively narrow until the

junction with the spherical spermatheca. The free ovi-

duct is slightly coiled and narrow, joining the short va-

gina just above the atrium.

The central teeth of the radula (fifth row from left)

are tricuspid, 9-10 wm wide and 14-15 ym long, rough-

ly the same size and shape as the first lateral teeth (Fig-

ure 44). The mesocones of both the central and first

lateral teeth are short, conical and erect, joining the rect-

angular basal plates nearly centrally and barely project-

ing beyond the plates’ anterior margin. The ectocones

are triagonal and short, only one third of the height of

the mesocones, joining the posterior edge of the basal

plates. The lateral teeth are asymmetrical; their endo-

cones are slightly taller then their ectocones. The mar-

ginal teeth are dorsoventrally compressed, tricuspid to

multicuspid, 8-10 wm wide and 10-12 wm long (Figure

45). The endocones of the marginal teeth are nearly the

same height as the mesocones while the ectocones are

much shorter, only one half to one third of their height

and often divided into three cusps.

Type Material: Holotype: UF 308238, J. Slapcinsky,

6 April 2002; Paratypes: Papua New Guinea, Milne Bay

Province: UF 299675 (2 specimens), UF 303594 (2 spec-

imens), Wowow Mountain, W of Naura, 10°16.9’ S,

150°9.9' E, 635 m altitude, J. Slapcinsky, 8 May 2002;

UF 299673 (1 specimen), UF 303595 (1 specimen), 4

April 2002, UF 303598 (1 specimen), 27 February 2003,

waterfall on Upalai Creek, 3 km WNW of Watunoa,

10°19.6’ S, 150°34.6’ E, 60 m altitude, J. Slapcinsky; UF

299670 (3 specimens), UF 299672 (6 specimens), 6 April

2002, UF 303596 (13 specimens), 7 March 2003, head-

water of Goilayoli River at road crossing, 30 km ENE

of Alotau, 10°18.7’ S, 150°37.3’ E, 275 m altitude, J.

Slapcinsky; BPBM (4 specimens), MNHW 977 (4 spec-

imens), PNGNM 004-104 (4 specimens), UF 303597 (45

specimens), small waterfall on Kinahidamadamana River

near Budo Village, 10°17.1’ S, 150°26.7’ E, 125 m alti-

tude, J. Slapcinsky, 4 March 2003.

Type Locality: Papua New Guinea, Milne Bay Prov-

ince, headwater of Goilayoli River at road crossing, 30

km ENE of Alotau, 10°18.7’ S, 150°37.3’ E, 275 m al-

titude.

Habitat: This species was active during the day in na-

tive forest on rocks and logs with moss and algae, and

on moist soil near streams. It was encountered in hilly

terrain at relatively low altitudes for the genus, ranging

from 60 to 635 meters.

Etymology: This patronym honors Mr. Benjamin Yawi

of Budo Village, Milne Bay Province, Papua New

Guinea. He and his family located, arranged permission

to visit, and helped to collect at many of the sites where

this species was found.

Page 40 THE NAUTILUS, Vol. 119, No. 1

lucida drawing of genitalia, UF 299672, maximum width 6.4 mm. 44-45. Scanning electron micrograph of radula, UF 299672,

field width of central and lateral teeth 63 zm, marginal teeth 48 ym.

J. Slapcinsky, 2005

Page 41

Remarks: Peripheral keels are unusual among known

Paryphantopsis species and are found only in P. elegans,

P. fultoni, and to a lesser extent in P. lebasii and P. la-

melligera. Paryphantopsis yawii is unlike all other spe-

cies, except for P. lebasii and P. fultoni, in having large,

rectangular periostracal extensions oriented parallel to

the keeled shell margin. These peripheral extensions are

all of equal length, unlike in P fultoni and they overlap,

forming a continuous periostracal fringe, unlike in P. le-

basii.

DISCUSSION AND CONCLUSIONS

The family Charopidae was previously considered to be

a minor component of the terrestrial molluscan fauna of

New Guinea, with relatively few species and genera, and

to lack the spectacular radiations exhibited By this and

the related Endodontidae in the oceanic islands of the

Pacific (Solem, 1983: 305). Ongoing surveys indicate that

this is not the case; inadequate : sampling, rather than low

diversity, is the cause of the perceived low number of

charopid species in New Guinea. Likewise, reports of

low generic diversity of charopids in New Guinea are

more likely the result of insufficient sampling and may

reflect the paucity of anatomic material available to de-

fine generic units (Solem, 1970: 241). Despite the short

duration and limited geographic scope of the current

survey, six new species of Paryphantopsis are reported

here, increasing the known diversity of the genus by al-

most 50%. In addition to Paryphantopsis, species be-

longing to several other charopid genera were also col-

lected; these will be treated in later publications.

On the eastern terminus of the Papuan Peninsula,

Paryphantopsis species that occur in close proximity or

sympatry share unique shell, genital, and radular char-

acters. For example, both species from the Mount Ma-

tawan area, P. matawanensis and P. koragae, have un-

usually short penial retractor muscles, long vaginas and

angled apertural margins. Species in the uplands of the

Cloudy Mountains, P ubwamensis and P. abstrusa, are

unusually tightly coiled and globose. The two lowland

species, P. lebasii and P. yawii, share distally rounded

rectangular periostracal processes and the unusual origin

of their mesocones from the center of their basal plates.

These unusual characters shared by different species in

close proximity or sympatry suggest that Paryphantopsis

species have speciated locally on a fine geographic scale;

speciation in these cases is presumably facilitated by

their poor dispersal ability. Because much of New

Guinea remains under-explored, the true diversity of the

islands’ Paryphantopsis, other charopids, and land snails

in general is almost certainly greatly underestimated.

The geographic distribution of the six Paryphantopsis

species appears to be limited to particular mountain

ranges on different terrains, despite the proximity of

these mountains to each other (Figure 1). Three species,

P. matawanensis, P. koragae, and P. yawii are restricted

to the Owen Stanley Range, part of the Kutu Terrane,

while three others, P abstrusa, P. lebasii, and P. ubwa-

mensis, are restricted to the Cloudy Mountains of the

Port Moresby Terrane. These distinct suites of endemic

species are consistent with the terrane-accretion hypoth-

esis (Davis et al. 1997) proposed for the formation of

the East Papua Composite Terrane and suggest that the

low vagility and high diversity of charopids and other

land snails may make them ideal to test hypotheses of

terrain accretional history.

ACKNOWLEDGMENTS

I thank the landowners of Alotau, Budo, Bunisi, Gadow-

alai, Ikara, and Naura for permission to work on their

land and for field assistance; J. Anamiato, I. Bigilale, F.

Kraus, F. Malesa, B. Uruwa, and B. Yawi for additional

field assistance; B. Lebasi for hosting my stay at Duabo

Mission Station; B. Yawi for help accessing land through-

out the Alotau area; G. Kula and D.Mitchell of Conser-

vation International for providing logistical support and

advice; PNG National Museum and Ae Gallery for pro-

viding in-country collaborative assistance; PNG Depart-

ment of Environment and Conservation, PNG National

Research Institute, and Milne Bay Provincial Govern-

ment for permission to work in Milne Bay Province; and

J. Worsfold for sharing bibliographic information. Field

work for this research was supported by National Sci-

ence Foundation grant DEB 0103794 and the University

of Florida Foundation, McGinty Endowment. K. Em-

berton, F. Kraus, G. Paulay, F. G. Thompson, and two

anonymous reviewers suggested improvements to earlier

drafts of this manuscript.

LITERATURE CITED

Bavay, A. 1908. Mollusques terrestres et fluviatiles. Nova

Guinea, Zoology 5: 269-292, pl. 14.

Coen, G. S. 1922. Descrizione di nuovo specie di molluschi

del Museo Civico di Genova. Annali del Museo Civico di

Storia Naturale, Genova 9(3): 359-363.

Fulton, H. C. 1902. Descriptions of new species of land Mol-

lusca from New Guinea. Annals and Magazine of Natural

History 7(9): 182-184.

Hedley, C. 1890. Description of a new Rhytida from New

Guinea. Annual Report of British New Guinea, 1888—

1889: 94.

Jutting, W. S. S. v. B. 1964. Non-marine Mollusca of West New

Guinea. Part 3, Pulmonata, I. Nova Guinea, Zoology 26:

-74, pls. 1-2.

Mollendorff, O. 1899. Neue arten aus der Strubell’schen sa-

mmlung. Nachrichtsblatt der Deutschen Malakozoolo-

gischen Gesellschaft 31(5): 89-92.

Pigram, C. J. and H. L. Davies. 1987. Terranes and the accre-

tion history of the New Guinea orogen. BMR Journal of

Australian Geology and Geophysics 10: 193-211.

Solem, A. G. 1970. The Endodontid land snail genera Pilsbry-

charopa and Paryphantopsis (Mollusca: Pulmonata). The

Veliger 12: 239-264.

Solem, A. G. 1983. Endodontoid land snails from Pacific Is-

lands (Mollusca: Pulmonata: Sigmurethra). Part II. Fam-

ilies Punctidae and Charopidae, Zoogeography. Field Mu-

seum of Natural History, Chicago ix + 336 p.

Page 42

Thiele, J. 1928. Mollusken vom Bismark-Archipel, von Neu-

Guinea und Nachbar-Inseln. Zoologische Jahrbiicher 55:

119-146.

Wiktor, A. 2003. Terrestrial gastropods (Mollusca) of province

THE NAUTILUS, Vol. 119, No. 1

Madang in Papua-New Guinea. Part III. Pulmonata:

Rathousiidae, Ellobiidae, Succineidae, Agriolimacidae,

Endodontidae (partim), Ariophantidae, Euconulidae, Sub-

ulinidae, Streptaxidae. Folia Malacologica 11(1/2): 1-21.

THE NAUTILUS 119(1):43-49, 2005 Page 43

First record of Akera Miiller, 1776, from the eastern Pacific,

with the description of a new species

Kelvin Barwick

City of San Diego

EMTS Laboratory

2392 Kincaid Road

Angel Valdés

Natural History Museum of Los

Angeles County

900 Exposition Boulevard

Los Angeles, CA 90007 USA

[email protected]

San Diego, CA 92101 USA

kbarwick@sandiego. gov

ABSTRACT

The description of the new species Akera julieae is based on a

complete specimen collected from southern California and

three empty shells from Costa Rica. These specimens consti-

tute the first record of Akera in the eastern Pacific. Akera ju-

lieae is distinguishable from the widespread Indo-Pacific spe-

cies Akera soluta by its radular and jaw morphology; the rach-

idian teeth of A. soluta are more solid with larger cusps and

flanking denticles, the mid-lateral teeth of A. soluta are den-

ticulated, whereas in A. julieae they are smooth, and the out-

ermost lateral teeth of A. julieae are proportionally more elon-

gate and straighter than those of A. soluta; the jaws of A. soluta

are well developed and composed of a number of rows of

strong rodlets, whereas in A. julieae the jaws are more rudi-

mentary with only five rows of fragile-looking rodlets. There

are no consistent differences between these two species in giz-

zard plate and adult shell morphology, but the protoconch of

A. soluta is slightly larger and more elongate. The western At-

lantic species Akera bayeri is distinguishable from A. julieae by

having a more elevated shell spire and stronger radular teeth

with denticles on all lateral teeth.

INTRODUCTION

The opisthobranch family Akeridae Mazzarelli, 1891,

includes opisthobranchs with an external, cylindrical

shell into which the animal cannot retract completely. It

contains the single genus Akera Miiller, 1776, and only

a few Recent valid species distributed throughout trop-

ical and temperate regions. The type species, Akera bul-

lata Miiller, 1776, hae | been reported from the northeast

Atlantic, from dhe Baltic shores of Denmark and Norway

to the British Isles, continuing on to the Atlantic and

Mediterranean costs of continental France and Spain

(Thompson, 1976), Italy (Rinaldi, 1988), and the Canary

Islands (Ortea et al., 2001).

In the Indo-Pacific, the widespread species Akera so-

luta (Gmelin, 1791) has been reported from South Af-

rica and Mozambique to Australia, Marshall Islands, and

the Philippines (Gosliner, 1987). Both Akera bicincta

(Quoy and Gaimard, 1833), from Australia, and Akera

constricta Kuroda 1947, from Japan, have been regarded

as junior synonyms of A. soluta, see Wells and Bryce

(1993) and Hamatani (2000) respectively. This expands

the known range for A. soluta. Other nominal Indo-Pa-

cific species, Akera tumida (A. Adams in Sowerby, 1850),

Akera tasmanica Beddome, 1882, and Akera aperta

Hedley, 1899, were all described from the southwestern

Pacific, and are poorly known. They are likely synonyms

of A. soluta, but because they were described based sole-

ly on shell morphology, their identities are unclear and

in need of revision. In the western Atlantic, Olsson and

McGinty (1951) reported for the first time a species of

Akera from Florida under the name “Akera thompsoni.”

Because the animal was figured but not described, this

species name is a nomen pain in accordance with Ar-

ticle 13.1 of the International Code of Zoological No-

menclature (ICZN, 1999). The first available name for

the western Atlantic Akera is Akera bayeri Ey. Marcus

and Er. Marcus, 1967, which has been reported from

the southwestern Caribbean Sea and Brazil (Ev. Marcus,

1970).

There are no Recent species of Akera known from

the eastern Pacific. Akera maga Vokes, 1939, the only

known species from this area, was described from the

lower to middle Eocene, Domengine Formation, Fresno

County, California (Vokes, 1939). Squires (2001) report-

ed this species from the Llajas Formation (also lower to

middle Eocene), Ventura County, California. In the

present paper we describe the first occurrence of Recent

Akera in the eastern Pacific based on a complete spec-

imen collected in the Channel Islands, California and

three shells from Guanacaste, Costa Rica.

MATERIALS AND METHODS

The specimen from California was collected as part of

the Southem California Bight 1998 Regional Marine

Monitoring Survey (Bight, 1998) eondmeted | in the sum-

mer of 1998. Infaunal samples were taken with a 0.1 m?

Van Veen grab sampler and screened through a 1 mm

mesh. The sample was then placed in a relaxant solution

of Epsom salts (magnesium sulfate heptahydrate—

Page 44

THE NAUTILUS, Vol. 119, No. 1

Table 1. Comparative material examined in this study. The specimen marked with an asterisk (*) was collected alive and included

soft parts.

Species Locality

Akera soluta Kii, Japan

Uala Reef, Quezon, Philippines

New Zealand

Hardwick Bay, South Australia

Phuket, Thailand

Akera bullata Finmark, Norway

Date Depth Number

— LACM 153414%

May 1959 3-15 m LACM 073035

LACM 153415

LACM 157942

LACM 153416

LACM 153417

LACM 046386

LACM 153418

MgSO,-7H,O) and freshwater for a minimum of 30 min-

utes. The sample was then fixed in 10% buffered for-

malin and preserved in 70% ethanol. The preserved

whole animal was photographed using a digital camera

mounted on a dissecting scope (Wild Epimarkroskop®

M450).

The Costa Rican shells were collected during the

Searcher 401 Expedition of the Natural History Muse-

um of Los Angeles County.

Several additional specimens and shells belonging to

other species of Akera were examined for comparison

purposes. These specimens are listed in Table 1.

The shell is very fragile and could not be dissected

intact from the specimens examined, which rendered

the description below incomplete. Once the shell was

removed, the internal organs were dissected and drawn

using a Nikon SMZ 1000 microscope equipped with a

drawing tube. The radula, jaws, and gizzard plates were

dissected and photographed using a Hitachi S-3000N

Scanning Electron Microscope (SEM). The apical region

of the shell was separated from the rest of the shell and

mounted for SEM photography.

SYSTEMATICS

Akeridae Mazzarelli, 1891

Akera Miiller, 1776

Type Species: Akera bullata Miiller, 1776, by mono-

typy.

Diagnosis: Shell external, fragile, translucent, cylin-

drical to slightly bulloid. Spire flattened to elevated with

a partially embedded protoconch. Aperture equal to the

length of the spire or slightly shorter. Thin periostracum

forming a raised flange at the keel. Animal unable to

retract fully into the shell, but can stretch over twice its

length; with a posterior pallial tentacle. Parapodia ex-

tending laterally, meeting mid-dorsally over the shell.

Radula with rachidian tooth having a median cusp and

smaller denticles on either side. Rachidian tooth flanked

by 21-52 lateral teeth. Gizzard with a number of irreg-

ular plates arranged in three tiers.

Akera julieae new species

(Figures 1-4)

Akera sp.—Behrens, 2004: 18, pl. 1D.

Holotype: LACM 3033, from type locality.

Paratypes: 3 shells, southeastern corner of Bahia

Jobo, off sand beach west of Bahia de Salinas, Guana-

caste Province, Costa Rica (11°02’22” N, 85°45'16” W),

14 Feb. 1972, 1.5-10.7 m depth (LACM 3034).

Type Locality: Southwestern corner of Santa Catalina

Island, California, USA (33°18’24" N, 118°22’05” W), 24

Jul. 1998, 40.6 m depth, in gray colored silt and clay

(LACM 3033).

External Morphology: The body is oval, 10 mm long

in the preserved holotype. The cephalic shield is trian-

gular, comprising about 4 of the body length in the pre-

served specimen (Figures 1A, 2A). The parapodia are

narrow and do not reach the midline of the body (Fig-

ures 1A—-C, 2A). The gill is unipinnate, with 11 simple

lamellae (Figure 2B). The color of the living animals is

unknown; the preserved holotype is grayish white.

Shell Morphology: The shell is bullomorph, fragile,

well-calcified (Figure 1). The protoconch is smooth, and

only the outer whorl is visible externally (Figure 3C). Its

maximum diameter is 200 um. The whorls are separated

by a deep, channelled suture, which is shallower on the

apical whorls. The periphery of the whorls, near the

apex, is angulated and forms a conspicuous keel, which

divides the whorls into two parts (Figure 1D, 3C). The

inner part has a characteristic pattern of strong, curved

wrinkles. The whorls are attached to the preceding

whorl just below the periphery of this whorl. The apical

region is flattened and the external whorls overlap the

most internal. The whorl sides are clearly curved, con-

vex. The aperture is broad below and narrow above,

where it extends into a deep sinus along the suture of

the upper lip. There is a thin callus in the columella.

The sculpture consists of numerous, low and thin spiral

ribs and axial growth lines. The shells are covered with

a thin brownish periostracum.

Anatomy: The buccal bulb is oval; it connects poste-

riorly to the long esophagus and the salivary glands (Fig-

A. Valdés and K. Barwick, 2005

Page 45

Figure 1. Akera julieae new species, photographs of the preserved holotype (LACM 3033). A. Dorsal view. Scale bar = 5 mm.

B. Ventral view. Scale bar as in A. C. Lateral view. Scale bar = 5 mm. D. Apical view of the shell. Scale bar = 5 mm.

ure 2C). Two strong retractor muscles attach laterally to

the buccal bulb. The radular formula is 19X23.1.23 in

the holotype. The rachidian teeth are broad, with a tri-

angular based, basally concave, and a pointed central

cusp (Figure 4A). There are 3-5 denticles on each side

of the cusp varying in shape and size. The two innermost

teeth on each row have a long and narrow base and a

conspicuously wider cusp bearing denticles on the inner

Page 46

THE NAUTILUS, Vol. 119, No. 1

Figure 2. Akera julieae new species, drawings of the preserved holotype (LACM 3033). A. Dorsal view of the complete body.

Scale bar = 1 mm. B. Detail of the gill. Scale bar = 1 mm. C. General view of the anatomy. Scale bar = 1 mm. D. General view

of the reproductive system. Scale bar = 0.5 mm. E. Penis and prosate. Scale bar = 0.5 mm. Abbreviations: ag, albumen gland:

am, ampulla; bb, buccal bulb; be, bursa copulatrix; cs, cephalic shield; dg, digestive gland; es, esophagus; esg, external seminal

groove; feo, female copulatory organ; ga, genital atrium; gl, gill; gz, gizzard; i, intestine; mg, mucous gland; pn, penis; pr, prostate;

ps, penial sheath; rm, retractor muscle; sh, shell; sg, salivary gland; sr, seminal receptacle.

and outer sides. The rest of the lateral teeth are hook-

shaped, with a long and narrow cusp and lack denticles

(Figure 4B). In the outermost teeth the base is shorter

and the cusp proportionally longer than in the mid-lat-

erals (Figure 4C). The jaws are composed of 6 rows of

simple, elongate rodlets (Figure 3B). The esophagus

opens into a large muscular gizzard, which contains sev-

eral gizzard plates. The gizzard plates vary in shape and

size; they are irregular with angular edges (Figure 3A).

The reproductive system is monoaulic (Figure 2D).

The ampulla is long and convoluted; it opens into the

genital atrium at the same point where the albumen and

mucous glands open. The bursa copulatrix is oval; it con-

nects with the genital atrium through a wide and curved

duct. The genital atrium is long and is connected to a

complex female copulatory organ near the opening.

From the gonopore an open seminal groove runs in an-

terior direction to the protrusible cephalic penis and the

prostate. The penis is long and externally enclosed in a

sheath (Figure 2E). The prostate is short and simple,

and connects proximally to the penis.

Etymology: Dedicated to Julie Barwick, the daughter

of the junior author.

DISCUSSION

Akera julieae has been included in Akera because of the

presence of a fragile, cylindrical external shell, a flat-

tened spire, and a partially embedded protoconch. The

radula of this species has a single broad, triangular rach-

idian tooth, with a median cusp and smaller denticles on

either side, as well as several hamate lateral teeth. The

gizzard contains a number of irregular gizzard plates ar-

ranged in three tiers. All these characteristics are diag-

nostic of the genus Akera (see above).

This is the first record of Akera in the eastern Pacific.

Behrens (2004) cited this species as Akera sp., based on

, oe i 3 Y tila

Page 47

Figure 3. Akera juliae new species and A. soluta (Gmelin, 1791). A-C. Akera julieae, scanning electron micrographs of the

preserved holotype (LACM 3033). A. Gizzard plate. Scale bar = 300 wm. B. Jaw. Scale bar = 50 ym. C. Protoconch. Scale bar

= 200 pm. D-E. Akera soluta (Gmelin, 1791), scar

ng electron micrographs of a specimen from Japan (LACM 153414). D.

Gizzard plate. Scale bar = 300 pm. E. Jaw. Scale bar = 50 wm. F. Protoconch. Scale bar = 200 pm.

the material here examined and information, photo-

graphs, and descriptions provided by the junior author.

The description of the new species is mainly based on

the holotype, collected from California, but the three

empty shells collected from Costa Rica seem to belong

to the same species. However, this needs to be verified

when complete specimens from Costa Rica become

available.

Akera julieae differs from other described species of

the genus. Several shells and one specimen of the Indo-

Pacific Akera soluta were examined for comparison (see

Table 1) confirming the presence of several external and

internal differences between these two species. For in-

stance, the rachidian radular teeth of A. soluta are more

solid with larger cusps and flanking denticles (Figure

4D). The three innermost lateral teeth have wide cusps

. 119, No. 1

Figure 4. Akera juliae new species and A. soluta (Gmelin, 1791). A-C. Akera julieae, scanning electron micrographs of the radula

of the holotype (LACM 3033). A. Rachidian and innermost lateral teeth. Scale bar = 50 pm. B. Mid-lateral teeth. Scale bar = 50

wm. C. Outermost lateral teeth. Scale bar = 50 ym. D-F. Akera soluta, scanning electron micrographs of the radula of a specimen

from Japan (LACM 153414). D. Rachidian and innermost lateral teeth. Scale bar = 50 um. E. Mid-lateral teeth. Scale bar = 50

ym. F. Outermost lateral teeth. Scale bar = 50 pm.

with denticles in A. soluta, whereas only the two inner-

most teeth of A. julieae have similar characteristics. The

mid-lateral teeth in A. soluta also have denticles (Figure

4E), whereas they are smooth in A. julieae. The outer-

most lateral teeth of A. julieae are proportionally more

elongate and straighter than those of A. soluta (Figure

4F). More importantly, the jaws of A. soluta are ‘well

developed and composed of a number of rows of strong

rodlets (Figure 3E), whereas in A. julieae the jaws are

more rudimentary with only 5 rows of fragile-looking

rodlets. There are no consistent differences maneen ihe

gizzard plates of A. julieae and A. soluta (Figures 3A,

3D). Externally, A. soluta has a slightly larger ead more

elongate protoconch. The adult shells of hase two spe-

cies are indistinguishable due to the morphological var-

iability in A. soluta.

Akera bayeri is the only other species of Akera known

from the Americas. The external morphology and anat-

A. Valdés and K. Barwick, 2005

Page 49

omy of this species was described in detail by Marcus

and Marcus (1967) and Marcus (1970). Akera bayeri dif-

fers from A. julieae in two important regards. The shell

of A. bayeri has a more elevated spire and the radular

teeth are stronger with denticles on all lateral teeth,

whereas in A. julieae the lateral teeth are smooth.

LITERATURE CITED

Behrens, D. W. 2004. Pacific coast nudibranchs, Supplement

II—New species to the Pacific coast and new information

on the oldies. Proceedings of the California Academy of

Sciences 55: 11-54.

Gosliner, T. 1987. Nudibranchs of Southem Africa. Sea Chal-

lengers, Monterey, 136 pp.

Hamatani, I. 2000. Family Akeridae. In: Okutani, T. (ed.), Ma-

rine Mollusks in Japan. Toaki University Press, Tokyo, xlvii

+ 1173 pp.

ICZN (International Commission of Zoological Nomenclature).

1999. International Code of Zoological Nomenclature, 4"

Edition. International Trust on Zoological Nomenclature,

London, 306 pp.

Marcus, Er. and Ev. Marcus. 1967. Opistobranchs from the

southwestern Caribbean Sea. Bulletin of Marine Science

17: 597-628.

Marcus, Ev. 1970. Opisthobranchs from northern Brazil. Bul-

letin of Marine Science 20: 922-995.

Olsson, T. and L. McGinty. 1951. “Akera thompsoni.” The

Nautilus 65: pl. 3, figs. 7-7b.

Ortea, J. A., L. Moro, J. J. Bacallado and R. Herrera. 2001.

Catélogo actualizado de los Moluscos Opisthobranquios

de las Islas Canarias. Revista de la Academia Canaria de

Ciencias 12: 105-134, pls. 14.

Rinaldi, E. 1988. Primi rinvenimenti di Akera bullata O. F.

Mueller, 1776 sulla costa Romagnola. Bollettino Malacol-

ogico 24: 25-26.

Squires, R. L. 2001. Additions to the Eocene megafossil fauna

of the Las Llajas Formation, Simi Valley, Southern Cali-

fornia. Contributions in Science 489: 1-40.

Thompson, T. E. 1976. Biology of Opisthobranch Molluscs,

Volume 1. The Ray Society, London, 207 pp.

Vokes, H. E. 1939. Molluscan faunas of the Domengine and

Arroyo Hondo formations of the California Eocene. An-

nals of the New York Academy of Sciences 38: 1-246, pls

1-22.

Wells, F. E. and C. W. Bryce. 1993. Sea Slugs of Western

Australia. Western Australian Museum, 184 pp.

THE NAUTILUS 119(1):50-54, 2005

Page 50

Description of Calliotropis pulvinaris new species (Gastropoda:

Trochidae: Eucyclinae: Calliotropini) from West Madagascar

Claude Vilvens

Rue de Hermalle, 113

B-4680 Oupeye

BELGIUM

[email protected]

ABSTRACT

Calliotropis pulvinaris new species is described from West

Madagascar and compared with similar species in the trochid

subfamily Eucyclinae, particularly with C. patula (Martens,

1904), C. concavospira (Schepman, 1908), C. blacki Marshall,

1979, and C. vaillanti (Fischer, 1882). The new species can be

separated from these by a rather depressed spire, a rounded

periphery, tumid whorls beari ing four spiral cords of which nod-

ules decrease in size and increase in number from adapical

cord to abapical cord, and five spiral cords on the base.

INTRODUCTION

The malacofauna of this area remains poorly known, de-

spite earlier surveys (1971-1973) by ORSTOM (Office

de la Recherche Scientifique et Technique Outre-Mer,

now IRD: Institut de Recherche pour le Développe-

ment) on the continental slope of Madagascar (Crosnier

and Jouannie, 1973). Independently of the inherent eco-

nomic interest, this endeavor yielded abundant zoologi-

cal material, more particularly mollusks now deposited

at the MNHN (Muséum national d’Histoire naturelle,

Paris).

Commercial fishing boats have trawled for deep-water

shrimp off Madagascar. The commercial dredging off

West Madagascar “Exoran (hese lest years brought various

specimens of trochid species, some of etn described

in the past (Watson, 1886; Martens and Thiele, 1904;

Thiele, 1925; Barnard, 1963), others recently named as

new species (Vilvens, 2001 and 2002). Two years ago,

Guido T. Poppe entrusted me with trochid shells col-

lected in deep water. These shells, originally labeled as

Calliotropis patula (Martens, 1904), are conspecific with

material trawled by French expeditions and deposited at

MNHN in the 1970s. Closer examination and compari-

son with the type of the supposed species leads me to

conclude that all these shells belong to an unnamed spe-

cies that is described here as new.

Text abbreviations used are: IRSNB: Institut royal des

Sciences naturelles de Belgique, Bruxelles, Belgium;

MNHN: Muséum national d’Histoire naturelle, Paris,

France; NMNZ: Museum of New Zealand Te Papa Ton-

garewa, Wellington, New Zealand; ZMA: Zodlogisch

Museum, Amsterdam, The Netherlands; ZMB: Zoolo-

gisches Museum of Berlin, Germany; P1, P2, P3, P4:

primary cords (P1 is the most adapical); stn: station; dd:

no live specimens present in sample.

I follow below the classification of Hickman and Mc-

Lean (Hickman and McLean, 1990) at the suprageneric

level.

SYSTEMATICS

Family Trochidae Rafinesque, 1815

Subfamily Eucyclinae Koken, 1897

Tribe Calliotropini Hickman and McLean, 1990

Genus Calliotropis Seguenza, 1903

Type Species: Trochus ottoi Philippi, 1844, Pliocene—

Pleistocene, Italy; by original designation.

Calliotropis pulvinaris new species

(Figures 1-6)

Description: Shell rather long for genus (height up to

18.3 mm, width up to 29.0 mm), rather depressed, rath-

er thin, cyrtoconoidal; spire rather low, height 0.6—

0.7Xwidth, height 2.3-4.5Xaperture height; umbilicus

deep and large. Protoconch about 300 zm wide, with

about | whorl, partially damaged on available specimens,

remaining part smooth. Teleoconch of up to seven con-

vex whorls! bearing four spiral granular cords and pro-

socline threads; nodules from cords produced by inter-

sections with axial folds on four first whorls; additional

axial threads not connecting nodules on last coThorl. Su-

ture visible, impressed, not canaliculated. First teleo-

conch whorl convex, sculptured by about 1S—20 proso-

cline smooth riblets, interspace between riblets twice as

wide as riblets; primary spiral cords P2 and P3 appearing

almost immediately, evenly spaced, similar in size and

shape, bearing rounded nodules produced by intersec-

tion with axial riblets. On second whorl, P2 and P3 stron-

ger, P1 appearing at end of whorl, close to P2. On third

whorl, nodules of P1 and P2 becoming sharp, with weak-

C. Vilvens, 2005

Page 51

Dn ee eal

ih fal EE

Figures 1-8. Calliotropis species. 1-6. Calliotropis pulvinaris new species. 1-3. Holotype MNHN, northwestern Madagascar,

18.1X29.0 mm. 4. Paratype IRSNB, West Madagascar, 18.1X26.6 mm. 5-6. Paratype, collection C. Vilvens, West Madagascar

18.3% 25.4 mm. 7-8. C. concavospira (Schepman, 1908), syntype ZMA, Indonesia, 6.08.7 mm.

Page 52 THE NAUTILUS, Vol. 119, No. 1

Figures 9-16. Calliotropis species. 9-12. Calliotropis patula (Martens, 1904). 9-10. Syntype ZMB 55919, between Zanzibar and

Brawa, 16.2X25.1 mm. 11-12. Syntype ZMB 109933, Somaly, 19.0X27.2 mm. 13-14. C. blacki Marshall, 1979, holotype NMUNZ

M226932, Raoul Island, Kermadec group, 11.3X13.0 mm. 15-16. C. vaillanti (Fischer, 1882), MNHN, Azores Islands, 10.6X12.2

mm.

C. Vilvens, 2005

Page 53

Table 1. Calliotropis pulvinaris. Shells measurements in mm for all types cited and type material (n=8). H: height; W: width; HA:

aperature height; TW: number of teleoconch whorls.

Measurement

TW H W HA H/W H/HA

Range 5.75—7.00 15.4-18.3 21.8-29.0 4.00-6.90 0.60—0.70 2,304.50

Mean 6.69 17.11 24.7 5.71 0.69 3.10

Standard deviation 0.41 0.97 2.13 0.93 0.03 0.70

ly adapically oriented tip; P4 partially covered by suc-

ceeding whorl, with nodules smaller and more numerous

than ones on other cords. From fourth whorl on nodules

on cords decrease in size and sharpness, and increase in

number from Pl to P4; cords evenly distributed on

whorl: axial threads in area between cords remain thin,

distance between threads similar to threads width. On

last whorl, P4 peripheral; no secondary spiral cords; pe-

riphery rounded.

Aperture almost circular, forming angle at junction of

inner and outer lips; this angle almost rounded and outer

lip slightly flared in fully mature specimens; inner lip

flanged in a curved arc projecting weakly over umbilicus;

parietal lip forming thin, transparent glaze. Columella

slightly curved, without tooth, abapical part prosocline.

Base moderately convex, with 5 granular spiral cords,

innermost one stronger than others and bordering um-

bilicus; interspace between cords twice as wide as cords,

smaller for two innermost cords on young adult speci-

mens; very fine axial lamellate threads between cords,

poorly visible. Umbilicus wide, funnel shaped, diameter

about 35% of shell diameter, with very fine crowded axial

lamellae and no spiral cord within. Color of protoconch

and teleoconch pinkish white, with no maculations; first

two whorls somewhat brownish.

Type Locality: West Madagascar, 22°17.0' S,

43°02.2' E, 640-660 m, Indian Ocean.

Type Material: Holotype MNHN unnumbered (dd),

Chalutages Vauban, stn. CH 112, coll. A. Crosnier, from

type locality, 18.129.0 mm; Paratype 1 MNHN un-

numbered (dd), 12°50’S, 48°09'E, 580-585 m, north-

western Madagascar, coll. A. Crosnier; eles 2

MNHN (dd), 1 IRSNB (IGnr 30 185) (dd), 1 ZMB

(Moll. 108.519) (dd), 1 G. T. Poppe collection (dd), 1

C.Vilvens collection (dd), all from West Madagascar, off

Mahajanga (formerly Majunga), commercial trawlers

said to be from 800 m.

Etymology: Of a cushion (Latin); with reference to

the soft and oval form of the shell, without angulations.

Remarks: Calliotropis pulvinaris new species is simi-

lar to C. patula (Martens, 1904) (Figures 9-12) from

East Africa (off Somalia and Zanzibar, 977-1019 m), but

this species differs from the new species by having a

subangulated periphery, less tumid whorls, four spiral

cords on base (instead of 5) and also by the nodules of

Pl and P2 that become bigger than those of P3 much

later (4 whorl).

The new species weakly resembles C. concavospira

(Schepman, 1908) (Figures 7-8) from Indonesia (also

deep water from 835 to 883 m), but this smaller species

has only three cords on the whorls and these whorls are

more angulated.

Calliotropis pulvinaris new species may also be com-

pared to C. blacki Marshall, 1979 (Figures 13-14), from

Kermadec Islands, but this Indo-Pacific species is small-

er for a similar number of whorls, has a more elevated

spire and only four spiral cords on the base.

The new ae is also superficially similar to C. vail-

lanti (Fischer, 1882) (Figures 15-16) and C. ambigua

(Dautzenberg and F ischer, 1896), both from eastern “Ne

lantic, and to C. actinophora (Dall, 1890) from western

Atlantic, but these three species have a more elevated

spire, only 3 spiral cords on the whorls and only four

spiral cords on the base.

ACKNOWLEDGMENTS

I would like to express my warm thanks to P. Bouchet

(Muséum national d’Histoire naturelle, Paris) for access

to the malacological resources of the MNHN, and V.

Héros (MNHN) or her help in my search for scientific

papers. Also, I am especially grateful to J. L. Van Goe-

them (Institut royal des Sciences naturelles de Belgique)

for his constant help, particularly with loan of types. I

also would like to thank F. Koehler (Museum fiir Na-

turkiinde, formerly Zoologisches Museum, Berlin), B. A.

Marshall (Museum of New Zealand Te Papa Tongarewa,

Wellington) and R. Moolenbeek (Zoélogisch Museum,

Amsterdam) for the loan of types from their institution.

Finally, I highly appreciate the judicious advice of R.

Houart.

LITERATURE CITED

Barnard, K. H. 1963. Contributions of South African marine

mollusca. Part IV. Gastropoda: Prosobranchiata: Rhipidog-

lossa, Docoglossa. Tectibranchiata. Polyplacophora. Solen-

ogastres. Scaphopoda. Annals of the South African Mu-

seum 47 (2): 201-360.

Crosnier, A. and C. Jouannic. 1973. Note dinformations sur

les prospections de la pente continentale malgache effec-

tuées par le N.O. Vauban (B athymétrie—Sédimentolo-

gie—Péche au chalut). Documents scientifiques du Cen-

tre ORSTOM de Nosy Bé 42.

Hickman, C. S. and J. H. McLean. 1990. Systematic revision

and suprageneric classification of trochacean gasteropods.

Page 54 THE NAUTILUS, Vol. 119, No. 1

Natural History Museum of Los Angeles County Science Vilvens, C. 2001. Description of a new species of Calliostoma

Series vi+ 169 pp. (Gastropoda: Trochidae: Calliostomatinae) from Madagas-

Martens, E. von and J. Thiele. 1904. Die beschalten Gastro- car. Novapex 2: 175-178.

poden der Deutschen Tiefsee-Expedition, 1898-1899. A. Vilvens, C. 2002. Description of Lischkeia mahajangaensis n.

Systematisch-geographischer Teil. Wissenschaftliche Er- sp. (Gastropoda: Trochidae: Eucyclinae: Calliotropini)

gebnisse der deutschen Tiefsee-Expedition auf dem from East Madagascar. Novapex 3: 127-131.

Dampfer “Valdivia” 1898-1899, 7 (A): 1-146. Watson, R. B. 1886. Report on the Scaphopoda and Gas-

Thiele, J. 1925. Gastropoda der Deutschen Tiefsee-Expedition tropoda collected by HMS Challenger during the

II Teil. Wissenschaftliche Ergebnisse der deutschen Tief- years 1873-1876. Report on the scientific results of the

see-Expedition auf dem Dampfer “Valdivia” 1898-1899, voyage of HMS Challenger, 1873-1876. Zoology 15: 1—

17(2): 35-282. 680.

Notice

THE 2005 R. T. ABBOTT VISITING CURATORSHIP

The Bailey-Matthews Shell Museum is pleased to invite applications for the 2005 R. T. Abbott Visiting Curatorship.

The Curatorship, established originally in accordance with the wishes of the late Dr. R. Tucker Abbott, Founding Director of

the Shell Museum, is awarded annually to enable malacologists to visit the museum for a period of one week. Abbott Fellows

will be expected, by performing collection-based research, to assist with the curation of portions of the Museum’s collection

and to provide one evening talk for the general public. The Museum collection consists of marine, freshwater, and terrestrial

specimens. A large percentage of our holdings have been catalogued through a computerized database management system.

A substantial portion of the time will be available for research in the collection, but field work in southwest Florida can be

arranged. The R. T. Abbott Visiting Curatorship is accompanied by a stipend of $1,500.

Interested malacologists are invited to send a copy of their curriculum vitae together with a letter detailing their areas of

taxonomic expertise and research objectives, and to provide a tentative subject for their talk. Send materials to:

Dr. José H. Leal, Director

The Bailey-Matthews Shell Museum

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Applications for the 2005 Visiting Curatorship should be sent no later than May 30, 2005. The award will be announced by

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Volume 119, Number 2

July 20, 2005

ISSN 0028-1344

CONTENTS

Guido Pastorino A revision of the genus Trophon Montfort, 1810

(Gastropoda: Muricidae) from southern South America................... 55

Alvar Carranza Latitudinal trends in shell characters of the neogastropod

Walter Norbis Olivancillaria urceus (Gastropoda: Olividae) in the

temperate SOuthwestempAtlanticn@ Cecil eee tere nen 83

ire

s -

THE NAUTILUS 119(2):55-82, 2005

Page 55

A revision of the genus Trophon Montfort, 1810 (Gastropoda:

Muricidae) from southern South America

Guido Pastorino

Museo Argentino de Ciencias

Naturales

Ay. Angel Gallardo 470, 3° piso lab. 57

C1405DJR Buenos Aires

ARGENTINA

[email protected]

ABSTRACT

The genus Trophon from southern South America is revised

and restricted to ten valid species from among the 36 nominal

species still currently used. In addition, a new species, Trophon

parodizi from Patagonian waters, is described. Adult specimens

of the new taxon are illustrated, described and compared with

other living species of the same genus and similar geographic

distribution. Redescription and re-illustration of types, based

on material from several institutions around the world are pro-

vided for Trophon geversianus (Pallas, 1774), T. plicatus (Light-

foot, 1786), T. patagonicus (d’Orbigny, 1839), T. acanthodes

Watson, 1882, T. pelseneeri Smith, 1915, T. amettei Carcelles,

1946, T. clenchi (Carcelles, 1953), T. wilhelmensis Ramirez-

Bohme, 1981, and T. bahamondei McLean and Andrade, 1982.

In addition, “Trophon” malvinarum Strebel, 1908, assigned to

genera incerta, is also redescribed and illustrated.

INTRODUCTION

Among many groups of marine gastropods from South

American waters in need of a modern comprehensive

revision, the high diversity and abundance exhibited by

the muricid genus Trophon Montfort, 1810, renders it

particularly interesting. Early collections include a large

and varied array of specimens obtained by 19" wentinnsy

expeditions from shallow waters of a vast area including

more than 5,000 km of coast in Argentina. A taxonomic

revision of these gastropods revealed that a large num-

ber of names should be placed in synonymy. The tax-

onomy and nomenclature thus clarified is sure to im-

prove the usefulness of this genus and its species as tools

for biogeographic and evolutionary interpretations, with-

out forgetting that clear specific delimitation is crucial

to other uses such as the commercial exploitation of the

type species (Trophon geversianus) in southern Chile.

The subfamily Trophoninae is one of the most con-

spicuous groups of marine gastropods living presently

around the southern tip of South America. The southem

origin of the group seems to be beyond doubt, as dis-

cussed by Griffin and Pastorino (2005), when revising

the numerous extinct species appearing in the fossil re-

cord since the late Oligocene.

This article constitutes a review of all living species of

Trophon from both coasts of southern South America.

The study involves only those taxa living in environments

associated with the continental shelf. Accordingly, T. mu-

crone Houart, 1991, from 1500-1575 m off Brazil and

the subantarctic T. veronicae Pastorino, 1999, are not

considered herein. These two deep-water species seem

to belong in a different group according to data available

on the radula, protoconch, and penis of T. veronicae.

Such anatomical data remain unknown for T. mucrone.

In addition, T. ohlini Strebel, 1904, with a distinct pro-

toconch and radula, different from those of other Pata-

gonian species and resembling the boreal Boreotrophon

truncatus, will be the subject ‘of a future paper. Finally,

T. triacanthus Castellanos et al., 1987, recently described

under Trophon, is also considered as belonging to a dif-

Apyxistus—according to several

differences in protoconch and shell morphology.

Houart (2003) recently described three new species

under the genus Trophon sensu lato from dredgings

more than 1000 m in depth. No radular, anatomical, or

protoconch information is included in the descriptions.

However, enough differences can be observed in the

shells that, as mentioned by Houart himself, a new genus

may be granted for these species. A similar situation is

true for “Trophon” malvinarum; however, as there are

no accurate illustrations or recent descriptions of this

species I decided to include it in this work as belonging

to an indeterminate genus.

For each of the species considered herein, adult spec-

imens, operculum, gross anatomy, radula, protoconch,

and ultrastructure of the shell are described whenever

enough material was available. This work is part of a

complete revision of the genus including all species from

South America and Antarctica.

MATERIALS AND METHODS

All the material examined is housed in the collections of

the Museo Argentino de Ciencias Naturales “Bermardino

Page 56

THE NAUTILUS, Vol. 119, No. 2

Rivadavia”, Buenos Aires (MACN-In); Museo de La Pla-

ta, La Plata (MLP): National Museum of Natural His-

tory, Smithsonian Institution, Washington, D.C.

(USNM) and American Museum of Natural History,

New York (AMNH). Part of the type material is housed

in the following museums: The Natural History Muse-

um, London, (BMNH); Zoologisches Institut und Zool-

ogisches Museum der Universitat Hamburg, (ZMH);

Swedish Museum of Natural History, Stockholm,

(NHRM); Museum national d’Histoire naturelle, Paris

(MNHN): Museo Nacional de Historia Natural, Santia-

go, Chile (MNHNS); Museo Nacional de Historia Nat-

ural, Montevideo, Uruguay (MNHNM); Academy of

Natural Sciences of Philadelphia (ANSP) and Los An-

geles County Museum of Natural History (LACM). Fi-

nally, several historical specimens from the Auckland In-

stitute and Museum, New Zealand (AK) were examined

for comparative purposes. Material from the USNM

originates from the United States Antarctic Program

(USAP) and was mostly collected by two ships: R/V

Hero and R/V ELTANIN. Material collected by the Uru-

guayan ship R/V ALDEBARAN is also included.

Dissections were performed on ethanol-preserved

specimens for study of gross anatomy, with emphasis on

the morphology of the anterior alimentary system, and

the pallial portions of the male and female reproductive

systems. Radulae were prepared according to the meth-

od described by Solem (1972) and observed using a

LEO 440 scanning electron microscope (SEM) at the

USNM and a Philips XL 30 at MACN. Radular termi-

nology follows Kool (1993, fig. 6B). Shell ultrastructure

data were procured from freshly fractured colabral sec-

tions taken from the central portion of the lip on the

last whorl of two individuals per taxon, whenever suffi-

cient material was available.

Photographs were taken using a digital scanning cam-

era. Several images were scanned from black and white

35 mm negatives using a slide scanner. All images were

digitally processed.

For the convenience of the reader in the Additional

Material Examined sections, “D” means that the speci-

mens were collected dead and “A” means alive.

SYSTEMATICS

Class Gastropoda Cuvier, 1791

Subclass Ortogastropoda Ponder and Lindberg, 1996

Superorder Caenogastropoda Cox, 1959

Order Sorbeoconcha Ponder and Lindberg, 1996

Infraorder Neogastropoda Wenz, 1938

Family Muricidae da Costa, 1776

Subfamily Trophoninae Cossmann, 1903

Genus Trophon Montfort, 1810

Type Species: Murex magellanicus Gmelin, 1791

(=Buccinum geversianus Pallas, 1774) by original des-

ignation. Polyplex Perry, 1811 (type species Polyplex bul-

bosa Perry, 1811 =Buccinum geversianum Pallas, 1774)

and Muricidea Swainson, 1840 (type species Murex ma-

gellanicus Chemnitz, 1780 (nomen nudum) =Murex ma-

gellanicus Gmelin, 1791 =Buccinum geversianum Pallas,

1774) are synonyms.

Description: Shells variable in size, ranging from

about 1 to more than 10 centimeters high, fusiform, su-

bquadrate to elongate. Protoconch paucispiral, with 2 to

2¥%2 asymmetrical, smooth, slightly globose, and regularly

convex whorls. Spire of about 4 whorls, equal or slightly

shorter than aperture height, never higher. Subsutural

ramp generally present, variously developed and slightly

inclined posteriorly, defining a conspicuous keel. Axial

sculpture variable, from weak growth lines to strong la-

mellae—in some instances even growing over the sub-

sutural ramp; axial sculpture generally better developed

than spiral sculpture. Lamellae along keel sometimes

growing into fairly strong, always open, spines, which

may even curve backwards. Spiral ornamentation vari-

ably developed; sometimes missing, but generally con-

sisting of slightly rounded and equally developed cords,

wider than interspaces and sometimes accompanied by

weaker secondary intercalated ones. Spiral omamenta-

tion usually restricted to surface abapical to keel, while

missing along subsutural ramp and in some cases only

present in the earliest whorls. Aperture subovoid; outer

lip sharp, sometimes slightly reflected, but always

smooth throughout. Siphonal canal always present and

open, although variably developed in length, sometimes

curved. Umbilicus variable, usually open, wide, although

in some species totally absent or represented by a chink.

Shell ultrastructure arranged invariably in 2 layers: in-

nermost layer of crossed lamellar aragonite, outer layer

of amorphous calcite. Relative thickness of both layers

variable according to species or—within a given spe-

cies—latitude at which the population lives.

Radulae rachiglossate, rachidian teeth with 3 median

cusps, the central one the larger and the lateral ones

with a denticle, sometimes obsolete but always present,

attached to the interior margin (never free). Rachidian

base always sinuous and with the base offset under the

proximal tooth. Marginal cusps always single, never bifid.

Lateral teeth always thin, with the attachment area also

thin. Operculum circular or suboval tear-shaped, attach-

ment area with horseshoe shaped scars.

Accessory salivary glands always developed, tubular,

single or coiled and unfilled. Esophagus with a loop run-

ning along the left side of the gland of Leiblein; esoph-

ageal glands externally invisible.

Egg-capsules always erect, never lenticular, usually

with nurse eggs.

Trophon geversianus (Pallas, 1774)

(Figures 1-21)

“Purpurschnecken” Knorr, 1769: 47, pl. 30, fig. 2.

“Buccin feuilleté” Knorr, 1770: 53, pl. 30, fig. 2.

Buccinum geversianum Pallas, 1774: 33, pl. 3, figs. 1, 2.

Buccinum foliaceum multifariam frondosum Chemnitz, 1780:

130, pl. 139, fig. 1297 [non-binominal, rejected by Opin-

ion 184 (ICZN, 1944) [Lectotype of Murex magellanicus

Gmelin, 1791 (Beu, 1978)].

G. Pastorino, 2005 Page 57

Figures 1-16. Trophon geversianus (Pallas, 1774). 1-3. MACN-In 36036, Punta Cavendish, Puerto Deseado, Santa Cruz province

in 5 m. 4-5. MACN-In 36042, Cueva del Indio, Puerto Deseado. 6-9. MACN-In 36041, Sierra Grande, Rio Negro province, in

tide pools. 10-11. MACN-In 36037, Punta Penas, San Julién, Santa Cruz Province in 2 m. 12. MACN-In 36043, Bahia Almanza,

Puerto Harberton, Tierra del Fuego, in 3 m. 13-14. MACN-In 36038, both specimens from Playa La Mina, San Julian, Santa Cruz

Province intertidal. Scale bar for all shells = 1 cm. 15-16. Two views of protoconch, arrow head the transition to teleoconch. Scale

bar = 500 pm.

Page 58

THE NAUTILUS, Vol. 119, No. 2

Figures 17-21. Trophon geversianus (Pallas, 1774). 17.

Buccinum fimbriatum Martyn, 1784: fig. 6

Murex magellanicus Gmelin, 1791: 3548 partim (var. B excl.):

d'Orbigny 1841: 451; Wood, 1828: 127, pl. 26, fig. 90;

Hanley, 1856: 132, pl. 26, fig. 90.

Aigpirenze jolkesen Réding, 1798: 116.

PMurex ventricosus Misia, 1810: 178.

Polyplex bulbosa Perry, 1811: pl. 9, fig. 5.

Murex foliatus Schumacher, 1817: 215, sensu Vokes 1971.

Fusus magellanicus Lamarck. Gray, 1839: 118.

?Murex varians d’Orbigny, 1839: pl. 42, figs. 4-7; d’Orbigny,

1841:452.

T. geversianus Pallas—Montfort, 1810: 483, fig.; H. and A.

Adams, 1853: 77, pl. 8, fig. « 2 C.; Tapparones Canefri, 1874:

15; Kobelt, 1878: 205, pl. 72, fig. 1-3; pl. 73, fig. 1; Sow-

erby oe 1880: pl. 404, figs. 7,8; ‘Tryon, 1880: 144, pl. 32,

figs. 337-340, 343-347; vl. 70, figs. 433: Watson, 1886:

164: Rochekeae and Mabille, 1889: H.53: Strebel, 1904:

173, pl. 4, figs. 11-23; pl. 5, figs. 2442; pl. 6, figs. 43-52;

Lamy, 1906: 3; Ihering, 1907: 404; Melvill andl. Standen,

1907: 106: Strebel, 1908: : 7, pl. 6, figs. 94 a, b.; Carcelles,

1946: 60, figs. 1-5; 1946: 69, figs. 6a,b; 7a,b,c,d; 8; Powell,

1951: 151, fig. L, 81; N, 107; Castellanos, 1970: 76, pl. 5

fig. 2; Dell, 1971: 210; Harasewych, 1984: 13, figs. 1-3,

19- 25. oe 1991: 7, fig.; 1992: 3, figs 1c, d; 3c, d; Kool,

1993: 47, figs. S14, 30-31; Gratailinaes and Landoni,

1993: 5 all I figs. 1-15, 18-21.

Radula, frontal view. Scale bar =

100 pm. 18. Lateral view of the same

radula. Scale bar = 30 xm. 19. Ultrastructure of the shell. Scale bar = 100 xm. 20. Penis, critical-point dried. Scale bar = 800

wm. 21. two views of the operculum. Scale bar = 1 cm.

Fusus intermedius Hupé in Gay, 1854: 166, pl. AL fig. 6, non

Cristofori and Jan, 1832 nomen nudum; nec A. J. Mich-

elotti, 1846 nomen nudum: nec G. Michelotti, 1847;

Rochebrune and Mabille. 1889: H.53.

F. geversianus Pallas—Hupé in Gay, 1854: 167; Gould, 1861:

pl. 16, fig. 277 a, b.

Trophon geversianus var. calva Kobelt, 1878: 305, pl. 75; HS il.

T. geversianus var. lirata Kobelt, 1878: 305, pl. 76, fig. 1,

T. philippianus Dunker in Kobelt, 1878: 277, pl. 72, figs. a 5;

Melvill and Standen, 1907: 107; Powell, 1951: 152.

T. philippinarum Dunker. Sowerby I, 1880: pl. ae. » Hg. Dil.

?T. varians (d’Orb.)—Carcelles, 1943: 431, figs. 1, 2, 3, 6,

Castellanos, 1970: 75, pl. 5, fig. 3, 4 Wellies, 1999: see

T. plicatus (Lightfoot).—Calvo, 1987: 135, fig. 99.

Trophon sp. Vokes 1991: 9, figs. 1-13.

Description: Shell large (up to 100 mm) and extreme-

ly variable, fusiform, subquadrate profile, chalky, whitish;

protoconch of 2 whorls, smooth, cylindrical, slightly glo-

bose, slightly asymmetrical; teleoconch of 4 shouldered

whorls, spire less than % of total shell height. Spire angle

about 50°; suture impressed; subsutural shelf straight,

aperture ovoid, interior glossy pinkish; anterior siphonal

canal moderately long (half the height of aperture); um-

bilicus closed or deep, some specimens with a pseu-

G. Pastorino, 2005

doumbilical chink; outer lip rounded, with reflected edg-

es; inner lip curved, adpressed.

Axial omamentation of irregular, low lamellose varices

on first whorls, becoming 8-10 well-defined lamellae on

last ones. Lamellae growing across entire whorl, attached

to the shell, sometimes curving adaxially. Lamellae end-

ing in shallow peripheral spine, in some specimens grow-

ing adapically.

Spiral ornamentation of about 15 cords beginning at

periphery of whorls. Smooth specimens common in in-

tertidal pools and mytilid banks. Coloration varying from

creamy white to dark brown. Growth lines regularly

spaced, present throughout shell. Geographic variation

conspicuous from north to south and from intertidal to

infralittoral specimens, expressed as a series of smooth

to profusely ornamented specimens according to area of

collection.

Shell ultrastructure composed of two layers; inner-

most layer (50% shell thickness) of colabrally aligned

crossed lamellar aragonite, outer layer (50% shell thick-

ness) with amorphous calcite.

Operculum oval, brownish, with terminal nucleus. Ex-

ternal surface covered by concentric, irregular, growth

lines. Inner surface attachment area with 3 horseshoe-

shaped scars, thick glazed rim present in all specimens.

Anatomical and radular characters as described by

Harasewych (1984) and Kool (1993).

Type Material: The type material upon which Pallas

(1774) based Buccinum geversianum could not be lo-

cated. It could not be found at the Zoological Institute

of the Russian Academy of Sciences (ZIL) St. Peters-

burg, where most of the material studied by that author

is housed (B. Sirenko, pers. comm. ).

Additional Material Examined: 53°39’ S, 70°55.5’ W,

5 A, R/V Hero Cruise 702, Sta. 466, 25 April 1970, 20

m (USNM 901605): 53°39’ S, 70°55.5’ W, 6 A, R/V

Hero Cruise 702, Sta. 473, 26 April 1970, 15-18 m

(USNM 901609); 53°30'48” S, 70°50'33” W, 4 A, R/V

Hero Cruise 692, Sta. 69-11, 24 April 1969, intertidal,

(USNM 901604); 53°24.8’ S, 69°39.2’ W, 1 D, R/V

Hero Cruise 702, Sta. 481, 27 April 1970, 18 m, (USNM

901605); 53°17’ S, 68°13’ W, 1 A, B/V Hero Cruise 712,

Sta. 71-2-6, 21 April 1971, 1 m, (USNM 886739);

53°51'32” S, 70°25'52” W, 1 A, R/V Hero Cruise 692,

Sta. 69-22; 13 May 1969, 2-3 m (USNM_ 886187):

53°37.9' S, 70°14’ W, 1 D, R/V Hero Cruise 702, Sta.

486, 28 April 1970, 292-296 m (USNM 901602); 54°59’

S, 64°53’ W, 1 D, R/V ELTANIN Cruise 11, Sta. 970, 11

February 1964, 586-641 m (USNM 870515); 53°48.7’ S,

70°24.1' W, 1 D, R/V HERO Cruise702, Sta. 489, 29 April

1970, 13-18 m (USNM 901601); 54°27’ S, 66°12’ W, 2

D, R/V ELTANIN Cruise 6, Sta. 453, 21 January 1963, 31

m (USNM 901600); 53°26’ S, 68°35’ W, 6 A, R/V EL-

TANIN Cruise 21, Sta. 297, 12 January 1966, 0 m (USNM

901607); 53°35’ S, 70°50’ W, 1 D, 1 A, B/V ELranin

Cruise 21, Sta. 292, 8 January 1966, 0 m (USNM

901606); 53°17’ S, 68°13’ W, 3 D, B/V ELTANIN Cruise

712, Sta. 71-2-6, 21 April 1971, 0-1 m (USNM 901608):

Page 59

Puerto Basil Hall, Isla de los Estados, 10 A, 21 May

1971, 0-1 m (MLP 4243); Buen Suceso Bay, Tierra del

Fuego, 4 A, 23 October 1941 (MLP 27218); San Julian,

Punta Pefias, 6 A, 1 March 1924 (MLP 526 partim);

Puerto Lobos, Chubut, 12 A, 2 February 1938 (MLP

2021); Puerto Golondrina, Ushuaia, 9 A, January 1962

(MLP 27201): Puerto Piramides, Chubut, 1 A (MLP

4715); San Julian, 8 A (MLP 1583); Puerto Hoppner,

Isla de los Estados, 7 A (MACN-In 22547): Punta Cav-

endish, Puerto Deseado, 7 A, 6 February 1961 (MACN-

In 26171); Punta Cavendish, Puerto Deseado, Santa

Cruz province, 5 m (MACN-In 36036); Chubut, 6 A

(MACN-In 4097); Tierra del Fuego, 4 A (MACN-In

5777-1); Cueva del Indio, Puerto Deseado (MACN-In

36042); Sierra Grande, Rio Negro province, in tide pools

(MACN-In 36041); Punta Pefias, San Julian, Santa Cruz

province, 2 m (MACN-In 36037); Playa La Mina, San

Julidn, Santa Cruz province intertidal (MACN-In

36038); Bahia Almanza, Puerto Harberton, Tierra del

Fuego, 3 m (MACN-In 36043); San Antonio Oeste, Rio

Negro, intertidal, 3 A (MACN 35387); 54°47'36” S,

64°22'35" W, 1 A, B/V ELTaANINn Cruise 712, Sta. 71-2-

44, 24 May 1971, intertidal (USNM 881127): 54°46'12”

S, 64°24'42” W, 1 A, B/V ELTANIN Cruise 712, Sta. 71-

2-46, 21 May 1971, intertidal (USNM 881131); 54°4536”

S, 64°02’36" W, 1 A, R/V ELTANIN Cruise 7151, Sta. 869,

23 October 1971, intertidal (USNM 881132): 54°45'45”

S, 64°09'55” W, 1 A, R/V Hero Cruise 712, Sta. 71-2-

40, 21 May 1971, intertidal (USNM 881130); 54°47'48”

S, 65°16’ W, 1 A, R/V HERO Cruise 712, Sta. 71-2-8, 23

April 1971, intertidal (USNM 881126); 54°48’, 65°14’ W,

1 A, R/V Hero Cruise 712, Sta. 71-2-14, 25 April 1971,

intertidal (USNM 881129): Punta Arenas, Cabeza de

Mar, Chile (ANSP 101444): Ushuaia (ANSP 316762):

Malvinas Is. (ANSP 277535 and 277538); Cape Fair-

weather, Santa Cruz province, Argentina (ANSP 78080);

Straits of Magellan (ANSP 36241 and ANSP 366497);

Puerto San Julian (ANSP 312324): Santa Cruz River

(ANSP 101445); Puerto Gallegos (ANSP 312319); Golfo

San Jorge (ANSP 178645); Puerto Madryn, Chubut

(ANSP 170471); Puerto Parry, Isla de los Estados, 54°46’

S, 64°23’ W (ANSP 402810): mouth of Santa Cruz River

(ANSP 88536); Punta Arenas (ANSP 88549): Puerto

Madryn (ANSP 170474); Punta Arenas, Chile (ANSP

199711).

Distribution: Trophon geversianus has the widest

geographic range of all species of Trophon, i.e., from

Buenos Aires province to Burdwood Bank in the south-

western Atlantic, Tierra del Fuego and Malvinas Is., and

the Magellan Strait in Chile. Literature records from

around Antarctica are almost certainly wrong assign-

ments, probably mistaking it for Trophon nucelliformis

Oliver and Picken, 1984, T: macquariensis Powell, 1957,

or T. albolabratus Smith, 1875.

Remarks: Trophon geversianus is the best-known spe-

cies of the entire genus. The great morphological vari-

ation can be appreciated from the large number of

names proposed for the different morphological variants

Page 60

in this species. Zaixso (1973) and Penchaszadeh (1976)

both studied the egg capsules of this species but only

the latter confirmed the existence of short-lived nurse

eggs in the capsules.

Trophon varians is a dubious species described by

dOrbigny from material he collected in northern Pata-

gonia (“au sud du Rio Negro”). This species has no la-

mellae, weak spiral cords, and its shell is unusually thick.

Houart (1998) illustrated a paralectotype (as syntype)

housed in Paris and Aguirre (1993) designated and fig-

ured the lectotype (BMNH 1854.12.4.539) from 13 syn-

types from the BMNH collection. According to

dOrbigny’s illustration (Plate 42, figs. 4, 5) and the lec-

totype and the paralectotype housed in Paris, it appears

that its distinctive characters are the thickness of the

shell and absence of lamellae. However, despite this dif-

ference between T. geversianus and these primary types

of T. varians, the rest of the paralectotypes are very sim-

ilar to other thin-shelled specimens of Trophon gever-

sianus usually found exposed during low tides in north-

em Patagonia, Golfo San Matias and around the Valdés

Peninsula. The anatomy and radula of the latter are

identical to those of T. geversianus. The specimens with

heavy shells are characteristic of the mouth of Rio Ne-

gro. Nothing is known about the anatomy and radula of

these heavy specimens. Perhaps these characters may

prove that it is only a local variation of Trophon gever-

sianus, as it was suggested originally by d’Orbigny him-

self (1841: 452). D’Orbigny (1841: 452; 1839, plate 42,

figs. 6-7) also described and illustrated the egg-capsules

from what he supposed were T. varians. The illustration

agrees better with capsules of Urosalpinx haneti (Petit,

1856).

Trophon plicatus (Lightfoot, 1786)

(Figures 22-42)

Le Sabot Magellanique Favanne, 1780: 342, pl. 79, fig. L., no-

men nudum.

Buccinum laciniatum Martyn, 1784: fig. 42, nomen nudum.

Murex plicatus Lightfoot, 1786: 104.

M. lamellosus Martyn —Gmelin, 1791: 3536; Wood and Han-

ley, 1856: 133, pl. 27, fig. 100

Polyplex gracilis Perry, 1811: pl. 9, fig. 4.

Fusus laciniatus Martyn.—Reeve, 1847: spec. 14, figs. a, b, c.;

Hupé in Gay, 1854: 168; Gould, 1861: pl. 16, fig. 278.

Trophon laciniatus Martini—H. and A. Adams, 1853: 77, pl.

8, figs. 3 a, b.; Kobelt, 1878: 280, pl. 72, figs. 6, 7; Tryon,

1880: 143, pl. 31, figs. 330-332. Rochebrune and Mabille,

1889: H.53; Strebel, 1904: 199, pl. 3, figs. 1-8; Lamy,

1906: 3; Strebel, 1908: 37; Castellanos, 1970: 74, pl. 5,

fig. 1.

>Trophon antarcticus Philippi, 1868: 225 (sensu Tryon 1880).

T. lacineatus Martyn. Sowerby II, 1880: pl. 404, fig. 13.

T. (Stramonitrophon) laciniatus (Martyn)—Powell, 1951: 156,

fig. L, 86.

T. (Stramonitrophon) lamellosa (Gmelin).—Dell; 1971: 212.

T. plicatus (Lightfoot, 1786)—Rehder, 1967: 20; Cemohorsky,

Iehyae la, fig. 18: Vokes, 1991: 7, fig. ; 1991b: 9, figs. 14—

16; 1992: 3, figs. 2, b, c, d.; Castellanos and Landoni, 1993:

5, figs. 16, 15, 22.

THE NAUTILUS, Vol. 119, No. 2

Description: Shell of medium to large size (to 50

mm), smooth, fusiform, thin, somewhat chalky; proto-

conch smooth of 2% whorls; teleoconch of 6 shouldered

whorls, spire less than % of total shell height. Spire angle

about 50°; suture impressed; subsutural ramp straight,

aperture small, ovoidal, interior pale brownish; anterior

siphonal canal long (more than half height of the aper-

ture), narrow, curved, open; umbilicus absent; outer lip

rounded with reflected edges; inner lip curved, adpres-

sed. Axial ornamentation of irregular, low lamellose var-

ices in the first whorls, that become 8—10 real lamellae

in the last ones. Lamellae growing over the entire whorl,

but attached to the shell, sometimes curving adaxially.

Lamellae ending in a peripheral spine, in some speci-

mens growing adapically. Spiral ornamentation poorly

developed to almost smooth, sometimes consisting of 6

weak cords on the base of the last whorl and the back

of the siphonal channel. Regular, very weak growth lines

present throughout shell.

Shell ultrastructure composed of two layers; inner-

most layer (25% of shell thickness), composed of cola-

brally aligned crossed lamellar aragonite, outer layer

thick (75% of shell thickness) with amorphous calcite

(Figure 42).

Operculum oval, with terminal nucleus. External sur-

face covered by concentric, irregular, growth lines. Inner

surface attachment area with two or three horseshoe-

shaped scars, glazed rim present in all specimens (Figure

36).

Animal of medium size relative to shell. Mantle large,

mantle roof thin. Cephalic tentacles broad in basal half,

flat, blunt, with rounded large black eyes in the middle;

mantle edge thickened, smooth; pallial organs arranged

as in other rachiglossans; brown osphradium less than

half of ctenidium length, slightly asymmetrical, with 50—

55 leaflets per side; ctenidium as wide as osphradium,

containing triangular leaflets. Hypobranchial gland

brownish and inconspicuous, rectum to the right of hy-

pobranquial gland.

Proboscis pleuroembolic, long, broad. Radular ribbon

extending beyond rear of buccal mass, long (0.69 X ap-

erture height vs. 0.78 X aperture height in Trophon gev-

ersianus). Esophagus loops beneath buccal mass, where

esphagus receives embedded ducts of salivary glands an-

terior to a small Leiblein valve. Esophagus curving dis-

tinctly and running along entire side of gland of Lei-

blein. Esophageal glands (“Glandule framboise”) whitish

in color, posterior to nerve ring slightly marked on the

external side of the esophagus. Large salivary glands en-

veloping Leiblein valve and accessory salivary glands. Ac-

cessory salivary glands distinctly large, sometimes dark-

colored, tubular, compact, coiled, embedded in salivary

glands. Gland of Leiblein conspicuous, brown, ending in

a medium size blind duct and very small ampulla.

Radula rachiglossan with rachidian teeth wide (~150

wm), central cusp thin, large; lateral cusps wider and

slightly shorter than central cusp, with inner edge

straight; with sharp straight denticle in upper third of

internal edge of lateral cusp, external edge with 6 very

G. Pastorino, 2005 Page 61

Figures 22-36. Trophon plicatus (Lightfoot, 1786). 22-24. MACN-In 36033, Bahia Ensenada, Ushuaia, Tierra del Fuego, Ar-

gentina. 25-26. MACN-In 9032-16, Comodoro Rivadavia, Chubut province, Argentina. 27. BMNH 19990384, 45°55.219’ S,

73°39.522' W, intertidal, Islet NE shore of Isla Huemules, Golfo Elefantes, Chile. 28-29. AK 133035, Station WS788 Discovery,

45°07’ S, 65°W. 30-31. MACN-In 36034, Punta Pefias, San Julian, Santa Cruz province, Argentina. 32. Same lot as Figure 6. 33.

MLP 27202, Bahia Golondrina, Ushuaia, Tierra del Fuego. 34-35. Two protoconchs, scale bars = 400 wm. 34. USNM 870535,

52°30’ S, 67°14’ W, in 82 m. 35. USNM 901620, 53°35’ S, 69°45’ W, 1 D, R/V Hero Cruise 692, Sta. 404, 37-46 m. 36. MLP

27232, operculum, external (left) and internal (right) views, scale bar = 1 em. Scale bar for all shells = 1 cm.

Page 62

THE NAUTILUS, Vol. 119, No. 2

Figures 37-42. Trophon plicatus (Lightfoot, 1786). 37. MLP 27232

g 27232, Dorsal view of radular-ribbon. Scale bar = 50 pm. 38.

MLP 526, San Julian, Punta Pefias, 1 March 1924, rachidian teeth. Scale bar = 30 ym. 39. Variations of rachidian teeth, Rocamora,

Ushuaia, 1 A, 8-10 m (MACN-In 36053). Scale bar = 30 pm. 40. MLP 27202, Bahia Golondrina, Ushuaia, Tierra del Fuego,

Argentina, detail of the rachidian teeth. Scale bar = 20 jxm. 41. Same specimen as 40, lateral view of rachidian teeth, scale bar =

30 ym. 42. USNM 870535, 52°30’ S, 67°14’ W, in 82 m, shell ultrastructure, fracture surface commarginal. Scale bar

= 30 pm.

G. Pastorino, 2005

Page 63

well defined denticles, always present. Base of rachidian

tooth sinuous, sliding beneath base of next tooth. Mar-

ginal area with single conspicuous cusp. Lateral teeth

with single, long and narrow cusp, slightly attached basal

plate. Cusps of rachidian teeth pyramid-like in lateral

view (Figures 38-40).

Male and female organs as in Trophon geversianus

(see Harasewych, 1984).

Type Locality: Islas Malvinas.

Type Material: Probably lost. According to Dance

(1966) part of the material from the Portland Catalogue

is in London (BMNH), however this is not the case of

T. plicatus.

Additional Material Examined: 52°30’ S, 67°14’ W,

2 D, R/V ELTANIN Cruise 11, Sta. 980, 14 February

1964, 82 m (USNM 870535): 53°35’ S, 69°45’ W, 1 D,

R/V Hero Cruise 692, Sta. 404, 37-46 m (USNM

901620): 52°56’ S, 75°00’ W, 1 D, R/V ELTaAnin Cruise

11, Sta. 958, 5 February 1964, 92-101 m (USNM

870423); 53°06’ S, 67°04’ W, 3 D, B/V HERO Cruise 702,

Sta. 450, 5 March 1970, 86 m (USNM 901622):

53°39'24” S_ 70°55'30"” W, 1 D, R/V HERO Cruise 702,

Sta. 467, 25 April 1970, 24 m (USNM 901623): 52°35’

S, 65°08’ W, 1 D, R/V ELTanIn Cruise 11, Sta. 976, 13

February 1974, 128 m (USNM 870525); 46°04’ S, 83°55’

W, 1 A, B/V Etranin Cruise 25, Sta. 326 9 October

1966, 298 m (USNM 901621): 53°48.7’ S, 70°24.1’ W,

1 D, R/V Hero Cruise 702, Sta. 489, 29 April 1970, 13—

18 m (USNM 901624); 53°32’ S, 64°57’ W, 2 A, R/V

ELTANIN Cruise 11, Sta. 974, 12 February 1964, 119-

124 m (USNM 881960): 53°39’ S, 70°55.5’ W, 1 A, R/V

HERO Cruise 702, Sta. 466, 20 m (USNM 901754): Ba-

hia Golondrina, Ushuaia, 1 A, (MLP 27202): Rocamora,

Ushuaia, 1 A, 8-10 m (MACN-In 36053): Bahia Ensen-

ada, Ushuaia, Tierra del Fuego, (MACN-In 36033);

45°07’ S, 65°W, 1 A, Discovery Station WS788, 13 De-

cember 1931, 82-88 m (AK 133035) illustrated in figs.

28-29: Comodoro Rivadavia, 17 D (MACN-In 9032-16):

54°34’ S, 64°00'18” W, 1 D, 1 A, B/V Hero Cruise 715,

Sta. 870, 24 October 1971, 84 m (USNM 881128): Bahia

Buen Suceso, 1 A, 23 October 1941, (MLP27230): Cabo

Colnett, Isla de los Estados, 1 A, R/V HERO Cruise 712,

Sta. 853, 20 October 1971, 91 m (USNM 869720): 2 D,

78 m (USNM 96176); Orange Harbor (USNM 5676);

Paso Richmond, Tierra del Fuego 55 m, (MACN-In

24940): 55°7’ S, 66°33’ W, 82 m (MACN-In 23938): Ti-

erra del Fuego (MACN-In 5240-2); Punta Pejfias, San

Julian, Santa Cruz province, Argentina, 4 A (MLP

27232); Punta Pefias, San Julidn, Santa Cruz, Argentina,

1 A (MLP 27212); Puerto San Julian, Santa Cruz prov-

ince, Argentina (49°15’ S, 67°39’ W), 4 A, 2-3 m

(MACN-In 36034); 45°55.219’ S, 73°39.522’ W intertid-

al, Islet NE shore of Isla Huemules, Golfo Elefantes,

Chile (BMNH 19990384).

Distribution: This is a typical species from the Ma-

gellanic province. It has been recorded from Peninsula

Valdés to Tierra del Fuego in Argentina and Chile to

49°S in the north (Reid and Osorio, 2000).

Remarks: The authorship of the name according to

Dance (1962) and Rehder (1967) should be credited to

J. Lightfoot, the anonymous compiler of the “Catalogue”

where the name was used for the first time, not to So-

lander.

There is an interesting range of conchological varia-

tion in Trophon plicatus, from almost smooth specimens

to highly lamellate. However, the profile is always slen-

der. Trophon bahamondei McLean and Andrade, 1982,

is a morphologically similar species, with peripheral

spines and shallow lamellae. On the other hand, T. ba-

hamondei has no spiral ornamentation and Trophon pli-

catus has 6 weak cords on the base of the last whorl and

the back of the siphonal canal.

The gross anatomy is that customary for most Pata-

gonian Trophoninae. However, a distinctive anatomical

feature is the morphology of the accessory salivary

glands, which are compact, tubular, somewhat coiled and

large, and completely embedded in the salivary glands.

Most Patagonian Trophoninae (e.g., T. bahamondei as

well as T: geversianus), have small, kidney-shaped, ac-

cessory salivary glands.

Radular features of T. plicatus are clearly different

from T. bahamondei (see Figures 26-27). The latter has

a distinctive small cusp on the outer margin of the lateral

cusp on the rachidian teeth. In addition, the rachidian

base is thinner and wider.

Trophon antarcticus Philippi, 1868, is probably a syn-

onym according to Tryon (1880); however, the type ma-

terial is missing and the description is rather obscure.

Powell (1951) described the subgenus Stramonitro-

phon to include only T. plicatus [(as T. laciniatus Martyn,

nomen nudum rejected by Opinion 456 (ICZN, 1957)]

a species with Stramonita-like radula, i.e., rachidian

teeth with three cusps, where the marginal side of the

lateral cusps bears several denticles. This radula illus-

trated by Powell (1951: 194, fig. L, 86) was dissected

from the specimen illustrated in figs. 28-29. The radular

morphology of T. plicatus is different from that of T.

geversianus. However, Coronium coronatum (Penna-

Neme and Leme, 1978) and TL. acanthodes Watson,

1882, bear the same denticles on the lateral cusps of the

rachidian teeth. This appears to be a common feature in

several Trophoninae from the southwestern Atlantic.

The shell of Trophon iarae Houart, 1998, shows some

similarity with some specimens of T. plicatus. It was

based on only two specimens collected by fishing boats

apparently from Brazil and off Uruguay. The anatomy

and soft parts of T. iarae remain unknown. The radula

apparently has been illustrated by Calvo, 1987 (although

according to Houart, 1998: 127, there is no certainty

about the identity of the specimen from where this rad-

ula was taken). In any event, this illustration does not

allow for a detailed comparison with other species. Fur-

ther comments about the validity of this species or its

Page 64

affinity with other species of Trophon mostly depends

on anatomical data which are not available.

Houart (1998: 127) mentioned the specimen of T. pli-

catus illustrated by Cermohorsky (1977) from Lively Is.

(Malvinas Is.) as belonging to his new species, T. iarae.

However, Cernohorsky’s specimen could be easily in-

cluded in the range of geographic distribution and mor-

phological variation of T. plicatus. He also illustrated two

specimens of T. patagonicus as T. plicatus (F igures 8-10

in Houart, 1998).

Trophon patagonicus (d’Orbigny, 1839)

(Figures 43-65)

Murex patagonicus d’Orbigny, 1839: pl. 62, figs. 2-3; 1941: 452

non Fusus patagonicus Sowerby, 1846 (=Trophon).

Trophon necocheanus Ihering, 1907: 404, pl. 16, fig. 106.

Trophon laciniatus (Martyn).—Carcelles, 1946: 70-72, figs. 6

A, lo, 7 a@=cl, &.

Trophon plicatus (Lightfoot, 1786)—Rios, 1985: 88, pl. 31, fig.

386; 1994: 114, pl. SU, fig. 483; Houart, 1998: 130, figs.

8-10 non Lightfoot, 1786.

Description: Shell large (up to 72 mm), biconic,

heavy, chalky or glossy white, sometimes brownish grey,

opaque; protoconch smooth of 24 (~2.0 X 1.5 mm)

asymmetrical whorls; transition to teleoconch well de-

fined; teleoconch of 7 shouldered whorls, spire less than

¥ of total shell height. Spire angle about 70°; suture

impressed; aperture oval, interior glossy white; anterior

siphonal canal rather long for the genus (half height of

aperture), narrow, open; pseudoumbilicus deep and

widely open; outer lip reflexed. Axial ornamentation of

irregular strong, sharp lamellae covering entire whorl

surface, numbering 9-12 in the last whorl. Continuum

of morphologies present from completely smooth shells

with no axial ornamentation (Figures 53-56), to some

incipient lamellae along peripheral keel (Figures 48—50),

to strongly developed lamellae (Figures 43-45). Spiral

ornamentation missing, except on first teleoconch whorl

which bears 34 invariably present spiral cords (Figures

62-63). Irregular growth lines present throughout shell

surface. Some uncommon specimens have about three

weak greenish or dark spiral lines on the last three or

four whorls.

Shell ultrastructure arranged invariably of two layers;

innermost layer (25% shell thickness), composed of

crossed lamellar aragonite, outer layer very thick (75%

shell thickness) of amorphous calcite (Figure 61).

Operculum oval, with terminal nucleus. External sur-

face covered by concentric, irregular, growth lines. Inner

surface attachment area with two or three horseshoe-

shape scars.

Rachiglossan radula with rachidian teeth distinct, cen-

tral cusp thin, large; lateral cusps slightly shorter than

central cusp, sharp straight denticle in the upper third

of the internal edge of the lateral cusp, external edge of

lateral cusps smooth. Base of rachidian tooth sinuous,

sliding beneath base of next tooth. Marginal area with

single conspicuous cusp. Lateral teeth with single, long

THE NAUTILUS, Vol. 119, No. 2

and narrow cusp, attached to basal plate (Figures 64—

65).

Penchaszadeh (1976) described the egg capsules of T.

patagonicus [identified as T. laciniatus (Martyn) and T,

varians (d’Orbigny)].

Type Material: Four syntypes are housed in the Nat-

ural History Museum, London under the number

1854.12.4.538, from “Baie de San Blas, Patagonie”. One

of them is herein illustrated (Figures 43, 44).

Additional Material Examined: Puerto Quequén,

Buenos Aires province, 3 A (MLP 26309): Miramar,

Buenos Aires province, 1 A, 1 D (MLP 417); Necochea,

Buenos Aires province, 3 A, 30 m (AMNH 173640); Ne-

cochea, 1 A, 30 m (AMNH 181220); Puerto Quequén

(ANSP 236034 and 236032); 18 miles off Puerto Que-

quén, 4 D (MACN-In 20441): 37°20’ S, 56°50’ W, 4 D,

55 m (MACN-In 15104): Mar del Plata, Buenos Aires

province, Argentina (MACN-In 10289; 12902: 2 D, 45

m, 9361-51; 7 D, 11374; 11587; 11587-1; 4 D, 10249:

10320; 2 D, 10742; 11118; 3 D, 12066; 3 A, 25775; 9361-

53; 5 D, 10290; 10190; 10248; 12216): 36°25’ S, 54°38’

W, 54 m, 1 D (MACN-In 23426): 34°40’ S, 52°18’ W, 1

A, 100 m (MACN-In 23491); Fondos de Querandi,

Buenos Aires province, 5 A (MACN-In 14334); 25 miles

off Puerto Quequén, 22 D (MACN-In 21138); Fondos

de Querandf, 1 D (MACN-In 25774); Necochea, 2 D,

40-50 m (USNM 710024); Mar del Plata, 4 D, 1 D

(USNM 568240: USNM 346826): off Necochea, 1 A, 30

m (USNM 876123, Bledsoe collection); Mar del Plata,

1 D (USNM 363768); Uruguay, 1 D (USNM 346786);

1 D, 36°30’ S, 54°44’ W, 26 m (MACN-In 24259).

Distribution: This species is common on the mussel

banks off southern Uruguay and Buenos Aires province

in depths of 25-40 m.

Remarks: Trophon patagonicus is a variable species,

endemic to the littoral of Buenos Aires province and the

Uruguayan coast. The species has had quite a confusing

taxonomic history. Those specimens with well-developed

lamellae have been usually identified as T. plicatus, and

those with smooth shells and no ornamentation as T.

geversianus or T. varians. Specimens collected on the

same location were identified as two different species

according to the presence or absence of lamellae. How-

ever, some specimens (Figures 50-52) are clearly inter-

mediate forms, and no another anatomical feature seems

to separate them. There is no reason to consider them

as two different species. The whole lot of specimens

studied by Carcelles (1946) shows clearly that he con-

fused d’Orbigny’s species with T. plicatus. Such a mis-

take was also made by Rios (1985, 1994) and several

other authors. Trophon plicatus is a typical magellanic

species with a thinner and more cylindrical profile, me-

dium size (never reaching more than 50 mm high).

Specimens of T. plicatus here identified were never col-

lected at latitudes north of 45°S. Its protoconch is always

smooth and with fewer whorls, and the lamellae along

the keel usually develop into strong triangular projec-

G. Pastorino, 2005 Page 65

Figures 43-59. Trophon patagonicus (d’Orbigny, 1839). 43-44. Syntype BMNH 1854.12.4.538, Bahia San Blas, Argentina. 45-

46. MACN-In 36040, Puerto Quequén, Buenos Aires. 47-49. MLP 417, Miramar, Buenos Aires, in 54 m, 50-52. MACN-In 12066,

Mar del Plata, Buenos Aires. 53-56. MACN-In 36031, Puerto Quequén. 57-59. MACN-In 21047, “Carmen de Patagones?”,

Buenos Aires. Scale bar for all figures = 1 cm.

Page 66

THE NAUTILUS, Vol. 119, No.

Figures 60-65. Trophon patagonicus (d’Orbigny, 1839). 60. Penis, critical-point dried. Lateral and frontal view. Scale bar = 1000

wm. 61. Shell ultrastructure, fracture surface commarginal. Scale bar = 300 um. 62-63. MACN-In 11374, protoconch, three views,

Mar del Plata, Buenos Aires province. Scale bar for all figures

65. Lateral view of rachidian teeth. Scale bar = 50 ym.

tions approximately parallel to the coiling axis. The ap-

erture is larger and subcircular in T. patagonicus, very

different from the smaller and almost circular one on T.

plicatus. Trophon geversianus usually has strong spiral

ornamentation on the entire shell surface, a feature also

observed in T. varians.

Ihering (1907) described T. necocheanus based on

(Quaternary?) specimens collected at Necochea, Buenos

Aires province, Carmen de Patagones, Buenos Aires

600 xm. 64. Dorsal view of radular ribbon. Scale bar = 50 pm.

province, and Sierra Laziar, Santa Cruz province (“For

mation araucanienne”). Most of the type material is lost,

but the remaining types fall within the range of variation

of T. patagonicus. One of the specimens, from a lot of

three, with uncertain locality (MACN-In 21047, rela-

beled “Carmen de Patagones?”) is here illustrated (Fig-

ures 57, 59). This specimen was acquired through an

exchange with Museu Paulista, Sao Paulo, Brazil (where

Ihering worked for most of his professional life) and is

G. Pastorino, 2005

Page 67

part of the original type series. All of the specimens fit

well in the original description of T. patagonicus and

there is no doubt that is the same species described by

d Orbigny.

Sowerby II (2"¢ of name) described in 1846 Fusus pa-

tagonicus from the Tertiary of San Julian (Santa Cruz

province, Argentina), a species that clearly belongs in

Trophon. Therefore, as the two species are quite distinct,

dOrbignys name has priority over Sowerby’ (Griffin

and Pastorino, 2005).

Trophon acanthodes Watson, 1882

(Figures 66-78)

Trophon acanthodes Watson, 1882: 386; 1886: 166, pl. 10, fig.

6; Cemohorsky, 1977: 112, fig. 9 (holotype); Pain, 1980:

8, fig.; Rios, 1985: 88, pl. 31, fig. 388; Castellanos, 1986:

22, fig.; Castellanos and Landoni, 1993: 8, pl. 2, fig. 28.

Fusus acanthodes (Watson).—Carcelles, 1947: 12, pl. 2, figs. 1,

2; pl. 3, figs. 3, 4 (not figs. 5, 6 which is Coronium coron-

atum).

Pagodula acanthodes (Watson).—Kaicher, 1980: fig. 2599.

“Fusinus” acanthodes (Watson)—Calvo, 1987: 153, fig. 127.

“Trophon” acanthodes Watson.—Rios, 1994: 37, pl. 37, fig. 486.

Description: Shell large, about 125 mm high, thick,

slender in profile, chalky white; protoconch worn in all

specimens; teleoconch of 7 shouldered whorls; spire less

than % of total shell height. Spire angle about 45°; suture

impressed, subsutural shelf oblique, somewhat convex,

aperture small, subcircular, interior glossy white; ante-

rior siphonal canal very long (longer than aperture

height) deep, straight or curved, always open; outer lip

rather reflected, rounded, inner lip adpressed. Axial or-

namentation of 12-15 regularly spaced, axial lamellae,

slightly raised, almost attached along the entire whorl,

producing open, conspicuous, regularly spaced spines at

periphery; spines becoming more closely packed on last

whorl. Spiral omamentation of 4-5 weak rounded

threads on the lower part of the first whorls, becoming

more than 20 in last one. Growth lines present through-

out shell, producing wrinkly surface by intersection with

spirals.

Operculum elliptical, nucleus terminal, older speci-

mens tear-shaped. External surface covered by concen-

tric, irregular, growth lines. Inner surface with a con-

spicuous marginal rim; attachment area covering almost

the whole operculum but the rim; horseshoe-shaped

scars present.

Rachidian teeth wide (~130 j:m), central cusp large;

lateral cusps wider and shorter than central cusp, inner

edge with an almost obsolete denticle (particularly in

adult specimens), external edge with 5-6 denticles de-

creasing in size towards the edge. Base of rachidian

tooth curved, somewhat sinuous, sliding beneath base of

next tooth. Marginal area with single cusp. Lateral teeth

with single, long cusp, slightly attached basal plate.

Shell ultrastructure composed of two layers: inner-

most layer (55% of total thickness of shell) composed of

crossed lamellar aragonite, outer layer (45%) of amor-

phous calcite.

Animal as in other Trophon species but with some size

differences. Tentacles long and well defined, joined at

base; eyes deeply marked. Salivary and accessory salivary

glands as in T. plicatus. Osphradium a bit more than one

third of ctenidium length, asymmetrical, with about 112

leaflets, ctenidium with about 250 leaflets twice as large

as osphradium. Typical pleuroembolic proboscis, shorter

than in T. geversianus, valve of Leiblein also smaller;

gland of Leiblein large with a long blind duct. Radula

long; digestive tract with the usual loop on the left side

of the gland of Leiblein, before the duct to the gland.

Male and female organs similar to T. geversianus.

Type Material: BMNH 1887.2.9.568, holotype

50°8'30" S, 74°41’ W, 229 m (illustrated by Cernohorsky,

1977, fig. 9).

Additional Material Examined: 52°53’ S, 74°05’ W,

3 D, R/V ELranin Cruise 23, Sta. 1605, 1 April 1966,

522-544 m (USNM 901756): off Mar del Plata, 5 A,

January 1962 (MLP 96283); 52°41’ S, 74°35’ W, 1 D,

R/V ELTANIN Cruise 21, Sta. 290, 6 January 1966, 188—

247 m (USNM 870115): 51°56’ S, 56°39’ W, 1 D, B/V

ELTANIN Cruise 7, Sta. 557, 14 March 1963, 855-866 m

(USNM 870345): 52°52’ S, 75°18’ W, 1 D, B/V ELTANIN

Cruise 21, Sta. 288, 119-329 m (USNM 901758): 52°51’

S, 74°13’ W, 1 D, B/V ELTANIN Cruise 21, Sta. 291, 523—

539 m (USNM 901757); 52°53’ S, 74°05’ W, 2 D, R/V

ELTANIN Cruise 23, Sta. 1605, 1 April 1966, 522-544 m

(USNM 897615); 40°15’ S, 57°40’ W, 1 A (MACN

18425): 40°03’ S, 57°00’ W, 1 D, 50 fathoms (91.5 m)

(MACN 15699): 30 miles off Mar del Plata (ANSP

262989 and 236028): off Mar del Plata, 1 A, 2 D (MACN

17671 and 16449): 39°26’ S, 56°40’ W, 1 A, 1 D, 90 m

(MACN 17040): 39°02’ S, 56°46’ W, 1 A, 1 D; East of

Punta Médanos (39°-39°30' S), 4 D, 50 fathoms (91.5

m) (MACN 14386): 39°55’ S, 57°50’ W, 1 D, 51 fathoms

(93 m); 38°25’ S, 56°30’ W, 3 D (MACN 16798): 39°35’

S, 57°10’ W, 1 A, 1 D (MACN 18342): 39°50’ S, 57°18’

W, 4 A, 52 fathoms (95 m) (MACN 21741): 39°00’ S,

57°10’ W, 2 D, 45 fathoms (82 m) (MACN 15216):

37°15’ S, 54°50’ W, 1 A, R/V ALDEBARAN, March 2000,

commercial otter trawl, 111 m (MACN-In 36032);

36°37’ S, 54°14’ W, 1 A, B/V ALDEBARAN, March 2000,

otter trawl, 104 m (MACN-In 36031); 39°02’, 57°02’ W,

2D, 46 fathoms (84.1 m) (MACN-In 25118); 37°35’ S,

54°55’ W, 1 D, 105 fathoms (192 m) (MACN-In 25165-

2),

Distribution: Off Rio Grande do Sul state in Brazil

(Rios, 1994), Uruguay, to Tierra del Fuego, Argentina.

Remarks: The shell of Trophon acanthodes is some-

what similar to that of Coronium coronatum, which in fact

could be granted that generic position. The operculum

and the radula are different in both species. The oper-

culum is somewhat triangular in Coronium with a weak

rim instead of the characteristic thicker one of the Tro-

phon species. The radula of Trophon acanthodes presents

the intermediate cusp between the central and the lateral

Page 68 THE NAUTILUS, Vol. 119, No. 2

Figures 66-76. Trophon acanthodes Watson, 1882. 66-68. MACN-In 25118, 39°02’, 57°02’ W, 46 fathoms (84.1 m). 69-71.

MACN-In 36031, 37°15’ S, 54°50’ W in 111 m. 72-73. MLP 26283, Mar del Plata. Scale bar for all shells = 1 cm.74. Two views

of the operculum of the specimen in Figures 69-71. Scale bar = 1 cm. 75. Ultrastructure of the shell. Scale bar = 200 um; detail,

large quadrangle. Scale bar = 50 wm. 76. Penis, critical-point dried (scale bar = 600 jm) with detail of the efferent conduct (scale

bar = 150 pm).

one of the rachidian teeth almost obsolete, while in Co- The protoconch in Coronium is multispiral and ex-

ronium coronatum it is very conspicuous. Castellanos tremely pointed, distinctive of the genus. All the studied

(1986) drew a stereotyped radula of T: acanthodes where specimens of T. acanthodes are worn; however, some of

denticles are wrongly placed on the margin of the rachi- them show the sinuated line of a typical Trophon pro-

dian tooth instead of the inner edge of the lateral cusp. toconch.

G. Pastorino, 2005

Page 69

Figures 77-78. Trophon acanthodes Watson, 1882. 77. MLP 26283, off Mar del Plata Radula of a large specimen, frontal view.

Scale bar = 50 pm. 78. Lateral view. Scale bar = 50 pm.

Trophon pelseneeri Smith, 1915

(Figures 79-93)

Trophon pelseneeri Smith, 1915: 92, pl. 2, figs. 6, 7; Rios, 1994:

115, pl. 38, fig. 484; Houart, 1991: 33.

Trophon sp.—Carcelles, 1944: 253.

Trophon orbignyi Carcelles, 1946: 81, pl. 12;

73, pl. 5, fig. 6

Castellanos, 1970:

Description: Shell small to medium in size (up to 35—

40 mm), fusiform, slender, chalky, pinkish with 2 weak,

diffuse reddish bands along the edge of last whorl la-

mellae; protoconch symmetrical, cylindrical, smooth, of

2% whorls; teleoconch of 7 rectangular whorls, spire 4

of total shell height. Spire angle about 45°, suture im-

pressed, subsutural shelf straight, aperture small, circu-

lar, interior glossy white; anterior siphonal canal long,

open, straight, with the tip adaxially curved, equal to

aperture height; umbilicus slightly open, sometimes only

a narrow slit; outer lip reflexed to form lamellae; inner

lip curved, with white adpressed callus. Axial ornamen-

tation of 9-11 regular, low lamellae, covering entire

whorl surface; lamellae forming a back-tumed spine at

periphery. Spiral ornamentation of 2 to 3 very weak

threads in first whorls becoming more than 7 in last one,

sometimes only visible along edge of last whorl lamellae.

Siphonal fasciolae slightly oblique, always present.

Operculum triangular (tear-shaped), with terminal nu-

cleus. External surface covered by irregular growth lines.

Inner surface attachment area with horseshoe-shaped

scars; glazed rim weak but present (Figure 88).

Radula rachiglossate with rachidian teeth wider than

high, central cusp thin, in a different plane than laterals;

lateral cusps wider and shorter than central cusp, with

inner edge oblique; sharp straight denticle in the upper

third of the internal edge of the lateral cusp, external

edge with almost obsolete denticles numbering 3-4.

Base of rachidian tooth slightly sinuous, sliding beneath

base of next tooth. Marginal area with single conspicuous

cusp. Lateral teeth with single, long and narrow cusp,

slightly attached basal plate (Figures 92-93).

Type Material: [T. pelseneeri] BMNH 1915.4.18.276~7,

two syntypes from west of Malvinas Is. in 229 m (Figures

79-82) and [T. orbignyi] MACN-In 24421, Puerto Que-

quén, Buenos Aires, holotype (Figures 83-85) and 10

paratypes.

Additional Material Examined: Macaé, Rio de Ja-

neiro, Brazil, August 1969, 2 A, 55 m (USNM 846550);

Macaé, Rio de Janeiro, Brazil, 1 A, 30 m (AMNH

187586); Rio de Janeiro, Brazil, 1 A (AMNH 241045);

off Rio de Janeiro, Brazil (ANSP 289807); 34°48'7” S,

54°21'9” W, 1 A, R/V ALDEBARAN, Cruise 9901, Sta. 25,

27 January 1999, 25 m, with Piccard trawl (MNHNM

15400).

Distribution: Rio de Janeiro, Brazil, to Uruguay and

Buenos Aires province, Malvinas Is. Rios (1994) cited it

from dredgings from 55 to 225 m off the Brazilian coast.

Remarks: This is a rare species occasionally collected

by fishermen on the mussel banks off Buenos Aires

province. It was originally described from Sta. 38 of the

British Antarctic (“Terra Nova”) Expedition, west of

Malvinas (Falkland) Islands in 125 fathoms depth. How-

ever, together with this species, the author mentioned

several others not reported before or since from this

latitude, but which occur at Station 42 of the same ex-

pedition, off Rio de Janeiro. This leads to the supposi-

tion that the material from these stations could have

been mixed up (Scarabino, 2003: 199).

Trophon amettei Carcelles, 1946

(Figures 94-100)

Trophon amettei Carcelles, 1946: 84, fig. 13; Carcelles and Wil-

liamson, 1951: 287.

Description: Shell small in size (up to 30 mm), fusi-

Page 70 THE NAUTILUS, Vol. 119, No. 2

Figures 79-93. Trophon pelseneeri E. A. Smith, 1915. 79-80. BMNH 1915.4.18.276-7 syntype. 81-82. BMNH 1915.4.18.276-7,

other syntype. 83-85. T. orbignyi Carcelles, MACN-In 24421 holotype. 86-87. MNHNM 15400, B/I ALDEBARAN, 34°d48’7" S,

54°21'9" W, in 25 m. Scale bar = 1 cm. 88. Two views of the operculum of the specimen in Figures 86-87. Scale bar = 2 mm.

89-90. USNM 846550, apical and lateral view of the protoconch. Scale bars = 400 and 300 «um respectively. 91. Detail of the

ultrastructure of the shell. Scale bar = 100 pm. 92-93. Dorsal and lateral view of the radula of the specimen in Figures 86-87.

Scale bars = 30 wm.

G. Pastorino, 2005

Page 71

Figures 94-100.

form, biconic, chalky whitish; protoconch elongate,

smooth, of 214 whorls: teleoconch of 6 strongly sulk

dered whorls, spire less than % of the total shell height.

Spire angle about 45°, suture impressed; subsutural shelf

short but straight, aperture small, ovoid, interior glossy

white; anterior siphonal canal comparatively long, almost

same height as aperture, open and straight; umbilicus

closed, inner lip adpressed. Axial ornamentation of 8

regular concave lamellae per whorl. Lamellae growing

atecied to entire whorl surface and producing concave

spine along periphery. Spiral ornamentation of 2 cords

in first felboconch whorls that soon become obsolete on

subsequent whorls. Last whorl with 6-10 cords on lower

part. Shell ultrastructure composed of two layers, similar

to T. geversianus. Operculum and soft parts unknown.

Type Material: Holotype and 15 paratypes (MACN-

In 23810) all from 45°09’ S, 66°27’ W anchorage Res-

tinga Aristizébal, Chubut province, in 8 ecihorns: (11.28

m), rocky bottom associated with the calyptraeids Cre-

pidula cachimilla and Calyptraea pileolus.

Remarks: This is a rare species known only from the

type locality. It has not been found again. The regular

concave lamellae, are in fact, unusual for the genus. The

Trophon amettei Carcelles, 1946. 94-96. MACN-In 23810, holotype, 45°09’ S, 66°27’ W in 11.28 m. 97-98.

MACN-In 23810, paratype. Scale bar = 1 cm. 99. MACN-In 23810, ultrastructure of the shell. Scale bar = 30 pm. 100. MACN-

In 23810, protoconch, uncoated SEM picture. Scale bar = 300 wm.

protoconch resembles those usually found in the genus

Fuegotrophon, however the typical fimbriate spiral or-

namentation is absent. Trophon pelseneeri has a com-

parable profile, however it is easily segregated: where

the latter has lamellae forming a back-turned spine at

periphery, T. amettei presents a characteristic and

unique concave lamellae. In addition T. pelseneeri has a

slightly open umbilicus while in T. amettei it is invariably

closed. Examination of the radula may beget a new ge-

neric allocation for this species.

Trophon clenchi (Carcelles, 1953)

(Figures 101-114)

Murex clenchi Carcelles, 1953: 7, figs. 23-28; Castellanos,

1970: 80, pl. 5, fig. 7; Fair, 1976: 31, fig. 15: Vokes, 1992b:

24- Rios, 1994: 115.

Poirieria (Poirieria) clenchi (Carcelles).—Vokes, 1970: 18.

Description: Shell medium in size, delicate, thin,

chalky, translucent white; protoconch known only from

the holotype, asymmetrical, globose, of 2-2% whorls; te-

leoconch of 5 shouldered whorls: spire short, less than

¥% of total shell height. Spire angle about 45° (without

spines); suture impressed, subsutural shelf short,

Page 72

THE NAUTILUS, Vol. 119, No. 2

Figures 101-110. Trophon clenchi (Carcelles, 195:

107. MACN-In 36269, aproximately 45°10’ S, 37°20) ne bar = 1 cm. 108. Holotype, protoconch. Scale bar = 1000 xm. 109.

External and internal views of operculum of specimen in Figures 105-107. Scale bar = 0.5 cm 110. MACN-In 25146, paratype.

straight; aperture suboval, interior glossy white; anterior

siphonal canal very long (longer than the aperture

height), deep and slightly curved in the beginning and

then straight, open but narrow; outer lip sharp, inner lip

somewhat" protruding,

7-9 regular axial lamellae growing across entire whorl

surface, but attached to shell producing open, long and

regular lg spaced spines along periphery. Spiral ornamen-

tation of 4—5 rounded cords on lower half of first whorls,

adpressed. Axial ornamentation of

. 101-104. MACN-In 25146, holotype, 38°24’ S, 55°36’ W in 89.61 m. 105-

becoming more than 20 in the last. Delicate growth lines

present on entire shell surface.

Shell ultrastructure composed of two layers following

the common pattern of the genus: innermost layer (40%

of total thickness of shell) composed of crossed lamellar

aragonite, outer layer (50%) of amorphous calcite.

Sometimes, depending on the fracture mode, a basal

aragonitic layer can be observed.

Aadulee rachiglossate, rachidian teeth with three me-

G. Pastorino, 2005

icy)

Page

Figures 111-114. Trophon clenchi (Carcelles, 1953). 111. USNM 901774, 51°58’ S, 56°38’ W, R/V ELTANIN Cruise 7, Sta. 558,

646-845 m, ultrastructure of the shell. Scale bar = 100 ym. 112. Poirieria zelandica, ultrastructure, commarginal fracture surface.

Seale bar = 100 wm. 113. Trophon clenchi MACN-In 36269. Lateral view of radula ribbon. Scale bar =

view of radular ribbon. Scale bar = 50 ym.

dian cusps, the central one the larger and the lateral

ones with a denticle, attached to the upper third of the

interior margin of the lateral cusp; external margin with

2-3 obsolete denticles. Rachidian base sinuous, with the

base offset under the proximal tooth. Marginal cusps sin-

gle, never bifid. Lateral teeth curved, thin, with attach-

ment area also thin (Figures 113-114).

Operculum triangular or suboval tear-shaped, attach-

ment area elliptical, with horseshoe shape scars.

Type Material: 38°24’ S, 55°36’ W, off Mar del Plata,

in 89.6 m, holotype and paratype (MACN-In 25146).

Additional Material Examined: 51°58’ S, 56°38’ W,

3 D, R/V ELTANIN Cruise 7, Sta. 558, 646-845 m

(USNM 901774), 41°51’ S, 57°34’ W, collected by Uru-

guayan fishermen, June 2002, 1062 m, 1A, MACN-In

36270; approximately 45°10’ S, 57°20’ Uruguayan fish-

ermen, 1 A, MACN-In 36269; 54°50’ S, 63°50.5’ W, 2.5

miles south Punta Fallows, Isla de los Estados, Tierra

del Fuego, 1 A, R/V Hero Cruise 715, Sta. 715/879, 28

October 1971, in 342-353 m (LACM 71-331).

100 pm. 114. Dorsal

Distribution: Known from off Buenos Aires province,

Patagonia and Isla de los Estados, in 90-1050 m depth.

Remarks: Carcelles (1953) described Murex clenchi

from two shells he received from the crew of the ArA

Bania BLANca, an Argentine Navy ship that occasionally

collected material for the collection of the Museo Ar-

gentino de Ciencias Naturales (MACN). Both holotype

and paratype were collected from the continental shelf

off Buenos Aires province. This material remained

housed at the MACN until it was sent on loan and it

appears to have been lost for almost 30 years (see Cas-

tellanos, 1986). The return of the material to MACN

made the type material again available for studies. Per-

haps because of these facts: the ordinary quality of the

illustration and the absence of soft parts, the species was

always reluctantly considered as the southernmost rep-

resentative of the genus Murex.

E. H. Vokes, in a fundamental paper published in

1970, stated that Murex clenchi belongs in the genus

Poirieria sensu stricto. Rios (1994) considered M. clenchi

THE NAUTILUS, Vol. 119, No. 2

as an anomalous specimen of Trophon acanthodes Wat-

son, a species that slightly resembles some specimens of

M. clenchi (e.g., the paratype).

The morphology of the shell as well as the radula and

penis of Murex clenchi allow the allocation in the genus

Trophon. In fact, this was already suggested by E. H.

Vokes (1992b). After studying the material of T. acan-

thodes housed at the MACN she proposed that M. clen-

chi belongs to Trophon sensu lato and not to Poirieria.

It bears no relationship with Poirieria despite some ap-

parent shell similarity. As it is shown in Figure 112, Poir-

ieria zelandica, type species of Poirieria, has no calcitic

layer on the shell but a thick aragonitic one instead (Fig-

ure 112a). In contrast, the entire group of Patagonian

Trophon (T. clenchi in particular), shows different de-

grees of development of the typical calcitic layer on the

distinct species.

The range of the very few lots studied falls within the

geographic distribution of other species of Trophon. Tro-

phon clenchi was known from two quite different geo-

graphic and bathymetric areas: off Buenos Aires prov-

ince (in about 90 m) and the slope off Patagonia. A new

lot is recorded here from Isla de los Estados, which sig-

nificantly increases the range of distribution of the spe-

cies. It is possible that like other Patagonian species of

the genus, T. clenchi would be angus to hard bot-

TONING, thus hampering the collection of material. Its fra-

gility, size, and inadequate sampling of the area adds

other reasons for the scarcity of records. It is suggested

that these variables accounts for the disjunt recorded

distribution of this species. In fact, this species started

to be repeatedly collected recently, when the fishery of

the Patagonian toothfish (Dissostichus eleginoides) on

the Argentinean slope provided the opportunity to catch

accidentally entangled specimens (F. Scarabino, pers.

comm. ).

Trophon wilhelmensis Ramirez-Bohme, 1981

(Figures 115-119)

Trophon (Enixotrophon) wilhelmensis Ramirez-Bohme, 1981:

6, fig. la, b.

Description: Shell of medium size, about 52 mm

high, slender in profile, chalky white; protoconch un-

known; teleoconch of 6 shouldered whorls; spire less

than % of total shell height. Spire angle less than 45°;

suture impressed, subsutural shelf straight, aperture

small, subcircular, interior glossy white; anterior siphonal

canal very long (longer than aperture height) deep,

straight or curved, always open; outer lip rather reflexed,

rounded, inner lip adpressed. Axial ornamentation of

12-15 regularly spaced, axial lamellae, slightly raised, al-

most attached along the entire whorl, producing open,

conspicuous, regularly spaced spines at periphery; spines

becoming more closely packed on last whorl. Spiral or-

namentation of 4—5 weak rounded threads on the lower

part of the first whorls, becoming more than 20 in last

one. Growth lines present throughout shell, producing

wrinkly surface by intersection with spirals.

Operculum elliptical, nucleus terminal. External sur-

face covered by growth lines. Inner surface with mar-

ginal rim; attachment area, horseshoe-shaped scars pres-

ent.

Rachidian teeth of about 90 zm wide, central cusp

large; lateral cusps shorter than central cusp, inner edge

with conspicuous denticle, external edge with 5 denticles

of equal size. Base of rachidian tooth straight, somewhat

sinuous, sliding beneath base of next tooth. Marginal

area with single cusp. Lateral teeth thin with single, long

cusp, slightly attached basal plate.

Type Material: Holotype in MNHN (unnumbered),

41°51’ S, 74°30'5” W West of Chiloé Island in 250 m.

Additional Material Examined: Boca del Guafo,

43°39'36" S, 73°51'11” W, southern Chile, 6 July 2002,

1 A, 200 m (MNHN unnumbered).

Distribution: Known only from the holotype and an-

other lot, both from the same area and depth, around

the Chiloe Is., Chile.

Remarks: Trophon wilhelmensis was recently de-

scribed from Chile with no mention of the similarity

with T. acanthodes. Both species are actually quite sim-

ilar. The species, known only from the holotype and an-

other specimen, may be distinguished from T: acantho-

des mainly by the upturned spines (compare Figures 66—

73 with Figures 115-117). In addition, slight differences

in the morphology of the rachidian teeth allow for fur-

ther differentiation of the two species. Notwithstanding,

future studies of specimens from intermediate localities

could demonstrate that these represent but a single spe-

cies.

Trophon bahamondei McLean and Andrade, 1982

(Figures 120-125)

Trophon bahamondei McLean and Andrade, 1982: 10, figs. 24—

25.

Description: Shell of medium size, up to 50 mm,

slender, with narrow profile, chalky or creamy white;

protoconch unknown (worn in all specimens); teleo-

conch of 6 shouldered whorls; spire less than % of total

shell height. Spire angle albowt 45°; suture impressed,

substan shelf well hence: straight; aperture subcir-

cular, interior chalky white; anterior siphonal canal very

long (equal to aperture height), narrow, and curved,

open; outer lip sharp, rounded, inner lip adpressed. Axial

ornamentation of 11-13 regular axial lamellae growing

on entire whorl surface, but attached to the shell and

producing open, short and regularly spaced spines along

periphery. Spiral ormamentation lacking. Growth lines

present throughout shell.

Operculum elliptical, nucleus terminal. External sur-

face covered by concentric, irregular growth lines. Inner

surface attachment area reaching upper side or center,

with horseshoe-shaped scars (Figure 123).

Radula with rachidian teeth very wide (~114 pm),

with narrow base, central cusp thin, large; lateral cusps

G. Pastorino, 2005

Page 75

ae as

ay. ee "1

on ey "

Figures 115-119. Trophon wilhelmensis Ramirez-Bohme, 1981. 115-117. MNHNS unnumbered, Boca del Guafo, 43°39'36" S,

73°51'11" W, southern Chile, 6 July 2002, 200 m. Scale bar =

1 cm. 118-119. Radula of the specimen on Figures 115-117, 118.

Frontal view. Scale bar = 50 wm. 119. Lateral view. Scale bar = 40 pm.

wider and shorter than central cusp, inner edge with

sharp, curved denticle, external edge with denticles de-

creasing in size toward lateral edge where they disap-

pear. Base of rachidian tooth sinuous, inserted in part

under base of subsequent tooth. Marginal area with sin-

gle conspicuous cusp. Lateral teeth with single, long and

very narrow cusp, slightly attached basal plate. Central

cusps of rachidian teeth curved back in lateral view (Fig-

ures 124—1925).

Animal very similar to T. plicatus. Osphradium less

than half ctenidium length, asymmetrical, with 60-70

leaflets. Ctenidium with 140 leaflets less than two times

larger than those of osphradium. Tentacles well defined;

eyes deeply marked. Same type of accessory salivary

glands, twisted and non-compact.

Male unknown, all specimens studied were females.

Female similar to T. geversianus.

Type Material: Holotype from off Pichilemu, Chile,

34°27’ S, 340 m (LACM 1982, but apparently on loan)

and two paratypes, 34°27’ S, 71°54’ W, 25 May 1976,

200-450 m (USNM 784739).

Additional Material Examined: 31°56’ S, 71°54’ W,

off Los Vilos, Chile, 2 A, 8 March 1977, 240-350 m

(LACM 72491); off Playa Blanca, Coquimbo, Chile, 400

m (ANSP 291065); off Coquimbo, Chile (LACM 75-88).

Distribution: This is a deep water Trophon known

from latitudes 30° to 34° S off the coast of Chile.

Remarks: Trophon bahamondei is apparently a very

Page 76

THE NAUTILUS, Vol. 119, No. 2

Figures 120-125. Trophon bahamondei McLean and Andrade, 1982. 120-122. Paratype USNM 784739, 34°27’ S, 71°54’ W,

Pichilemu, Chile, in 200-450 m. Scale bar = 1 cm. 123. Operculum, external (right) and internal (left) views. Scale bar = 1 cm.

124. Dorsal view of radular ribbon. Scale bar = 30 wm. 125. Lateral view of rachidian teeth. Scale bar = 30 pm.

consistent species from the morphological standpoint. It

was only recently described despite its apparently wide-

spread presence in shrimp trawls. The deeper water

habitat of species (more than 200 m) probably rendered

the species less accessible in the past.

General morphology of shell and gross anatomy shows

at first glance some similarities with those of T. plicatus;

however close examination of shell and radula confirms

the presence of a different species.

Trophon parodizi new species

(Figures 126-137, Table 1)

Description: Shell medium in size (up to 23 mm),

very thin, chalky; protoconch smooth, of 1% (1.41 X

1.39) very asymmetrical whorls; transition to teleoconch

well defined: teleoconch of 4 convex whorls, spire less

than ¥% total shell height. Spire angle about 40°; suture

impressed; aperture subov oid, interior glossy white; an-

terior siphonal canal medium in size (less than half

height of aperture), narrow, open; umbilicus absent; out-

er lip rounded. Axial ornamentation of irregular, low,

rounded ridges occupying entire whorl surface, num-

bering up to 9 in early whorls, but vanishing and un-

dilewe on last whorl. Spiral ornamentation poorly devel-

oped or almost smooth, when present consisting of ob-

solete, weak cords mostly developed on last whorl. l-

regular growth lines present throughout shell.

‘Shell “Tilrectranettnnie composed of two layers; inner-

most layer thin (35% of shell thickness), composed of

colabrally aligned crossed lamellar aragonite, outer layer

thick (65% of shell thickness) with amorphous calcite.

Operculum oval, with terminal nucleus. External sur-

face covered by concentric, irregular, growth lines. Inner

surface attachment area with two or three horseshoe-

shape scars. Animal unknown.

Radula rachiglossan with rachidian teeth distinctive,

central cusp ‘livin. large; lateral cusps almost same size

as central cusp, sharp denticle on upper third, irregular

external denticles present. Base of rachidian tooth sin-

G. Pastorino, 2005 Page 77

Figures 126-137. Trophon parodizi new species. 126-128. USNM 896397, holotype, 54°56’ S, 65°03’ W, in 229-265 m. 129-

130. USNM 896397, paratype, coated with ammonium chloride. Scale bar = 1 cm. 131. Ultrastructure, fracture surface commar-

ginal. Scale bar = 20 wm. 132. USNM 896397, operculum, external view. Scale bar = 0.5 cm. 133-135. Protoconch of the

paratype, three views. Scale bar for all figures = 400 wm. 136. Dorsal view of radular ribbon. Scale bar = 40 xm. 137. Lateral

view of rachidian teeth. Scale bar = 40 pm.

Page 75

THE NAUTILUS, Vol. 119, No. 2

Figures 138-144.

uous, sliding beneath base of next tooth. Marginal area

with a conspicuous cusp. Lateral teeth with single, long

cusps and narrow, slightly attached basal plate.

Type Material: Holotype and one paratype from

54°56° S, 65°03° W, Le Maire Strait, R/V ELTANIN Cruise

11, Sta. 969, 10 February 1964, 229-265 m, Blake trawl

(USNM 896397) (Map 1).

Material Examined: Only holotype and paratype.

Distribution: Known only from the type locality.

Etymology: This species is named after Juan José Par-

odiz, one of the pioneers of malacology in Argentina,

Curator Emeritus at the Carnegie Museum, Pittsburgh,

and a good friend.

Remarks: Trophon parodizi is an unusual species

within Trophon. The small, very thin shell, almost com-

pletely smooth and devoid of lamellae, is an exception

for the Patagonian Trophon. However, the radula and

protoconch match those typical of the genus. The rad-

ular morphology resembles that of T. plic atus where the

denticles of the lateral cusp of the rachidian teeth are

placed along the entire external edge of the lateral cusp.

In T. parodizi the lateral cusp is thinner and the denti-

cles are more clearly separated from it and placed mainly

along the marginal area as in T. bahamondei. In addi-

tion—a character also present in T. bahamondei—most

internal denticle lies against the lateral cusp and is clear-

ly differentiated from the rest.

“Trophon” malvinarum Strebel 1908. 138-139. NHRM 1041, holotype, 52°29’ S, 60°36’ W. 140-142. MACN-

In 23944-2, 55°07’ S, 66°33’ W. 143-144. Protoconch. Scale bar = 400 wm.

“Trophon” malvinarum Strebel, 1908

(Figures 138-144)

Trophon malvinarum Strebel, 1908: 44, figs. 16 a—c; Carcelles

and Williamson, 1951: 289; Castellanos and Landoni,

1993: 10, pl. 2, fig, 29.

Description: Shell small, up to 30 mm, thick, fusi-

form, profile slender, chalky, whitish; protoconch glo-

bose, somewhat cylindrical, with two whorls: teleoconch

of 4% shouldered whorls, spire less than ¥% of total shell

height. Spire angle about 45°; suture impressed; subsu-

Aural shelf alberdlley oblique, aperture subquadrate, an-

terior siphonal canal moderately long (same height as

aperture); umbilicus closed, outer ‘lip polygonal to

rounded; inner lip slightly curved, adpressed. Axial or-

namentation of 8-9 regular, distinct low varices, which

never develop into all -fledged lamellae. Spiral orna-

mentation of about 3 cords, filling interspaces between

consecutive varices, and beginning at periphery of

whorls.

Radula and anatomy unknown.

Table 1. Measurements of the type specimens of Trophon

parodizi new species in mm.

Species Length Width Whorls

T. parodizi

Holotype 93 ll 4

Paratype 21.1 10.6 4

G. Pastorino, 2005

San Matias

Golfo

\ san Jorge

~ .Malvinas Is.

* hoe

Figure 145. Map showing the type locality of T. parodizi

new species ( @).

Type Material: 52°29’ S, 60°36’ W, West Falkland

(Malvinas) Is., Svenska Sudpolar Expedition, 11 Septem-

ber 1902, Sta. 58, 197 m (NHRM 1041).

Additional Material Examined: 55°07’ S, 66°33’ W,

1 D, 83 m (MACN-In 23944-2).

Distribution: Around Malvinas Is. and the Magellanic

region.

Remarks: “Trophon” malvinarum is a rare species

known from only one specimen other than the holotype.

It was originally described in the genus Trophon; how-

ever, the protoconch is somewhat shorter and symmet-

Page 79

rical and the shell never develops true lamellae but low

varices. It probably could be better assigned in the genus

Urosalpinx. Assessment of its real affinities must wait

until specimens with soft parts preserved can be studied.

CONCLUDING REMARKS

The geographic distribution of the species of the genus

Trophon sensu stricto is restricted to the southwestern

Atlantic and the southeastern Pacific Oceans. This dis-

tribution is certainly a consequence of their larval biol-

ogy. Planktonic larval development is unknown to occur

in the genus. Moreover, the larvae do not need to move

any significant distance either for feeding or reproduc-

tive purposes—those from shallow water habitats live on

mussel banks on which they feed and mate. Such a con-

dition is reflected in the enormous morphological vari-

ation shown by the shells of the different species, par-

ticularly in T. geversianus. While most of the species are

well known, some others were only collected in a single

location (e.g., T. amettei) and therefore their range of

variation remains to be known.

Radular and anatomical features suggest that the en-

tire Trophon group from Patagonia is very homoge-

neous. All the species included in this genus have several

common radular features. The most remarkable are: the

intermediate denticle attached to the upper third of the

internal edge of the lateral cusp of the rachidian teeth;

a single marginal denticle in the external edge of the

base of the rachidian teeth; the attachment area of the

marginal teeth are always (no exceptions known) narrow,

thin, with the free part of same thickness, and the cen-

tral cusp of the rachidian is always thin and larger than

the laterals (see also Pastorino, 2002).

The Antarctic species so far assigned to Trophon and

the boreal ones recently included in the genus Boreo-

trophon (see Egorov, 1993) have the inner denticle be-

tween central and lateral cusp of the rachidian teeth al-

ways free, attached to the base of the teeth. In addition,

most of the radulae of these northeastern Atlantic spe-

cies—according to Bouchet and Warén’s revision

(1985)—have a broad attachment of the marginal teeth.

Conchological features are so variable that I consid-

ered them as secondary. However, the protoconchs are

actually very different and allow for the division in at

least two clearly defined groups. There is no ornamen-

tation on the protoconchs of Patagonian representatives

of Trophon, whereas most of the boreal species of Bor-

eotrophon have a delicate pattern of irregular threads.

The Antarctic species have also no ornamentation in the

protoconch with only one exception: T. scotianus Powell,

1951 which has apparently the same pattern observed in

North Atlantic Boreotrophon species.

There are several anatomical features that character-

ize the group of species living along the South American

coast. The accessory salivary glands, when known, are

usually tubular; the esophagus produces a typical esoph-

ageal loop after the valve of Leiblein and posteriorly

runs appressed to the left side of the gland of Leiblein;

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THE NAUTILUS, Vol. 119, No. 2

the esophageal glands in the mid-esophagus are incon-

spicuous, not externally visible; and finally, the penises

are always dorso-ventrally flattened, with a large papilla

and a simple vas deferens either closed by the overlap-

ping sides of the penis or open.

The features mentioned above and the geological sto-

ry of the two areas, Antartica and Patagonia, allow for

the clear differentiation of these two groups.

ACKNOWLEDGMENTS

I thank the following people for making available spec-

imens and type material for study or advice when re-

quested: P. Greenhall and T. Nickens (USNM); K. Way

(BMNH), D. Reid (BMNH):; A. Warén (NHRM); P.

Bouchet and V. Heros (MNHN); J. H. McLean and L.

Groves (LACM); P. Mikkelsen (AMNH); G. Rosenberg

and P. Callomon (ANSP); B. Sirenko (ZIL); O. Galvez

Herrera (MNHNS); C. Osorio (Santiago, Chile); F. Scar-

abino (DINARA) and C. Ituarte (MLP). P. Lozouet

(MNHN) took the photographs of specimens from Paris.

S. Horta (DINARA) and H. Racz-Lorenz (Montevideo)

kindly provided some of the specimens of Trophon clen-

chi that allowed part of the present work. Emily Vokes

(Tulane University) and Bruce Marshall (Museum of

New Zealand Te Papa Tongarewa, Wellington) helped

with excellent reviews that considerably improved on the

original manuscript. Finally special thanks to M. G. Har-

asewych (USNM), who guided this work from the be-

ginning, and to M. Gala for good advice.

Part of this study was condinctied! during a Postdoctoral

Fellowship granted by the Consejo Nacional de Inves-

tigaciones Cientificas y Técnicas (CONICET), Argenti-

na, to support my tenure at the National Museum of

Natural History, Smithsonian Institution, Washington,

DC. The study was also supported in part by a Resear

Award from the NSF-USAP United States Antarctic

Program [Contract No. OPP-9509761], a Grant Award

from Conchologists of America and the Walter E. Sage

Memorial Award, in addition to the projects PICTs 02-

01795 and 03-14419 from the National Agency for Sci-

entific and Technical Promotion, Argentina.

LITERATURE CITED

Adams, H., and A. Adams. 1853-1854. The Genera of Recent

Mollusca, arranged according to their organization. Vol-

ume 1. John van Voorst, Paternoster Row, London, 484

pp: [pp. 1-156, 1853; pp. 257-484, 1854].

Aguirre, M. L. 1993. Type specimens of Quaternary marine

gastropods from Argentina. Ameghiniana 30: 23-38.

Bouchet, P., and A. Warén. 1985. Revision of the northeast

Atlantic bathyal and abyssal Neogastropoda excluding Tur-

ridae (Mollusca, Gastropoda). Bolletino Malacologico,

supplemento 2: 123-296.

Calvo, I. S. 1987. Radulas de Gastrépodes Marinhos Brasilei-

ros. Editora da Fundacgao Universidade do Rio Grande,

Rio Grande, 201 pp.

Carcelles, A. 1943. Observaciones sobre Trophon varians

d'Orbigny. Notas del Museo de La Plata, Zoologia 8(72):

431437.

Carcelles, A. 1946. Observaciones sobre algunas especies ac-

tuales y fésiles de Trophon de la Reptblica Argentina. No-

tas del Museo de La Plata, Zoologia 11(93): 59-89.

Carcelles, A. 1947. Notas sobre algunos gastropodos marinos

del Uruguay y la Argentina. I-VI. Comunicaciones Zool-

égicas del Museo de Historia Natural de Montevideo

2(40): 1-27.

Carcelles, A. 1953. Nuevas especies de gastropodos marinos de

las Reptiblicas Oriental del Uruguay y Argentina. Comun-

icaciones Zoolégicas del Museo de Historia Natural de

Montevideo 4(70): 1-18.

Carcelles, A. and S. Williamson. 1951. Catalogo de los molus-

cos marinos de la provincia magallanica. Revista del In-

stituto Nacional de Investigacién de las Ciencias Naturales

2, Ciencias Zoolégicas: 225-383.

Castellanos, Z. J. A. de. 1970. Catalogo de los moluscos mari-

nos bonaerenses. Anales de la Comisién de Investiga-

ciones Cientificas dé la provincia de Buenos Aires 8: 9—

365.

Castellanos, Z. A. de. 1986. Aclaracién sobre Trophon acan-

thodes Watson: 1883 (Mollusca Gastropoda). Neotrépica

32, (87): 22.

Castellanos, Z. J. A. de. and N. Landoni. 1993. Catélogo des-

criptivo de la malacofauna marina magallanica 9. Neogas-

tropoda: Muricidae y Thaisidae. Comisién de Investiga-

ciones Cientificas de la Provincia de Buenos Aires, 26 pp.

Castellanos, Z. J. A. de., E. Rolén and S. Bartolotta. 1987.

Nuevos micromoluscos de la plataforma inferior Argentina

y talud superior (Moll. Gastropoda). Revista del Museo de

La Plata, Seccion Zoologia 14(156): 93-102.

Cernohorsky, W. O. 1977. The taxonomy of some southem

ocean mollusca (Gastropoda) mainly antarctic and subant-

arctic. Records of the Auckland Institute and Museum 14:

105-119.

Chemnitz, J. H. 1780. Neues systematisches Conchylien-Cab-

inet / Geordnet und beschrieben von Friedrich Heinrich

Wilhelm Martini und unter dessen Aufsicht nach der Na-

tur gezeichnet und mit lebendigen Farben erleuchtet.

Raspe, Niiremberg. Vol. 4, 344 pp. pls. 122-159.

Cossmann, M. 1903. Essais de Paléoconchologie comparée. F.

R. de Rudeval, Paris, 151 pp.

Cristofori, J. D., and G. Jan. 1832. Catalogus in IV sectiones

divisus rerum naturalium in Musaeo exstantium / J. de

Cristofori et G. Jan ... complectens adumbrationem or-

yctognosiae et geognosiae atque prodromum faunae et flo-

rae italiae superioris. Mediolani, Parmae, Sectio 1, 1-8 pp.

Dance, P. 1962. The authorship of the Portland Catalogue

(1786). Journal of the Society of Bibliography of Natural

History 4(1): 30-34.

Dance, P. 1966. Shell Collecting: An Illustrated History. Berke-

ley, University of California Press, Berkeley, 305 pp.

Dell, R. K. 1971. The marine mollusca of the Royal Society

Expedition to southern Chile, 1958-1959. Records of the

Dominion Museum 7(17): 155-233

Dell, R. K. 1990. Antarctic Mollusca with special reference to

the fauna of the Ross Sea. Bulletin of the Royal Society

of New Zealand 27: 1-311.

dOrbigny, A. D. 1834-1847. Mollusques. In: Bertrand, C.P,,

(ed., Voyage dans Amerique Meridionale (Le Bresil, La

Republique Orientale de LUruguay, La Republique Ar

gentine, La Patagonie, La Republique du Chili, La Re-

publique de Bolivia, La Republique du Perou), execute

G. Pastorino, 2005

Page 81

pendant les annees 1826, 1827, 1828, 1829, 1830, 1831,

1832 et 1833. Chez Ve. Levrault, Paris, 5(3) 758 pp.

Egoroy, R. 1993. Trophoninae (Muricidae) of Russian and ad-

jacent waters. Ruthenica Supplement: 11-49.

Fair, R. H. 1976. The Murex Book: an illustrated catalogue of

Recent Muricidae (Muricinae, Muricopsinae, Ocenebri-

nae). Ruth H. Fair, Honolulu, Hawaii, 138 pp.

Favanne Montcervelle, J. D. 1780. La conchyliologie, ou his-

toire naturelle des coquilles de mer, d’eau douce, terres-

tres et fossiles; avec un traite de la Zoomorphose, ou rep-

resentation des animaux qui les habitent: Ouvrage dans

lequel on trouve une nouvelle Methode de les diviser. De

Bure, G. Paris, 2, 848 pp

Gmelin, J. F. 1791. Caroli a Linné Systema Naturae per Regna

Tria Naturae, Secundum Classes, Ordines, Genera, Spe-

cies, cum characteribus, differentiis, synonymis, locis. Ed-

itio decima tertia, aucta, reformata, cura J. F. Gmelin.

Georg. Emanuel. Beer, Leipzig, 1, part 6 Vermes, pp.

3021-3910.

Gould, A. A. 1852-1861. Atlas. Mollusca and Shells, p. 512,

United States Exploring Expedition. During the years

1838, 1839, 1840, 1041, 1842. Under the command of

Charles Wilkes, U. S. N. Volume 12. C. Sherman and Son,

Philadelphia.

Gray, J. E. 1839. Molluscous Animals. In: The Zoology of Cap-

tain Beechey’s Voyage; compiled from the collections and

notes made by Captain Beechey, the officers and naturalist

of the expedition, During a voyage to the pacific and Beh-

ring’s Straits performed in his Majesty's ship Blossom, un-

der the command of Captain F. W. Beechey, in the years

1825, 26, 27 and 28. H. G. Bohn, London 1: 103-142.

Griffin, M. and G. Pastorino. 2005. The genus Trophon Mont-

fort, 1810 (Gastropoda: Muricidae) in the Tertiary of Pat-

agonia. Journal of Paleontology 79: 296-311.

Hanley, S. 1856. Index Testaceologicus, an illustrated catalogue

of British and foreign shells ... by W. Wood. A new and

entirely revised edition. Willis and Sotheran, London. xx

+ 234 pp

Harasewych, M. G. 1984. Comparative anatomy of four prim-

itive muricacean gastropods. Implications for Trophoninae

phylogeny. American Malacological Bulletin 3: 11-26.

Houart, R. 1991. The southeastern brazilian muricidae col-

lected by RV Marion-Dufresne in 1987, with the descrip- _

tion of three new species. The Nautilus 105: 26-37.

Houart, R. 1998. Description of Trophon iarae n. sp., a new

muricid from southern West Atlantic (Gastropoda: Muri-

cidae) with illustration of related species. Apex 13: 127-—

130.

Houart, R. 2003. Description of three new species of Trophon

s. |. Montfort, 1810 (Gstropoda: Muricidae) from Chile.

Novapex 4: 101-110.

Hupé, L. H. 1854. Mollusca, p. 499. In: C. Gay (ed.), Historia

Fisica y Politica de Chile. Zoologia 8. Maulde y Renou,

Paris and Santiago.

International Commission on Zoological Nomenclature

(ICZN). 1944. Opinion 184. On the status of names first

published in volumes 1 to 11 of Martini (F. H. W.) and

Chemnitz (J. H.), Neues systematisches Conchylien-Cab-

inet, Ntirnberg, 1769-1795. Opinions and Declarations

rendered by the ICZN 3(3): 25-36.

International Commission on Zoological Nomenclature

(ICZN). 1957. Opinion 456. Rejection of the work by

Thomas Martyn published in 1784 with the title “The Uni-

versal Conchologist” as a work which does not comply

with the requirements of Article 25 of the “regles” and

which therefore possesses no status in zoological nomen-

clature and rejection also of a proposal that the forgoing

work should be validated under the plenary powers. Opin-

ions and Declarations of the ICZN 15(22): 395-410.

Thering, H. von 1907. Les Mollusques fossiles du Tertiaire et

du Cretace superieur de lArgentine. Anales del Museo

Nacional de Buenos Aires (3 V7: 1-611.

Knorr, G. W. 1757-1773. Verniigen der Augen und des Ge-

miiths, in Vorstellung einer allgemeinen Sammlung von

Muscheln und andern Geshépfen, welche im Meer ge-

funden werden. Niirnberg. 1, 40 pp., 30 pls. (1757); 2, 56

pp.. 30 pls. (1764); 3, 52 pp., 30 pls. ve 4, 48 pp., 30

pls. (1769); 5, 46 pp., 30 pls. (1771); 6, 18 pp., 40 pls.

(1773).

Knorr, G. W. 1760-1773. Les délices des yeux et de lespirit,

ou collection générale des différentes espéces de coquil-

lages que la mer renferme. Nuremberg. 1, 52 pp., 30 pls.

(1760); 2, 65 pp., 30 pls. (1765); 3, 55 pp., 30 pls. (1768);

4, 54 pp., 30 pls. (1770); 5, 48 pp., 30 pls. (1771); 6, 76

pp:, 40 pls. (1773).

Kobelt, W. 1878. Die geschwiinzten und bewehrten Purpur-

schnecken (Murex, Ranella, Tritoniwm, Trophon, Hindsia).

In: Abbildungen nach der Natur mit Beschreibungen. Be-

gonnen von H. C. Kiister, fortgesetzt und beendet von W.

Kobelt, Systematisches Conchylien-Cabinet von Martini

und Chemnitz. Neu herausgegeben und vervollstandigt

von H. C. Kuster (nach dessen Tode fortgesetzt von W.

Kobelt und H. C. Winkauff). Verlag von Bauer and Raspe,

Nurnberg 3 (2), 336 pp.

Kool, S. P. 1993. Phylogenetic analysis of the Rapaninae (Neo-

gastropoda: Muricidae). Malacologia 35: 55-260.

Lamy, E. 1906. Gastropodes Prosobranches et Pélécypodes:

Expédition Antarctique Frangaise (1903-1905) Comman-

deé par le Dr. Jean Charcot. Masson et Cie. Paris, Sci-

ences Naturelles, 20 pp.

Lightfoot, J. 1786. A catalogue of the Portland Museum, lately

the property of the Duchess Dowages of Portland, de-

ceased: which will be sold by auction by Mr. Skinner and

Co. on Monday the 24th of April, 1786, and the thirty

seven following days, at twelve o'clock London, 194 pp.

Martyn, T. 1784. The Universal Conchologist, exhibiting the

figure of every known shell, accurately drawn, and painted

after nature: with a new systematic arrangement by the

author Thomas Martyn. Volume 1. T. Martyn, London,

pls. 1-80.

McLean, J. H. and H. Andrade. 1982. Large archibenthal Gas-

tropods of Central Chile Collection from expedition of the

R/V Anton Bruun and the Chilean Shrimp Fishery. Con-

tributions in Science 342: 1-20.

Melvill, J. C., and R. Standen. 1907. The marine mollusca of

the Scottish National Antarctic Expedition. Transactions

of the Royal Society of Edinburgh 46: 119-157.

Michelotti, A. J. 1846. Mitteilungen an Professor Bronn geri-

chtet. Neues Jahrbuch fiir Mineralogie, Geognosie, Geo-

logie und Petrefakten-Kunde, 1846: 52-56.

Michelotti, G. 1847. Description des fossiles des terrains mio-

cénes de [Italie septentrionale. Natuurkundige verhan-

delingen van de Bataafsche Hollandsche Maatschappye

der Wetenschappen te Haarlem 3(2): 1-408.

Molina, G. I. 1810. Saggio sulla storia naturale del Chile. Bo-

logna, 306 pp.

Montfort, P. D. D. 1810. Conchyliologie systématique, et clas-

sification méthodique des coquilles; offrant leurs figures,

Page 82

leur arrangement generique, leurs descriptions caracter-

istiques, leurs noms: ainsi que leur synonymie en plusieurs

langues. F. Schoell, Paris, 2, 676 pp.

Pain, T. 1980. A gallery of Muricids from around the world.

Hawaiian Shell News 28: 8-9.

Pallas, P. S. 1774. Spicilegia Zoologica quibus novae imprimis

et obscurae animalium species iconibus, descriptionibus

atque commentariis illustrantur. Berolini, 1 part 10, 41

pp.. 44 pls.

Pastorino, G. 2002. Systematics and phylogeny of the genus

Trophon Montfort, 1810 (Gastropoda: Muricidae) from

Patagonia and Antarctica: morphological patterns. Bollet-

tino Malacologico 38 (supplemento 4): 127-134.

Pastorino, G. and M. G. Harasewych. 2000. A Revision of the

Patagonian genus Xymenopsis Powell, 1951 (Gastropoda:

Muricidae). The Nautilus 114: 38-58.

Penchaszadeh, P. 1976. Reproduccién de gastropodos proso-

branquios del Atlantico suroccidental. E] género Trophon.

Physis, Seccién A, 35 (90): 69-76.

Penna-Neme, L., and J. L. Moreira Leme. 1978. Novas espé-

cies e novas ocorréncias de Gastropodos marinhos da Cos-

ta Brasileira (Prosobranchia, Neogastropoda). Papéis Avul-

sos de Zoologia 31: 283-297.

Perry, G. 1811. Conchology, or the natural history of shells:

containing a new arrangement of the genera and species,

illustrated by coloured engravings executed from the nat-

ural specimens, and including the latest discoveries. Bul-

mer and Co, London, 4 pp., 61 pls.

Petit de la Saussaye, M. S. 1856. Description de Coquilles nou-

velles. Journal de Conchyliologie 5: 87-92.

Philippi, R. A. 1868. Conchylia nova potissimum magellanica.

Malakozoologische Blatter 15: 223-226.

Powell, A. W. B. 1951. Antarctic and subantarctic Mollusca:

Pelecypoda and Gastropoda. Discovery Reports 26: 47—

196.

Ramirez-Bohme, J. 1981. Trophon (Enixotrophon) wilhelmen-

sis n. sp. (Mollusca, Gastropoda, Muricidae). Museo Na-

cional de Historia Natural, Noticiario Mensual 24 (301—

302): 5-7.

Reeve, L. 1847. Monograph of the genus Fusus. Conchologia

Iconica. Vol. 4, pls. 1-14.

Rehder, H. A. 1967. Valid zoological names of the Portland

Catalogue. Proceedings of the United States National Mu-

seum 121 (3579): 2-21.

Rios, E. 1985. Seashells of Brazil. Editora da Fundacéo Univ-

ersidade do Rio Grande, Rio Grande, 328 pp.

Rios, E. 1994. Seashells of Brazil. 2n¢ ed. Editora da Fundagao

Universidade do Rio Grande, Rio Grande, 368 pp.

Rochebrune, A. T. D. and J. Mabille. 1889. Mollusques. Mis-

sion Scientifique du Cap Horn. 1882-1883. Volume 6,

Zoologie, 2. Gauthier-Villars et fils, Paris, pp. H1-129.

Réding, P. F. 1798. Museum Boltenianum sive catalogus ci-

meliorum e tribus regnis naturae quae olim collegerat.

Joa. Fried Bolten, M. D. pd Pars Secunda continens con-

chylia sive testacea univalvia, bivalvia and multivalvia,

Hamburgo, 199 pp.

Scarabino, F. 2003. Lista sistemdtica de los Cephalopoda vi-

vientes de Uruguay. Comunicaciones de la Sociedad Ma-

lacolégica del Uruguay 8: 197-202.

THE NAUTILUS, Vol. 119, No. 2

Schumacher, C. F. 1817. Essais d'un nouveau systéme des hab-

itations des vers testacés. Schultz, Copenhagen, 287 pp.

Smith, E. A. 1915. Mollusca Part I Gastropoda Prosobranchia,

Scaphopoda and Pelecypoda. British Antarctic (“Terra

Nova’) Expedition, 1910. Natural History Report, Zoology

2: 61-112.

Solem, A. 1972. Malacological application of Scanning Electron

Microscopy, II. Radular structure and functioning. The

Veliger 14: 327-336.

Sowerby, G. B. II. 1846. Descriptions of Tertiary fossil shells

from South America. In: Darwin, C. Geological Obser-

vations on South America. Smith Elder Co, London, Pp.

249-967.

Sowerby, G. B. II. 1847-1887. Thesaurus Conchyliorum of

Monographs of Genera of Shells, London.

Strebel, H. 1904. Beitrige zur Kenntnis der Molluskenfauna

der Magalhaen-Provinz. Zoologischen Jahrbiichern. Ab-

teilung fiir Systematik, Geographie und Biologie der Ti-

ere, 21 (2): 171-248.

Strebel, H. 1908. Die Gastropoden. Wissenschaftliche Ergeb-

nisse der Schwedischen Sudpolar-expedition 1901-1903

Unter Mitwirkung Zahlreicher Fachgenossen Herausge-

geben von Otto Nordenskjold Leiter der Expedition. 6,

Zoologie (2), 111 pp.

Tapparone-Canefri, C. 1874. Zoologia del viaggio intormo al

globo della Regia Fregata Magenta durante gli anni 1865—

68. Malacologia (Gasteropodi, Acefali e Brachiopodi). Me-

morie della Reale Accademia delle Scienze di Torino (2)

28: 161.

Tryon, G. W. 1880. Muricinae, Purpurinae. Manual of Con-

chology (1) 2, 289 pp.

Vokes, E. H. 1970. Cenozoic Muricidae of the western Atlantic

region. Part 5. Pterynotus and Poirieria. Tulane Studies in

Geology and Paleontology 8 (1): 1-50.

Vokes, E. H. 1971. Catalogue of the genus Murex Linné (Mol-

lusca: Gastropoda); Muricinae, Ocenebrine. Bulletins of

American Paleontology 61: 5-141.

Vokes, E. H. 1991. Collecting trophons in Argentina. Part I,

Tierra del Fuego. American Conchologist 19: 7—10.

Vokes, E. H. 1991b. Collecting trophons in Argentina. Part II,

The Valdes Peninsula.. American Conchologist 19: 8=11.

Vokes, E. H. 1992. Argentine trophons revisited—or Dr. Pow-

ell, I owe you an apology. American Conchologist 20:

3-4.

Vokes, E. H. 1992b. Cenozoic Muricidae of the Western At-

lantic Region, Part IX. Pterynotus, Poirieria, Aspella, Der-

momurex, Calotrophon, Acantholabia and Attiliosa, addi-

tions and corrections. Tulane Studies in Geology and Pa-

leontology 25: 1-108.

Watson, R. B. 1882. Mollusca of H. M. S. “Challenger” Ex-

pedition. Part 13. The Journal of The Linnean Society,

Zoology 16: 358-392.

Watson, R. B. 1886. Report on the Scaphopoda and Gastero-

poda collected by H. M. S. Challenger during the Years

1873-76, London, 15 Zoology (42), 756 pp.

Wood, W. 1828. Index Testaceologicus, an illustrated catalogue

of shells, British and foreign, arranged according to the

Linnean System. Second Edition. W. Wood, London, 212

pp:

Zaixso, H. 1973. Observaciones sobre el desove y embriologia

de Trophon geversianus (Pallas, 1774). Neotropica 19

(60): 152-155.

THE NAUTILUS 119(2):83-89, 2005

Page 83

Latitudinal trends in shell characters of the neogastropod

Olivancillaria urceus (Gastropoda: Olividae) in the temperate

southwestern Atlantic Ocean

Alvar Carranza!

Walter Norbis

Seccién Oceanologia

Facultad de Ciencias

Igua 4295

11400 Montevideo

URUGUAY

ABSTRACT

Variation in shell characters of the neogastropod Olivancillaria

urceus (Réding, 1798) was analyzed at the central part of its

geographical range. Nine shell dimensions were measured

from specimens from seven localities between 32°10’ S and

40°33’ S along the Atlantic coast of South America. Significant

effects of collection site were detected on all measured vari-

ables. The first three components in principal component anal-

ysis together explained nearly 80% of the observed variance in

morphometric measures. Variables also differ in their allome-

tric trajectories among localities: all measured shell dimensions

displayed both positive and negative allometric coefficients

along the latitudinal axis. Allometric effects were then removed

to allow for determination of size-free variation in shape. High-

spired, elongated forms occurred more frequently in higher

latitudes. Spire width, maximum width, maximum width/height

and shell thickness decreased as latitude increased. Spire

length measured on the apertural side showed an opposite

trend. No relationship with latitude was found for fasciolar-

band length and spire length (measured along the abapertural

shell side). The existence of extreme forms within the analyzed

sample showed high phenotypic and ecological plasticity in the

populations assigned to O. urceus.

INTRODUCTION

Gastropod shell shape reflects a trade-off among func-

tional requirements, energetics of construction and

maintenance, rules governing growth, and the imprint of

evolutionary ancestry. The result of the interactions be-

tween these factors on the realized shell is also affected

by environmental features and tied to architectural con-

straints (Vermeij, 1993). Phenotypic plasticity in shell

characters appears to exert a strong influence on small

and large-scale morphological variation in marine snails

(Trussel and Etter, 2001).

Different patterns in shell variation are found as dif-

‘ [email protected]

ferent spatial scales and taxonomical or ecological hier-

archies are considered: well-disseminated eco-geograph-

ical rules, suitable for many taxa, predict that animals at

high latitudes should have larger body sizes and life

spans than at low latitudes (Mayr, 1956; Atkinson and

Sibly, 1997). In most mollusks growth is faster at high

temperatures and in the presence of food than in cold,

nutrient-poor conditions (Vermeij, 1993).

Latitudinal variation in shell characters was detected

in northern hemisphere gastropods assemblages. This

variability was related to efficiency of calcium carbonate

utilization. It was shown that more heavily calcified and

ornamented shells occur with decrease in latitude

(Grauss, 1974).

When considering variation in shell features at the in-

traspecific level, it could be interpreted as a response to

environmental conditions, which may include biotic and

abiotic factors. Biotic interactions have been proposed

as driving forces for morphological changes. Certain

shell characters interpreted as anti-predatory features

such as low spire and elongate aperture were found to

be significantly different between comparable gastropod

assemblages from both sides of the tropical Atlantic and

Indo-West Pacific (Vermeij, 1978).

Within-location variation in shell shape and shore-lev-

el size gradients have been found for several species of

intertidal mollusks (Vermeij, 1972). This intra-popula-

tional variability is often coupled with shifts in shell

growth and has been related experimentally with food

supply and density-dependant processes: specimens of

Littorina littorea (Linnaeus, 1758) with plentiful supply

of seaweed grow faster and develop low-spired shells

than other group maintained under crowded conditions;

in the latter growth was slower and higher spired shells

were developed (Kemp and Bertness, 1984). De Wolf et

al. (1998) documented large-scale patterns of shell var-

iation in Littorina striata, a planktonic-developing peri-

winkle from Macaronesia, confirming expectations based

Page 84

BRAZIL

Cassino

A La Coronilla

La Paloma

Punta del Este

ARGENTINA

(A. Mar dei Plata

A Puerto Militar

ATLANTIC OCEAN

San Antonio

Figure 1. Map showing parts of the Atlantic coast of Brazil,

Uruguay, and Argentina that include the collection sites of Oli-

vancillaria urceus.

on morphological patterns observed among other pros-

obranchs for increasing shell size and weight in southern

sites.

There is also ontogenetic related variation due to al-

lometric increase of spire length relative to other shell

dimensions (Vermeij, 1993). In this context, remotion of

allometric effects is needed in order to detect changes

in shell shape.

However, most of the studies dealing with morpho-

logical variability in gastropods are ecied to intertidal

rocky- shore snails, in particular those belonging to Lit-

torinidae. We know of no attempt made to analyze or

even describe patterns of intraspecific variation in neo-

gastropod shell features along a latitudinal axis in the

Southern Hemisphere.

The genus Olivancillaria d’Orbigny, 1839, is widely

distributed along the Atlantic coast of South America. It

comprises seven species occurring from tropical areas

(Espirito Santo, Brazil) to temperate regions (Golfo San

Matias, Argentina) (Burch and Burch, 1964; Klappen-

bach, 1964; 1965: 1966: Rios, 1994). Olivancillaria ur-

ceus (Réding, 1798) is distributed along the entire lati-

tudinal range of the genus in subtidal soft bottoms, rang-

ing from ine surf zone to at least 35 m depth (Juanics

and Rodriguez-Moyano, 1976; Milstein et al., 1976; Es-

cofet et al., 1979; Scarabino, 1984) and exhibiting con-

spicuous differences in shell shape as noticed by Bara-

tinni and Ureta (1961). For these reasons, this species

is suitable for the examination of patterns of morpho-

logical variation in the region.

In this paper we document large-scale variability in

shell features of Olivancillaria urceus in the temperate

THE NAUTILUS, Vol. 119, No. 2

Figure 2. Dorsal, ventral, and apical view of Olivancillaria

urceus showing measurements taken for statistical analysis.

Scale bar = 1 em. Abbreviations: LWL: last whorl length from

suture to anterior end; AL: aperture length, distance along out-

er lip from suture to anterior end; FB: distance between an-

terior and posterior ends of fasciolar band along internal side

of the aperture; MW: maximum linear distance from outer lip

to opposite side; MWH: maximum linear distance perpendic-

ular to MW to anterior end; ST: shell thickness; SW: diameter

of spire base from tip of callus above aperture to opposite point

on suture; SL2: lateral spire length from tip of callus above

aperture to protoconch tip on outer lip side in ventral view,

parallel to growth axis; and SL1: same length measured on the

opposite side

sector of it distribution range and analyze this species

variability along the emenal axis. We also discuss pos-

sible ecological implications of the observed pattern.

MATERIALS AND METHODS

A total of 193 specimens collected in seven sites be-

tween 32° and 40° S along the Atlantic coast of South

America were analyzed. Collection sites were Cassino

Beach, Brazil (32°10’ S, 52°20’ W, Site 1), La Coronilla

(33° 57’ S, 53°30’, Site 2), La Paloma (34°38’ S, 54°08’

W, Site 3) and Punta del Este (34°36’ S, W 58° 18’ W,

Site 4), Uruguay, and Mar del Plata (38°02’ S, 57° 32’

W, Sites 5), Puerto Militar (38°44’ S, 62°10’ W, Site 6)

and San Antonio Oeste (40°33’ S, 64°50’ W, Site 7), Ar-

gentina (Figure 1). Examined material is deposited at

Museo Argentino de Ciencias Naturales “Bermardino Ri-

vadavia” (Buenos Aires, Argentina) and Museo Nacional

de Historia Natural (Montevideo, Uruguay).

Nine shell dimensions were measured with vernier

A. Carranza and W. Norbis, 2005

Page 85

Table 1. Sample size, % of variation accounted by PC] and allometric coefficients for morphometric variables in each collection

site. Abbreviations: LWL: last whorl length from suture to anterior end; AL: aperture length, distance along outer lip from suture

to anterior end; FB: distance between anterior and posterior ends of fasciolar band along internal side of the aperture; MW:

maximum linear distance from outer lip to opposite side; MWH: maximum linear distance perpendicular to MW to anterior end;

ST: shell thickness; SW: diameter of spire base from tip of callus above aperture to opposite point on suture; SL2: lateral spire

length from tip of callus above aperture to protoconch tip on outer lip side in ventral view, parallel to growth axis; and SL1: same

length measured on the opposite side.

Locality

Variable 1 2 4 5 6 it

Sample size Q7 17 18 43 2) 28

% of variation accounted

for by PCL 65.33 67.54 71.81 94.23 87.82 75.44

Allometric coefficients

LWL 1.258 0.492 1.098 1.144 1.063 0.985 0.890

AL 1.451 0.374 1.166 0.965 1.072 1.028 1.125

FB 1.544 0.617 1.207 1.086 1.142 1.05 0.948

MW 1.101 0.382 1.089 1.102 1.134 1.123

MWH 1.386 0.459 1.163 1.041 1.134 1.026 1.062

ST 1.214 0.485 1.143 1.159 1.118 1.043 1.151

SW 1.294 0.760 1.338 1.101 1.136 1.165 1.285

SLI 0.025 2.573 0.426 0.619 0.623 0.750 0.765

SL2 —0.274 2.858 0.340 0.796 0.610 0.819 0.650

calliper, within an accuracy of 0.05 mm. Abbreviations

for the variables are as follows: LWL: last whorl length

from suture to anterior end; AL: aperture length, dis-

tance along outer lip from suture to anterior end; FB:

distance between anterior and posterior ends of fasciolar

band along internal side of the aperture; MW: maximum

linear distance from outer lip to opposite side; MWH:

maximum linear distance perpendicular to MW to an-

terior end; ST: shell thickness; SW: diameter of spire

base from tip of callus above aperture to opposite point

on suture; SL2: lateral spire length from tip of callus

above aperture to protoconch tip on outer lip side in

ventral view, parallel to growth axis; and SLI: same

length measured on the opposite side (Figure 2). Col-

lection sites were grouped a priori for discrimination of

differences among samples from different geographic

regions. For this we used the non-parametric Kruskall-

Wallis ANOVA median test. Allometric coefficients for

the seven collection sites were calculated for all vari-

ables. The method used was based on Jolicoeur (1963)

with additions from Kowalewski et al. (1997). Data were

log-transformed and subjected to principal component

analysis (PCA). The first principal component (PC1) was

regarded as a size axis. Correlation between sampling

size and explained variance was examined by means of

regression analysis. The allometric coefficient for each

original variable was estimated by dividing the PC1 load-

ing for that variable by the mean PC] loading over all

variables. Estimation of 95% confidence intervals for the

allometric coefficients was made by bootstrapping spec-

imens (2000 bootstrap replicates were made, not

shown).

Size effects were then removed, following Lleonart et

al. (2000), by scaling all individual to the same size

(mean LWL) adjusting their shape to the one they would

have in the new size according to allometric change.

PCA was run on transformed data and loads of the eight

remaining variables (i.e., all but total length) were cal-

culated. This removal method is one of the most pow-

erful tools available, since it is in turn a theoretical gen-

eralization of the technique used by Thorpe (1975,

1976), which was recorded as one of the most efficient

methods in the empirical evaluation done by Reist

(1985).

A cluster analysis (single linkage, Euclidean distances)

was then performed to discriminate among possible

morphological groups using site mean value for each var-

iable. The cophenetic correlation coefficient (CCC) was

calculated to measure the internal distortion of the clus-

ter analysis following Sokal and Rolf (1962). Finally, pos-

sible relationships between latitude and shell characters

were examined by regression analysis.

RESULTS

All variables showed significant differences between sites

(non-parametric ANOVA: p < 0.01). PCA performed

with log-transformed data showed that variance ex-

plained by PCI (Factor 1, regarded as size axis) differed

among sampling sites and varied between 65.33% (site

1) and 94.23% (site 5). Explained variance was not cor-

related with sample size (regression analysis, F = :0.010,

p-level = 0.922). Allometric coefficients calculated for

the nine variables also varied among collection sites. All

variables displayed both positive (b > 1) and negative

(b < 1) allometric trajectories (Table 1).

The first three factors in size-free PCA explained to-

gether 79.38% of the total variance. Variables MW,

MHW, ST, and SW grouped together with positive loads

on Factor 1. SL1 and SL2 formed another highly cor-

Page 86

THE NAUTILUS, Vol. 119, No. 2

Table 2. Results of size-free PCA Analysis. % of variance ac-

counted by each factor and loading of each variable are shown.

Abbreviations: LWL: last whorl length from suture to anterior

end; AL: aperture length, distance along outer lip from suture

to anterior end: FB: distance between anterior and posterior

ends of fasciolar band along internal side of the aperture; MW:

maximum linear distance from outer lip to opposite side;

MWH: maximum linear distance perpendicular to MW to an-

terior end; ST: shell thickness; SW: diameter of spire base from

tip of callus above aperture to opposite point on suture; SL2:

lateral spire length from tip of callus above aperture to pro-

toconch tip on outer lip side in ventral view, parallel to growth

axis; and SLI: same length measured on the opposite side.

Factor loadings

Variable PCl PC2 PC3

% of variation 41.54 22.9 14.95

AL 0.422 —0.474 0.550

FB 0.173 —().487 0.716

MW 0.874 0.2125 —0.063

MHW 0.888 0.083 0.034

ST 0.858 0.226 —0.11]

SW 0.811 0.246 —0.044

SLI —().094 0.815 0.446

SL2 —0.396 0.735 0.399

related group loading positively on Factor 2, and AL and

FB did the same on sEnctons ‘(Table 2).

Two different groups were discriminated when the

collection-site mean value for each variable were plotted

onto a hierarchical cluster, one of these groups com-

posed by individuals belonging to sites 6 and 7 (Figure

3). The calculated value for CCC was 0.93.

Latitudinal-related patterns of variation were found in

variables MW, MHW, ST, AL, and SW. This group of

variables showed a decreasing trend with increasing lat-

itude. On the other hand, SL2 tend to increase with

latitude, whereas SL1 and FB displayed no relationships.

The coefficient of determination (1?) explained less than

30% for all variables (Figure 4).

DISCUSSION

The present work documents shell shape variation in O.

urceus along a latitudinal gradient. The results show that

shell features vary Ihe sites. The analyses presented

above show that northern forms are stunt, low-spired,

and conical, whereas southern specimens (mostly those

from sites 6 and 7) are high-spired, narrower and elon-

gated.

Within location variation may be due to local differ-

ences in habitat conditions, for example, contrasting

beach morphodynamics, which are in turn associated

with different sediment features. However, the poor

geographical definition of collection sites did not allow

us to assign in all cases a particular individual to a par-

ticular habitat. Notwithstanding, intraspecific phenotypic

plasticity in a small spatial gealle is known to occur in

mollusks. For example, observations made on the mur-

icid Nucella lapillus (Linnaeus, 1758) showed that spec-

imens from very exposed shores are short-spired when

compared with those from sheltered localities (Cooke,

1895; 1915).

When considering variation at a geographic scale, two

possible scenarios are suggested to explain the among-

collection sites differences. These are either: (1) a grad-

ual response to shifts in environmental conditions along

the latitudinal gradient that generate a shape gradient

(in this scenario, macro-scale variation in physical pa-

rameters such as sea water temperature and calcium car-

bonate availability could be correlated with the observed

variation); or (2) different morphs associated with par-

Figure 3.

Extreme forms of Olivancillaria urceus in the study area. A. dorsal and apertural view of specimens from site 3. B.

Dorsal and apertural views of specimens from site 7. Scale bar = 1 cm

A. Carranza and W. Norbis, 2005

Page 87

36| MW=45.70 -0.60* Lat”

Maximum Width (mm)

©).00

> DER

eal

‘

pPHENPEP>

IDR P> > >

Max. Width Height (mm

See Us

ipa

> ae.

RED

28 a SW = 38,74 - 0,64 * Lat

r= 0.281

Spire Width (mm)

QErs

eae

Dae

—

NES

PE ROMDRD Dm

PipeRe

eaRNBO b>

Shell Thickness (mm)

NNN

je) NS

paea

| acaba

DIRE D>

Rae >

38} AL = 36.68 -0.14* Lat

36| r= 0.050

Aperture Length (mm)

(ee)

SESE

ro)

>>>

>DppEEDP >:

28 A

Spire Length 2 (mm)

(oo)

DIP RPP > >

31 32 33 34 35 36 37 38 39 40 41 42

MWH = 42.97- 0.47 * Lat ~

34| =0.284 |

> Dene >

D> EDP I> bP

ST = 39.92-0.55*Lat 4

32 I

30| 12=0.244 ‘A

28) 4 :

pa re

#& 0) CO

DPIDPp>> >

SL2 = -1.39 + 0.22 * Lat a

r= 0.153 t

1 32 33 34 35 36 37 38 39 40 41 42

Latitude

Figure 4. Olivancillaria urceus, relationship between standardized shell characters and latitude. Coefficients of determination (17)

and linear model fitted are shown.

ticular habitats, such as exposed sandy beaches or shel-

tered bays: broader specimens with short spire in high-

energy habitats and high-spired, elongated forms in pro-

tected habitats. In this case, likelihood of occurrence of

one particular morph depends on environmental char-

acteristics of geographic locations and may be consid-

ered local populational phenomena.

A related species, Olivancillaria vesica (Gmelin, 1791)

also shows two geographical forms, recognized as sub-

species, which are easily separable along its distribution

range (Klappenbach, 1966). This species shows an op-

posite latitudinal pattern: northern forms are narrower

and longer compared with the broader, “auriculated”

(ear-shaped) southern forms. In accordance with (2), the

latter form extends along the southern coast of Brazil

(Santa Catarina), Uruguay, and Argentina as far as

Puerto Quequén, Province of Buenos Aires, the same

geographical area in which O. urceus displays broader

and low spired shells. These broader shell forms could

be interpreted as associated with a large foot that en-

hances its ability of “anchoring” to the substratum in

higher energy environments, thus improving individual

fitness. However, more study is necessary on the taxo-

nomic status on O. vesica forms before any analysis of

its morphological variation.

It can be argued that differences in spire length and

width reflect shifts in growth patterns associated with

the adaptive process. It should be noticed that there is

a coincidence between occurrences of high-spired forms

(associated with slower growth mode) in higher lati-

Page 88

LINKAGE DISTANCE

Pto. Militar

San Antonio

Punta del Este

La Paloma

THE NAUTILUS, Vol. 119, No. 2

La Coronilla Cassino

Mar del Plata

COLLECTION SITES

Figure 5. Olivancillaria urceus, cluster analysis. Collection sites were grouped on the basis of similarities in the mean value for

each morphometric variable.

tudes. This could be supporting the hypothesis that both

latitudinal and environmental effects may be coupled to-

gether in a combined effect that masks the isolated ef-

fects of each factor.

As demonstrated by PCA analysis, nearly half of the

variance is explained by PC1 (42% of total variance),

which in turn is mainly affected by MW, MWH, ST, and

SW. This fact implies that changes in shell shape de-

tected within the analyzed sample are due mostly to dif-

ferences in its variables. Variation observed in SL2 and

SL1 are of much less importance as form determinants

when considering total variance components. However,

height and shape of the spire and protoconch characters

have been regarded as important specific (Lépez et al.,

1988) and supraespecific (Tursch, 1988) features, for

which these characters deserves further analysis. Nev-

ertheless, it seems that macrogeographic variation in

shell morphology strongly depend on local conditions,

which make adequate sampling and data treatment very

difficult.

ACKNOWLEDGMENTS

The authors wish to thank Mr. Fabrizio Scarabino (Di-

reccién Nacional de Recursos Acudticos, Montevideo),

Dr. Sergio Martinez and Dr. Alejandro Brazeiro (Facul-

tad de Ciencias, Montevideo), who provided useful bib-

liography and suggestions that helped us to improve this

manuscript. J. Lleonart made his own designed software

available for statistical analysis on remotion of allometric

effects. Authors would also like to express their gratitude

to the two anonymous referees for their great help in

reviewing and correcting the original version of the man-

uscript. A.C. thanks Msc. Estela Delgado for encourage-

ment and support and J. de los Santos for assistance with

the figures.

LITERATURE CITED

Atkinson, D. and R. M. Sibly. 1997. Why are organisms usually

bigger in colder environments? Making sense of a life his-

tory puzzle. Trends in Ecology and Evolution 12: 235-239.

Barattini, L. P and E. H. Ureta. 1961. La fauna de las costas

del este (invertebrados). Publicaciones de Divulgacién

Cientifica “Museo Damaso Antonio Larrafiaga,” Monte-

video, 108 pp.

Burch, J. Q. and R. L. Burch. 1964. The genus Agaronia J. E.

Gray, 1839. The Nautilus 77: 110-114, pls. 6-7.

Cooke, A. H. 1895. The Cambridge Natural History, 3 Mol-

luscs and Brachiopods. London, Macmillan, London, 535

eats A. H. 1915. The geographical variation of Purpura la-

pillus. Proceedings of the Malacological Society of London

40: 319-327.

De Wolf, H., T. Backeljau, S. Van Dongen and R. Verhagen.

A. Carranza and W. Norbis, 2005

1998. Large-scale patterns of shell variation in Littorina

striata, a planktonic developing periwinkle from Macaro-

nesia (Mollusca: Prosobranchia). Marine Biology 131(2):

309-317.

Escofet, A., N. Giannuca, S. Maytia and V. Scarabino. 1979.

Playas arenosas del Atlantico Sudoccidental entre los 29°

y 43°S.: consideraciones generales y esquema biocenol6-

gico. Memorias del Seminario sobre Ecologia Bentonicas

y Sedimentacién de la Plataforma Continental del Atlan-

tico Sur, 1: 245-258. UNESCO, Montevideo.

Graus, R. 1974. Latitudinal trends in the shell characteristics

of marine gastropods. Lethaia 7: 303-314.

Jolicoeur, P. 1963. The multivariate generalization of the allo-

metric equation. Biometrics 19: 97-499.

Juanicé, M. and M. Rodriguez-Moyano. 1976. Composicién

faunistica de la comunidad de Mytilus edulis platensis

dOrbigny, 1846, ubicada a unas 55 millas al SE de La

Paloma. Comunicaciones de la Sociedad Malacolégica del

Uruguay 4 (29): 113-116.

Kemp, P. and M. D. Bertness. 1984. Snail shape and growth

rates: evidence of plastic shell allometry in Littorina lit-

torea. Proceedings of the National Academy of Sciences

81: 811-813.

Klappenbach, M. A. 1964. A new species of Olivancillaria from

Uruguay and Brazil. The Nautilus 77: 132-134.

Klappenbach, M. A. 1965. Consideraciones sobre el género

Olivancillaria dOrbigny, 1840 (Mollusca, Gastropoda) y

descripcién de dos nuevas especies de aguas argentinas y

uruguayas. Comunicaciones Zoolégicas del Museo de His-

toria Natural de Montevideo 8 (104): 1-10, 2 pls.

Klappenbach, M. A. 1966. Olivancillaria vesica (Gmelin, 1791)

has priority over Olivancillaria auricularia (Lamarck,

1810) (Mollusca, Gastropoda). Archiv fiir Molluskenkunde

95 (1/2): 75-77.

Kowalewski, M., E. Dyreson, J. Marcot, J. Vargas, K. Flessa

and D. Hallman. 1997. Phenetic discrimination of bio-

metric singletons: paleobiological implications of morpho-

species in the lingulide brachiopod Glottidia. Paleobiology

23: 444469.

Lopez, A., M. Montoya and J. Lopez .1988. Two new species

of the genus Agaronia (Olividae) in the Panamic province

Page 89

and the description of two new species from Nicaragua.

The Veliger 30: 295-304.

Lleonart, J., J. Salat and G. J. Torres. 2000. Removing allo-

metric effects of body size in morphological studies. Jour-

nal of Theoretical Biology 205: 85-93.

Mayr, E. 1956. Geographical character gradients and climatic

adaptation. Evolution 10: 105-108.

Milstein, A., M. Juanicé and J. Olazarri. 1976. Algunas asocia-

ciones benténicas frente a las costas de Rocha, Uruguay.

Resultados de la campafia del R/V “Hero”, viaje 72-3A.

Comunicaciones de la Sociedad Malacolégica del Uruguay

4 (30): 143-164.

Reist, J. D. 1985. An empirical evaluation of several univariate

methods that adjust for size variation in morphometric

data. Canadian Journal of Zoology 64: 1363-1368.

Rios, E. C. 1994. Seashells of Brazil. Museu Oceanogréfico

Eliézer de Carvalho Rios da Fundagao Universidade do

Rio Grande, Rio Grande, 328 pp.

Scarabino, V. 1984. Clave para el reconocimiento de moluscos

litorales del Uruguay, I. Gastropoda. Contribuciones del

Depto. de Oceanografia de la Facultad de Humanidades

y Ciencias 1 (2): 12-22.

Sokal, R. R. and F. J. Rohlf. 1962. The comparisons of den-

drograms by objective methods. Taxon 11: 33-40.

Thorpe, R. S. 1975. Quantitative handlings of characters useful

in snake systematycs with particular references to intra-

specific variation in the ringed snake Natrix natrix (L.).

Biological Journal of the Linnean Society 7: 27-43.

Thorpe, R. S. 1976. Biometric analysis of geographic variation

and racial affinities. Biological Review 51: 407-452.

Trussell, G. C. and R. J. Etter. 2001. Integrating genetic and

environmental forces that shape the evolution of geo-

graphic variation in a marine snail. Genetica 112-113:

321-337.

Tursch, B. 1988. Protoconch measurements as supraespecific

characters in the family Olividae. The Veliger 31: 244-251.

Vermeij, G. J. 1972. Intraspecific shore level size gradients in

intertidal mollusks. Ecology 53: 693-700.

Vermeij, G. J. 1978. Biogeography and adaptation. Pattemms of

marine life. Harvard University. Press, Cambridge, 332

PP:

Vermeij, G. J. 1993. A natural history of shells. Princeton Uni-

versity Press (Eds.) 207 pp.

THE NAUTILUS 119(2):90-91, 2005 Page 90

Errata

Due to an editorial lapse, the Introduction section was omitted from Kohler and Glaubrecht’s (2005) article in the

most recent issue of The Nautilus. The missing Introduction is printed below (literature references are in the main

article), with apologies to the authors and readers.

Fallen into oblivion—the systematic affinities of the enigmatic Sulcospira Troschel, 1858 (Cerithioidea:

Pachychilidae), a genus of viviparous freshwater gastropods from Java

Frank Kohler

Matthias Glaubrecht

INTRODUCTION

The first volume of the well-known work “Das Gebiss der Schnecken zur Begriindung einer natiirlichen Klassifikation”

[Establishing a natural classification of snails from their dentition’] by erie iieamneritn Troschel (1810-1882) was

published in parts between 1856 and 1863. The work is an important historic landmark in the enduring challenge of

zoologists to create a natural classification of the living gastropods. In his work, Troschel (p. 117) described the genus

Sulcospira within the tribus “Pachychili’ mainly Thawed on features of the operculum and the radula (Figure 1). Ac-

cording to Robertson (1957), that description was published in 1858. Within the non-marine Cerithioidea, Sulcospira

represents one of the least known genera of Southeast Asian Pachychilidae, a group of viviparous freshwater gastropods

we have been focusing on in the last few years.

Recent research aiming to propose a phylogenetic systematics hypothesis and to establish a natural classification of

this limnic gastropod family will also allow a better understanding of their evolution, morphology, and ecology (Glau-

brecht, 1996; 1999: Kohler and Glaubrecht, 2001; 2002: 2003; Glaubrecht and Rintelen, 2003). Only recently, the

Pachychilidae Troschel, 1857, have been shown to represent a monophyletic group clearly distinct from the Thiaridae

Troschel, 1857, and from the other limnic Cerithioidea. This new concept has been suggested by analyses of mor-

phological data (e.g. Glaubrecht, 1996; 1999) and is corroborated by molecular data suggesting that Thiaridae (e.g.

Melanoides, Thiara. Tarebia) and Pachychilidae (e.g. Paracrostoma Cossmann, 1900, Pachychilus Lea, 1850) are “tat

very Closely related to each other (see phy logenetic Teconetanetion in Lydeard et al., 2002: figs. 1, 2). This classification

conflicts with the traditional view of most earlier authors who treated pachychilid taxa as Melemidas = = Thiaridae

(among others, Thiele, 1929; Rensch, 1934; Benthem-Jutting, 1956; Brandt, 1968; 1974) or Pleuroceridae (e.g. Ponder

and Warén, 1988; Vaught, 1989).

However, our current knowledge of the phylogeny and systematics of freshwater Cerithioidea in general and the

Pachychilidae in particular is still limited, since many taxa remain poorly known. Only recently, systematic studies

using morphological as well as molecular genetic data have shed some light on the relationships of several other genera

within the Pachychilidae. For instance, Kéhler and Glaubrecht (2001) presented comparative morphological data on

taxa traditionally assigned to Brotia H. Adams, 1866, by various authors, revealing that this genus as previously per-

ceived actually comprises four lineages, each characterized most conspicuously by distinct reproductive morphologies.

Subsequent studies including molecular phylogenetics data put special emphasis on two of these lineages, Jagora

Kohler and Glaubrecht, 2003, endemic to the Philippines (K6hler and Glaubrecht, 2003) and Tylomelania F. and P.

Sarasin, 1898, endemic to Sulawesi (Rintelen and Glaubrecht, 1999; 2003), suggesting an independent generic status

for each. In addition, the properties of another Australasian pachychilid genus, Psewdopotamis Martens, 1894, have

been extensively described by Glaubrecht and Rintelen (2003). Hence, a agiclelle portion of crucial biological infor-

mation on pachychilids has been amassed, helping to facilitate a better understanding of pachychilid systematics,

phylogeny, and evolution.

Nevertheless, there remain a number of systematic and taxonomic problems and difficulties related to this group

of freshwater snails. One of them will be dealt with in this study: The taxonomy and systematic position of Sulcospira.

Among the various generic names that have been introduced for Southeast Asian pachychilid taxa, Sulcospira is

clearly the oldest one. Its description predates that of other names, such as Brotia, Antimelania Fischer and Crosse,

1892, or Pseudopotamis; a complete annotated list of the introduced supraspecific names within the Southeast Asian

Pachychilidae is given in Kohler and Glaubrecht (2002). Consequently, every other supraspecific pachychilid taxon is

valid only with the reserve that it is not a junior synonym of Sulcospira. In spite of this significant taxonomic role of

Sulcospira, this taxon has been widely ignored, especially by modern systematists. Thus, another aim of this article is

to compile all available information on Sulcospira and to provide new data from our own examinations of the limited

Errata, 2005 Page 91

material from museum collections. In addition, implications for pachychilid taxonomy and systematics are discussed

in relation to Sulcospira.

LITERATURE CITED

Kohler, F. and M. Glaubrecht. 2005. Fallen into oblivion—the systematic affinities of the enigmatic Sulcospira Troschel, 1858

(Cerithioidea: Pachychilidae), a genus of viviparous freshwater gastropod from Java. The Nautilus 119: 15-26.

In the recently published article by Ardila and Valdés (2004), please note the following corrections:

On page 134, caption of Figure 3, replace “Armina muelleri Thompson, Cattaneo and Wong, 1990”, for “Armina

muelleri (Ihering, 1886)”;

On the same page, the adjacent subsections “Type Material” and “Type Locality” should be replaced by:

Material Examined: MHNMC INV MOL3901, 32 mm length, alive, from off Salamanca Island, Colombia (11°5’46”

N, 74°40'35” W), 20 m depth, hard bottom with pennatulaceans (Renilla reniformis and Renilla muelleri).

LITERATURE CITED

Ardila, N. E. and A. Valdés. 2004. The genus Armina (Gastropoda: Nudibranchia: Arminidae) in the southern Caribbean, with the

description of a new species. The Nautilus 118: 131-138.

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erick

CONTENTS

INO Ga Baie

Volume 119, Number 3

October 6, 2005

ISSN 0028-1544

Cristian Ituarte

Shannon M. Carpenter

The Sphaeriidae (Bivalvia) from northwestern Argentina including three

new species Olt JERSTCHADN stove wacoy Be aN Cathe Oo Be Ee Ta hee SON Soe EU ee 93

Mysella pedroana, a commensal bivalve (Lasaeidae) on two decapod

GHUSEACE ATIBLLOS LSet a armnmne ary erat ke carn Ora Hey betes Moats cnr ae e fee Faber Rie 105

Paolo Mariottini Coralliophila trigoi (Gastropoda: Muricidae), a new species from the

Carlo Smriglio morinensuenn Auk QWs ,..6n00s0ccvocs ns esonassouosunsenon 109

Emilio Rolan

Book Review............ 116

THE NAUTILUS 119(3):93-104, 2005

Page 93

The Sphaeriidae (Bivalvia) from northwestern Argentina including

three new species of Pisidium

Cristian Ituarte

Division Zoologia Invertebrados

Museo de La Plata

1900 La Plata

Buenos Aires

ARGENTINA

[email protected]

ABSTRACT

Knowledge on the Sphaeriidae fauna in southern South

America is significantly improved with the description of three

new species of the genus Pisidiwm from Salta and Jujuy

provinces (northwestem Argentina). This paper provides the

first record of sphaeriids in restrictive high-altitude South

American environments, particularly from very small water

courses found in “vegas”, exceptionally localized areas of

“cushion vegetation” or “cushion peat bogs” developing

between 2000-4000 m altitude in the Argentine pre-Andean

ranges. Furthermore, the geographic distribution range of

Pisidium chiquitanum Ituarte, 2001, only reported to date from

the type locality in sub-Andean regions of central Bolivia, is

considerably enlarged.

INTRODUCTION

The Sphaeriidae fauna from Argentina is poorly known;

the first reports were those by Strobel (1874) on

Musculium argentinum (d’Orbigny, 1835) and Pilsbry

(1911) describing several new species of Pisidium C.

Pfeiffer, 1821, and Musculium Link, 1807, from

Patagonia. Later on, Doello-Jurado (1921) described

the first species of Eupera Bourguignat, 1854, from

Argentina, and Ituarte (1989, 1994) and Ituarte and

Dreher-Mansur (1993) described three new species of

Eupera from Iguazti, Uruguay and Parana River basins

in northeastern Argentina. Regarding the species di-

versity of Pisidiuwm C. Pfeiffer, 1821, twelve species are

known from Patagonia and Northeastern provinces

(Ituarte, 1996, 1999, 2000).

Only two species of Sphaeriidae have been reported

from northwestern Argentina: Sphaerium lauricochae

(Philippi, 1869), from Jujuy Province (Ituarte, 1995)

and Musculium argentinum from Mendoza Province

(Strobel, 1874). The species diversity of Pisidiwm

has essentially not been documented. In the

present paper, three new species of Pisidium

from lowland and high-altitude habitats in the pre-

Andean mountain ridges are described. Based on new

findings the knowledge on the geographic distribu-

tion of Pisidium chiquitanum Ituarte, 2001, is

updated.

MATERIALS AND METHODS

Materials for the present study were obtained during

three field trips to Northwestern Argentina (Tucuman

and Salta provinces in March 1999, Salta and Jujuy

provinces in December 2001 and March 2004, and

Catamarca Province in March 2004). Figure 1 shows the

location of collecting sites; more detailed information on

the source of specimens is given in the Systematics

section. The collected specimens were fixed immediately

after collecting in 70° alcohol after being relaxed through

a short rinse (around 20 seconds) in warm water (about

50°C). Specimens for scanning electron microscopy

(SEM) were cleaned by repeated rinsing in distilled

water followed by a short treatment (about 5 seconds) in

10% sodium hypochlorite solution. Soft anatomy was

studied after decalcification of valves through a 12-hour

rinsing in a 5% formaldehyde and 2% acetic acid

solution. Linear measurements (shell length [SL], shell

height [SH], shell width [SW] and presiphonal suture

length [PSS]), shape indices and morphometric ratios

(height index [HI= SH/SL], convexity index [Ci= SW/

SH], ratio of hinge length [HiL] to shell length [HiL/

SL]), were calculated according to the criteria followed

by Ituarte (1996). For each calculation (n= 10, unless

otherwise stated), mean and standard deviation values

are given.

Type specimens are deposited at Museo de La Plata,

La Plata (MLP), Museo Argentino de Ciencias Naturales

“Berardino Rivadavia’, Buenos Aires (MACN), Funda-

cion Miguel Lillo, Tucuman (FML) and Muséum

National d'Histoire Naturelle, Paris (MNHN). Types of

Pisidium chiquitanum, housed at Museo de Historia

Natural “Noel Kempff Mercado”, Santa Cruz de La

Sierra, Bolivia (MHNB) and MLP were also used for

comparative purposes.

Page 94

THE NAUTILUS, Vol. 119, No. 3

Tumbayae

\—Termast’ Juju

Jujuy

de Reyes | ae

SS. WW

Salta

68° Ww

Figure 1.

SYSTEMATICS

Pisidium omaguadca new species

(Figures 2-15)

Diagnosis: Shell markedly oval, high and anteriorly

produced, beaks depressed, displaced backward, not

projecting from shell surface and only barely visible

above dorsal margin; ligament internal; anal and

branchial mantle openings present; two demibranchs

present, nephridia of closed type, with lateral lobe not

visible in lateral or dorsal views.

Description: Shell thin, translucent, small to medium

size (mean SL = 2.73 + 0.17, maximum observed size:

3.7 mm), rather high (mean HI = 85 + 1) (Figure 2),

not inflated (mean Ci = 58 + 3) (Figures 2, 3). Shell

Location map. Stars in black indicate type localities of Pisidium omaguaca new species, Pisidium ocloya new species, and

Pisidium chicha new species.

outline markedly oval, anteriorly elongated. Anterior end

produced in a sharp curve, posterior end short, widely

rounded, sometimes slightly truncated and_ straight

(Figures 2, 4, 5, 8). Dorsal margin short, weakly

connected with anterior margin, which slopes markedly

towards anterior end; sometimes, a gentle angle marks

joining point of dorsal and posterior margins (Figure 8).

Beaks very low, depressed and wide, not raised from

shell surface, only slightly projected above dorsal

margin, displaced backward, located at about 59% of

SL (Figures 2, 4, 5). Shell surface finely and somewhat

irregularly striated, glossy, amber.

Hinge plate strong, hinge line short (HiL/SL = 53 +

3), strongly curved. Hinge: Left valve (Figures 4, 6):

cardinal teeth well-developed, the inner one (Cg) thin,

short at base, bent upward, slightly oblique with respect

C. Ituarte, 2005

‘ > O5

Page 95

nig 2-7. Pisidiwm omaguaca from Termas de Reyes, Jujuy. 2. Holotype (MLP 5496-1-1): outer view of right valve. 3-7.

Paratypes (MLP 5496-1-2). 3. SPottenor view. 4. Left valve, inner view. 5. Right valve, inner view. 6. Left valve, detail of hinge. 7

Right valve, detail of hinge. Scale bars = 500 pm.

to antero-posterior axis, rounded at the tip, outer

cardinal tooth (C4) a narrow, slightly wider at posterior

end, uniformly curved blade, quite obl ique, overlapping

C3 at posterior half; anterior lateral tooth (AII) very

strong, short, nearly straight, cusp high, pointed,

displaced forward; posterior lateral tooth (PII) minute,

straight and strong, cusp high, distally displaced. Right

valve (Figures 5, 7): cardinal tooth (C3) not strong,

somewhat displaced forward, curved in the middle, quite

narrow at anterior half, slightly enlarged in a posterior,

aera CYR

Page 96

THE NAUTILUS, Vol. 119, No. 3

Figures 8-13. Pisidiwm omaguaca. 8. Paratype (MLP 5485). Right valve, outer view. 9. Paratype (MLP 5496-1

-2). Dorsal view,

detail of the escutcheon. 10-11. Inner view of left and right valves ‘ofa specimen from Tiraxi, Jujuy (MLP 6535). 12, 13. Outer and

inner views of right and left valves of a specimen from a “peat bog” between Yavi (Salta) and Santa Victoria (eer (MLP6559). Scale

bars (8, 10-13) = 1000 um; (9) = 200 Lm.

elongated cup. Lateral teeth short and robust, inner

anterior lateral (AI) curved, cusp subcentral or slightly

displaced anteriorly; outer anterior lateral tooth (AIT)

quite short, cusp distal: inner posterior lateral (PI) nearly

straight, short, cusp subcentral; outer posterior lateral

(PIII) minute, with distal cusp.

Ligament-pit enclosed, deep, inner margin straight or

slightly concave (Figures 6, 7). Ligament relatively short,

representing 20 + 1% of shell length, strong, internal,

never visible from outside (Figure 9). Escutcheon

inconspicuous (Figure 9).

Anatomy: Anal siphon and branchial inhalant mantle

opening present. Presiphonal suture about 9% of shell

length (Figure 15). Eight or nine well-marked muscle

scars located away from pallial line correspond to inner

radial mantle muscles. Muscle scars corresponding to

anal siphon retractors are coalescent with that of

C. Ituarte, 2005

Hin (MA

Figures 14-15. Pisidiwm omaguaca new species. 14. Gross

anatomy. 15. Mantle muscles. (as: anal siphon; asr: anal

siphon retractor; ia: inhalant aperture; id: inner demibranch;

isr: inhalant siphon retractor; irm: inner radial mantle muscles;

n: nephridium; od: outer demibranch; pss: presiphonal suture).

posterior adductor muscle (Figures 4, 5). Bundles of

fibers of inner radial mantle muscles strong, converging

anteriorly, except for two posterior bundles (Figures 4,

5, 14).

Inner and outer demibranchs present. Outer demi-

branch much smaller, formed by 10-12 very short

descending filaments, reaching back to the 14" filament

of inner demibranch (Figure 14). Up to three large

embryos (1.3 mm length) were found within each brood

pouch of a specimen 3.7 mm L. Nephridia of closed

type, dorsal lobe, usually subquadrate, completely

covering pericardial part of nephridium (Figure 14).

Type Locality: A small watercourse opening into Reyes

River at Termas de Reyes, 24°10'19" S, 65°29'27” W,

1754 m altitude, Jujuy Province, Argentina (Figure 1).

Type Material: Holotype (MLP 5496-1-1) and 42

paratypes from the type locality (16 paratypes MLP

5496-1-2; 6 paratypes MLP 5485; 4 paratypes MACN-In

Page 97

36361; 12 paratypes FML 14506; and 4 paratypes

MNHN.

Other Material Examined: Jujuy Province: numer-

ous specimens from the type locality (MLP 5496-1-3);

Tumbaya, small pool with vegetations at the side of

national road No. 9 (23°47'28” S, 65°28'37” W), 2070 m

(MLP 6548); small stream flooding from springs in

highland areas covered with “cushion vegetation”,

between Yavi (Jujuy) and Santa Victoria (Salta)

(22°07'11" S 65°13'05” W), 4150 m (MLP 6559): small

pool at the side of Rio Grande River (22°58’14" S

65°27'01" W), 3950 m (MLP 6530); small springs at side

of Manzanito Rivulet, near Huertas (22°14'20" S

65°00'31” W), 2740 m (MLP6551); small stream at the

side of the road to Yala (24°07'20" S 65°24'16” W),

1430 m (MLP 6531). Salta Province: unnamed brook on

provincial road No. 57, near Cachi (25°05'24" S

66°07'33” W), 2340 m (MLP 6540).

Distribution: Highlands of Jujuy and Salta provinces,

Argentina, between 1400 and 4100 m altitude.

Etymology: The name refers to the Omaguacas,

ancient aboriginal inhabitants of the Quebrada de

Humahuaca, the spectacular 150 km long valley of the

Rio Grande River (Figure 1), which underwent a major

cultural change during the past 10,000 years.

Remarks: Pisidiwm omaguaca new species differs

from Pisidium meierbrooki Kuiper and Hinz, 1984, in

being higher and less obese, having lower and not

inflated beaks, and a more broadly rounded posterior

end. Pisidiwm omaguaca new species is similar to

Pisidium chiquitanum Ituarte, 2001, in soft anatomy,

but differs in having a more solid and higher shell, with

posterior end shorter and anterior half of dorsal margin

sloping markedly towards the anterior end.

Pisidium ocloya new species

(Figures 16-33)

Diagnosis: Shell rather trapezoidal and high, small size,

presence of only one (anal) mantle aperture and one

demibranch.

Description: Shell thin, translucent, of small to

medium size (maximum observed SL = 3.2 mm), high

(mean HI = 85 + 2), quite convex (mean Ci = 77 = 4),

shell outline rather trapezoidal. Anterior end somewhat

produced in a sharp curve, posterior end _ short,

truncate, somewhat oblique (Figures 16, 17). Beaks

full, wide at base, markedly raised from shell surface

and projected above dorsal margin, subcentral or

slightly displaced backward, located at about 58% of

SL (Figures 16, 19, 20). Shell surface finely and

irregularly striated (Figures 16, 18), glossy, whitish or

yellowish.

Hinge plate not strong, narrow in middle, hinge line

rather long (HiL/SL = 56 + 2), arcuate. Hinge on right

valve (Figures 20, 22): cardinal tooth (C3) delicate,

THE NAUTILUS, Vol. 119, No.

Figures 16-22. Pisidiwm ocloya new species from Burrumayo River, Jujuy. 16. Holotype (MLP 5499-1): outer view of right valve.

17-22. Paratypes (MLP 5499-2). 17. Posterior view. 18. Posterior view, detail of ligament. 19. Left valve, inner view. 20. Right

valve, inner view. 21. Left valve, detail of hinge. 22. Right valve, detail of hinge. Scale bars (16, 17, 19-22) = 500 um; (18) =

200 pm.

rather weak, slightly curved, narrow on anterior half,

enlarged into a well-marked, blunt, posterior cup.

Lateral teeth robust, inner anterior lateral (AI) well-

developed, cusp displaced distally; outer anterior lateral

tooth (AIII) shorter and weaker, cusp distal; inner

posterior lateral (PI) gently curved, not long, cusp

distally displaced; outer posterior lateral (PIII) reduced

in size, with distal cusp. Hinge on left valve (Figures 19,

91): cardinal teeth minute, inner one (Cs) short and

high, horizontal with respect to antero-posterior axis,

C. Ituarte, 2005

outer one (C4) short, oblique, slightly overlapping Cz at

posterior end; anterior lateral tooth (AII) very strong,

cusp high, distal; posterior lateral tooth (PII) shorter and

weaker, cusp high, distal.

Ligament-pit enclosed, deep, inner margin slightly

and evenly curved (Figures 21, 22), Ligament narrow,

relatively long, representing 24 + 1% of shell length,

visible from outside and somewhat protruded at anterior

half of ligament length. Escutcheon slightly marked by

a delicate lanceolate line (Figures 17, 18).

ANATOMY: Only one demibranch (inner) present

(Figure 23). Only anal mantle opening present. Inner

radial mantle muscles weak, 6—7 bundles of few weak

fibers converging anteriorly, attached just above pallial

line (Figure 23); sometimes scars corresponding to inner

radial muscles coalescent with pallial line (Figures 19,

20). Anal siphon retractors attached immediately ven-

trally to posterior adductor muscle. Nephridium with

lateral loop visible dorsally (Figure 24).

Type Locality: Small flooded areas on the banks of

Burrumayo River (24°10'18” S, 65°22'43” W), 1201 m

altitude, in the neighborhood of Jujuy City, Jujuy

Province, Argentina; and unnamed brook opening into

Zapla River (24°16'03” S, 65°07'09” W), 946 m altitude,

Zapla, Jujuy Province, Argentina.

Type Material: Holotype (MLP 5499-1) and 15

paratypes from the outskirts of Jujuy City (6 paratypes

MLP 5499-2; 10 paratypes FML 14505; 56 paratypes

from Zapla (36 paratypes MLP 6899-2; 10 paratypes

MACN-In 36362; 10 paratypes MNHN).

Other Material Examined: Catamarca Province:

unnamed brook at national road No. 40 at La Ciénaga

de Abajo, between La Ciénaga and Belén (27°31'05" S,

66°59'08" W), 1520 m (MLP 7201); Jujuy Province:

numerous specimens from the type locality (MLP 7369);

unnamed brook on side of provincial road No. 4 at

Guerrero (24°11'13” S, 65°26’51” W), 1650 m (MLP

7379); small spring on side of provincial road No. 4, near

Termas de Reyes (24°10'36" S 65°28'18” W), 1730 m

(MLP 5497-1, MLP 7370-1); small water course on side

of national road No.9 at Tumbaya (23°51’26" S,

65°27'57” W), 2020 m; Los Cedros Rivulet, south to

“El Carmen” (24°28'0” S 65°17'08” W), 1190 m (MLP

6545-1); small spring mear Tiraxi (23°59'57" S,

65°19’39" W), 1576m (MLP 6553); on the road

between Termas de Reyes and Laguna Yala, flooded

areas at the bottom of hills (24°07'47” S, 65°28'58” W),

1920 m (MLP 7371). Salta Province: unnamed brook at

national road No.40 at Molinos (25°18'53” S,

66°14’58”" W), 2155 m (MLP 6526); small spring at road

No. 33 to Cachi at Los Laureles (25°06'27" S,

65°36'10” W), 1360 m (MLP 6529): Tucumdan Province:

flooded areas at side of the Nio River, near Rio del Nio

City (26°25.60’ S 64°55.60’ W), 886 m (MLP 7403); on

side of Medina River, on provincial road No. 305,

between El Tipal and Aserradero (MLP 7404-1); small

Page 99

Figures 23-24. Pisidiwm ocloya new species. 23. Gross

anatomy. 24. Detail of nephridium. (id: inner demibranch;

irm: inner radial mantle muscles; n: nephridium; p: pericar-

dium; pa: posterior adductor; r: rectum.

pond on side of Potrero de Las Tablas River, Raco,

880 m (MLP6992), La Angostura dam, on Los Sosa

River (26°55/21” S, 65°41'02” W), 2000 m (MLP 6897-

2).

Distribution: Catamarca, Jujuy, Salta, and Tucuman

provinces, Argentina, between 880 and 2155 m altitude.

Etymology: The name of the new species refers to the

Ocloyas, ancient aboriginal inhabitants of the lands in

the surroundings of the type locality.

Remarks: Pisidiwm ocloya new species strikingly

differs from other northwestern Argentine Pisidiwm

species by its relatively small size and trapezoidal shell

outline. Compared with Pisidium vile Pilsbry, 1987,

a small species from the eastern drainage system of the

Rio de La Plata Basin, P. ocloya new species differs in

being larger and comparatively lower. Ituarte (1999)

described Pisidium huillichum from southern Chile,

another small-sized Pisidium species with one mantle

opening and one demibranch, which differs from P.

ocloya in having a non trapezoidal shell outline, lower

beaks, very strong lateral teeth, and strongly marked

commarginal ribs of the shell surface.

The height/length ratio, the convexity index and the

degree at which part of the ligament is protruded

showed a relatively wide variability in samples of

Pisidium ocloya from different localities (Figures 25—

33); larger specimens were in general more convex with

much inflated and pronounced beaks, and more sharply

defined trapezoidal outline.

Page 100

THE NAUTILUS, Vol. 119, No. 3

Figures 25-33. Pisidiwm ocloya new species. 25-27. Specimens from Termas de Reyes, Jujuy (MLP 5497-1). 28-30. Specimens

from Zapla, Jujuy (MLP 6899-2). 31-33. Specimens from Cachi, Salta (MLP 6529). Scale bars =

Pisidium chicha new species

(Figures 34-43)

Diagnosis: Rather elliptic shell outline, sub-centrally

located beaks, somewhat inflated and markedly raised

from shell surface, are diagnostic features. The presence

of only one mantle aperture and one demibranch is also

distinctive.

Description: Shell small, maximum observed shell

length 2.7, moderately high (mean HI = 79 + 1), not

convex (mean Ci = 68 + 5): shell outline strikingly oval.

Dorsal and ventral margins broad, dorsal margin, slightly

arcuate, ventral margin uniformly curved; anterior end

500 um.

evenly curve and only slightly projected forward,

posterior end slightly nested (Figures 34-37). Beaks

wide, widely rounded at tip, somewhat inflated, raised

above dorsal mar gin but low, sub-central, slightly

displaced backward, located at about 57-58% of SL.

Shell surface glossy, amber, sculptured with well marked

fine and rather regularly spaced striae (Figure 34).

Hinge plate narrow, hinge line somewhat short, HiL/

SL = 53 + 3% of SL (n = 8), widely curved. Hinge on

left valve (Figures 36, 38): cardinal teeth well-developed,

the inner one (Cg) thin, long, straight at base, bent

upward distally, parallel with respect to antero-posterior

axis, rounded at tip, outer one (C4) a slender, slightly

C. Ituarte, 2005 Page 101

Figures 34-41. Pisidiwm chicha new species. 34. Holotype (MLP 6899-1-1): outer view of right valve. 35-41. Paratypes (MLP

6899-1-2). 35. Posterior view. 36. Left valve, inner view. 37. Right valve, inner view. 38. Left valve, detail of hinge. 39. Right valve,

detail of hinge. 40. Right valve, detail of cardinal tooth and ligament. 41. Posterior view, detail of ligament. Scale bars (34-37) =

500 tm; (38, 39) = 200 um; (40) = 250 um; (41) = 100 um.

Page 102

THE NAUTILUS, Vol. 119, No. 3

curved blade, quite oblique, overlapping Cy on posterior

half; anterior lateral tooth (AII) very strong, straight,

cusp high, somewhat acute, displaced distally; posterior

lateral tooth (PII) short, strong, cusp high, distal. Hinge

on right valve (Figures 37, 39, 40): cardinal tooth (C3)

well-developed, narrow, and evenly curved on anterior

half, quite enlarged into triangular, slightly grooved cup

at posterior end; slightly hanging from inner margin of

hinge plate. Lateral teeth short and robust, inner

anterior lateral (AI) somewhat curved, cusp displaced

forward; outer anterior lateral tooth (AIII) very short,

straight, with distal cusp; inner posterior lateral one (PI)

short, straight, slender and low, cusp sub-central; outer

posterior lateral one (PII) minute, with distal cusp.

Escutcheon lanceolate, long, outline demarcated by

a very delicate line. Ligament-pit enclosed, inner margin

gently sinuous (Figures 40, 41). Ligament moderately

strong, internal, slightly visible from exterior, but not

protruded, representing about 23% of shell length.

Anatomy: Only one (anal) mantle opening present.

Only inner demibranch present (Figure 42). Inner radial

mantle muscles weak, inserted just above or coalescent

with pallial line. Nephridium with dorsally visible lateral

loop (Figure 43).

Type Locality: Unnamed brook flooding into Zapla

River in the neighborhood of Zapla City (24°16'01” S,

65°07'09" W), 946 m altitude, Jujuy Province, Argen-

tina, and small springs on bank of Manzanito Rivulet, on

the road from Santa Victoria East to Yavi near Huertas

(22°14'20" S, 35°00'31" W), 2740 m, Jujuy Province,

Argentina.

Type Material: Holotype (MLP6899-1-1), 9 para-

types (7 paratypes MLP 6899-1-2; 2 paratypes MACN-

In 36363) from 24°16’ S, 65°12’ W; 62 paratypes from

22°14'20" S, 35°00'31" W (52 paratypes MLP 6550; 5

paratypes FML 14775; 5 paratypes MNHN).

Other Material Examined: Catamarca Province:

flooded areas at side of an unnamed river on the road

to Singuil, (27°38'25" S, 65°57'23"” W), 2000 m

(MLP7203). Jujuy Province: Los Cedros Rivulet, close

to Las Maderas Dam (24°28’40" S 65°17'08" W),

1190 m (MLP 6545-2). Tucuman Province: small pools

at the side of Medina River, on provincial road 305

between El Tipal and Aserradero (MLP 7404-2).

Distribution: Catamarca, Jujuy and Tucuman pro-

vinces, Argentina, between 940 and 2740 m altitude.

Etymology: The name of the new species alludes to

the Chichas, a small ethnic group that was a part of the

Omaguaca people, who inhabited the lands in the

neighborhood of the type locality.

Remarks: The shell shape, quite ovate and nearly

equilateral, and the marked sculpture of Pisidium chicha

new species are distinctive features that allow for easy

identification of the new species among other Pisidiwm

species from northwestern Argentina. A moderate shell

Figures 42-43. Pisidiwm chicha new species. 42. Gross

anatomy. 43. Detail of nephridium. (be: brooding embryos; id:

inner demibranch; n: nephridium; pa: posterior adductor; sr:

siphon retractor).

variation was observed: the specimens from Manzanito

Rivulet (MLP 6550) show shells slightly more convex

and higher than those of the specimens from Zapla

(MLP 6899-1); the striae were slightly coarser and more

marked in the former group. Specimens from Catamarca

Province (MLP 7203) show slightly inequilateral shells,

with posterior end slightly shorter. Pisidium chicha

shares with P. ocloya the same number of mantle

openings and demibranchs and the nephridium with

lateral lobe dorsally visible; but the former species

strikingly differs in having a smaller shell with quite an

oval shell outline. The presences of only one demibranch

and one mantle opening in P. chicha clearly separate this

species from P. chiquitanum. In addition, this latter is

a larger species of relatively similar shell outline but that

also differs from P. chicha in having an almost smooth

shell surface, lower, more backward displaced beaks,

and somewhat truncated posterior end.

Pisidium chiquitanum Ituarte, 2001

(Figures 44-47)

Pisidium chiquitanum Ituarte, 2001: 50; figs. 2-14 (La Siberia,

West of Comarapa, Santa Cruz de La Sierra, Bolivia,

holotype MHNB 34734).

Diagnosis: Rather elongate shell outline, slightly

truncated at posterior end, low and posteriorly located

beaks, ligament position, internal but externally visible,

presence of branchial and anal openings, two demi-

branchs on each side and nephridia of closed type.

C. Ituarte, 2005

Page 103

Description: Shell thin, small to medium size (mean

SL = 3.9 + 0.25: maximum observed size: 4.2 mm),

not very high (mean HI = 80 + 1), not convex (mean

Ci = 61 = 4), shell outline markedly oval, elongate,

anteriorly produced, posterior end short, widely

rounded, or slightly truncated and _ nearly straight

(Figures 44, 46, 47). Beaks low, depressed, slightly

projected above dorsal margin, located at about 62% of

SL (Figures 44-46). Shell surface dull glossy, straw-

yellowish, with fine and low commarginal striae,

moderately more accentuated towards the shell margin

(Figures 44, 46).

Hinge plate solid, hinge line rather long (HiL/SL =

56 + 2), Hinge on right elhies Right cardinal tooth (C3)

strongly curved in middle, quite narrow on anterior half,

enlarged into gently sulcated, rounded, or somewhat

elongate cup. Right lateral teeth robust, inner anterior

lateral (AI), widely curved, long, cusp sub-central or

slightly displaced forward; outer anterior lateral tooth

(AIII) quite short, cusp distal; inner posterior lateral (PI)

nearly straight, short, cusp sub-central; outer posterior

lateral tooth (PII) reduced in size with distal cusp. Left

valve (Figure 47): cardinal teeth short, inner one (C3)

short, oblique with respect to antero-posterior axis, outer

one (C4) a narrow curved lame, quite oblique, over-

lapping Cs on posterior half; anterior lateral tooth (AII)

strong, straight, cusp sub-central; posterior lateral tooth

(PII) narrow and weak, cusp distal.

Ligament-pit enclosed, deep, inner margin slightly

sinuous, concave at posterior end. Escutcheon well

marked by a delicate line; ligament long, internal, but

visible from outside in anterior half through a very

narrow and sometimes rather long gap between valves,

never protruded. Ligament length is 23 + 1% of shell

length.

Anatomy: Anal siphon and branchial mantle opening

present. Presiphonal suture rather long, representing 11

+ 2% of SL. Anal siphon well-developed, pair of

powerful siphonal retractors present. Inner radial mantle

muscles, 8 bundles as rule, inserted away from pallial

line, scars of those corresponding to anal siphon

retractors coalescent with posterior adductor muscle

scars. Inner and outer demibranchs present. Outer

demibranch reduced in size, composed of 11-15 Tea

short descending filaments, reaching back to the 14-16

filament of inner demibranch. Nephridia of closed type,

dorsal lobe variable in shape, commonly subquadrate,

with lateral loop not visible in dorsal view.

Material Examined: Holotype (MHNB 34734) La

Siberia, West of Comarapa, Santa Cruz de La Sierra,

Bolivia; Tucuman, Argentina: 27°01'24” S, 65°39'29" W

(MLP 6554): Cerro Munoz, Santa Cruz, 26°54’ S,

65°46'42” W, 2400 m (MLP 6991); La Angostura dam,

26°56’ S, 65°41'03” W, 1800 m (MLP 6527): Jujuy,

Argentina: Los Laureles, 25°06/27" S, 65° 36/ 10” W,

1360 m (MLP 6528); Los Toldos, Santa Victoria De-

partment, 1770 m (MLP 6993); small brook near Tiraxi

(23°59'57” S, 65°19'39” W), 1576 m (MLP 6552): small

Figures 44-47. Pisidium chiquitanum Ituarte, 2001. 44, 45.

Paratype (MLP 5362). 44. Outer view of right valve. 45.

Posterior view. 46, 47. Specimen from Tiraxi, Jujuy (MLP

6552). 46. Outer view of right valve. 47. Inner view of left

valve. Scale bars = 500 Lm.

spring at the side of provincial road No. 4, near Termas

de Reyes (24°10'36" S, 65°28'18” W) (MLP 5497-2).

Distribution: Ranging from sub-Andean regions in

Siberia (west of Comarapa) in central Bolivia (1800 m

altitude) southward to northwestern Argentina (between

1360 and 2400 m altitude).

Remarks: Pisidiwm chiquitanum can be easily identi-

fied among South American Pisidium species by its oval

shell outline with low beaks and internal (however

visible from the outside) ligament. It is also character-

ized by two, inner and outer, demibranchs on each side,

two siphonal openings, and nephridia of closed type.

Page 104

Pisidium chiquitanum resembles Pisidiuwm meierbrooki

Kuiper and Hinz, 1984, from Peru and Bolivia, which is

the only known species from tropical South America

with both, branchial and anal, siphonal openings

(Ituarte, 1995). Pisidiwm meierbrooki differs from P.

chiquitanum in having a more convex shell (according to

data in Kuiper and Hinz, 1984, the Ci varies between 77

and 80), fuller and more backward displaced beaks. As

pointed out by Ituarte (2001)Pisidium chiquitanum is

similar to specimens from Ecuador and Peru reported

by Kuiper and Hinz (1984), as Pisidium casertanum

(Poli, 1791), an Eurasian species extremely variable in

shell shape, currently reported as cosmopolitan (Burch,

1975; Kuiper, 1983; Kuiper and Hinz, 1984; Holopainen

and Kuiper, 1982). However, these specimens are larger

than P. chiquitanum, having more central beaks and less

produced anterior end. P. chiquitanum also differs from

P. casertanum in having less convex shell, lower and

narrower beaks, decidedly displaced backward. The

specimens from northwestern Argentina slightly differ

from the ones from Bolivia in being generally higher,

with beaks slightly less displaced in posterior direction

(Figure 46).

ACKNOWLEDGMENTS

The author is grateful for the warm friendship and kind

support received from colleagues Gabriela Cuezzo,

Fatima Romero, and Carlos Molineri during the three

field trips to northwestern Argentina. This study was

funded by grant: PIP 554/98 from the Consejo Nacional

de Investigaciones Cientificas y Técnicas (CONICET),

Argentina. The author is researcher of the CONICET.

LITERATURE CITED

Burch, J. B. 1975. Freshwater Sphaeriacean clams (Mollusca:

Pelecypoda). Malacological Publications, Hamburg, Mi-

chigan, 96 pp.

THE NAUTILUS, Vol. 119, No. 3

Doello-Jurado, M. 1921. Una nueva especie de Eupera del Rio

de la Plata. Physis 5: 72-75.

Holopainen, I. J. and G. J. Kuiper. 1982. Notes on the

morphometry and anatomy of some Pisidiwm and

Sphaerium species (Bivalvia, Sphaeriidae). Annales Zool-

ogici Fennici 19: 93-107.

Kuiper, J. G. J. 1983. The Sphaeriidae of Australia. Basteria 47:

3-52.

Kuiper, J. G. J. and W. Hinz. 1984. Zur Fauna der

Kleinmuscheln in den Anden. Archiv fiir Molluskenkunde

114/1983]: 137-156.

Ituarte, C. F. 1989. Los géneros Byssanodonta dOrbigny,

1846 y Eupera Bourguignat, 1854 (Bivalvia: Sphaeriidae)

en el area parano-platense. Descripcién de Eupera

iguazuensis n. sp. del rio Iguazii, Misiones, Argentina.

Neotropica 35: 53-63.

Ituarte, C. F. 1994. Ewpera guaraniana n. sp. (Pelecypoda:

Sphaeriidae) del rio Uruguay, Argentina. Gayana, ser.

zool. 58: 1-7.

Ituarte, C. F. 1995. Nuevos registros de Pisidium Pfeiffer,

1821 y Sphaerium Scopoli, 1777 (Bivalvia: Sphaeriidae) en

Chile, Bolivia y Noroeste argentino. Neotropica 41: 3141.

Ituarte, C. F. 1996. Argentine species of Pisidium Pfeiffer,

1821, and Musculium Link, 1807 (Bivalvia: Sphaeriidae).

The Veliger 39: 189-203.

Ituarte, C. F. 1999. Pisidium chilense (d’Orbigny, 1846) and

new species of Pisidium C. Pfeiffer, 1821 from southern

Chile (Bivalvia, Sphaeriidae). Zoosystema 21: 249-257.

Ituarte, C. F. 2000. Pisidiwm taraguyense and Pisidium

pipoense, new species from Northeastern Argentina

(Bivalvia: Sphaeriidae). The Veliger 43: 51-57.

Ituarte, C. F. 2001. Pisidium chiquitanum new species from

Santa Cruz de la Sierra, Bolivia (Bivalvia: Sphaeriidae).

The Nautilus 115; 50-54.

Ituarte, C. F. and M. C. Dreher-Mansur. 1993. Eupera

elliptica n. sp., una nueva especie en el rio Iguazt,

Misiones, Argentina. Neotropica 39: 11-16.

Pilsbry, H. A. 1911. Non-marine Mollusca of Patagonia.

Reports of the Princeton University Expedition to

Patagonia 1896-1899, 3(Part 5): 513-633.

Strobel, P. 1874. Materiali per una malacostatica de terra e di

acqua dolce dell’Argentina. Pisa.

THE NAUTILUS 119(3):105-108, 2005

Page 105

Mysella pedroana, a commensal bivalve (Lasaeidae) on two

decapod crustacean hosts

Shannon M. Carpenter

Santa Barbara Museum of Natural

History

Department of Invertebrate Zoology

2559 Puesta del Sol Road

Santa Barbara, CA 93105-2998 USA

[email protected]

ABSTRACT

Mysella pedroana (Dall, 1898), lives commensally on Isocheles

pilosus (Holmes, 1900), and Blepharipoda occidentalis Randall,

1839. Because of their small size at reproductive maturity,

specimens attached to Isocheles pilosus were previously

referred to in the literature as an undescribed species.

However, preliminary comparisons between the variability of

hinge dentition and internal structure of those specimens and

small individuals of Mysella pedroana indicate that they

represent instead a single species. Variability of characters

including size at reproductive maturity and shell morphology

suggests that Mysella pedroana is a highly variable species.

INTRODUCTION

A small bivalve living attached to hermit crabs has

perplexed biologists for over a decade. It was referred to

as Mysella sp. H by Valentich-Scott and Barwick (2001),

as an undescribed species. It was shown to be

reproductively mature at 1.0 mm in length.

Bivalves in the family Lasaeidae have been taxonom-

ically problematic (Dall, 1898; Gage, 1966a; 1966b;

O Foighil and Eermisse 1988; Morton and Scott,

1989). These bivalves combine features of immaturity

induced by changes in environmental factors, perhaps

due to their commensal nature. Mysella pedroana

(Dall, 1898), is no exception. It is found both as

a commensal and free-living (Scott, 1987); however,

due to its prevalence on hosts, it may prefer a commensal

habitat.

Mysella pedroana was previously thought to be host-

specific and found only on the sand crab Blepharipoda

occidentalis Randall, 1839, (Burch and Burch, 1944;

Boss, 1965a; Lafferty, 1993; Boyko and Mikkelsen,

2002). Originally described by Dall in 1898 from a single

valve, M. pedroana was then redescribed (Boyko and

Mikkelsen, 2002) and associated with B. occidentalis as

host. The discovery of its occurrence on another host,

the hermit crab Isocheles pilosus (Holmes, 1900),

previously undocumented variation in shell morphology,

and small size at reproductive maturity has led to this

present study.

Blepharipoda occidentalis (Decapoda: Albuneidae) is

found in the northeastern Pacific from Stinson Beach,

Marin County, California, USA, to Bahia Santa Rosalia,

Baja California, Mexico (Morris et al., 1980) both

intertidally and subtidally, burrowing in sand. Isocheles

pilosus (Decapoda: Diogenidae) ranges from Bodega

Bay Harbor, Sonoma County, Califormia, USA, to Estero

de Punta Banda, Baja California, Mexico (Rickets et al.,

1985). Like B. occidentalis, it is intertidal, but also occurs

in mud flats, bays and estuaries including depths

offshore up to 55 meters. Isocheles pilosus most

commonly inhabits shells of Polinices or Kelletia (pers.

comm. Scott, 2004) and crawls on the sand or buries

with only its eyes and mouth visible (Fager, 1968).

Mysella pedroana is either attached to the crab’s setae or

in its branchial chambers.

MATERIALS AND METHODS

Preserved specimens identified by Valentich-Scott and

Barwick (2001) as Mysella sp. H from the collections in the

Santa Barbara Museum of Natural History (SBMNH)

were studied. Additional material included SBMNH

specimens associated with preserved specimens of Iso-

cheles pilosus and B. occidentalis, along with specimens

from two living I. pilosus and one B. occidentalis collected

at Sands Beach, Santa Barbara. Specimens removed from

I. pilosus are deposited as SBMNH 351472351480 and

47635. Material of M. pedroana examined includes

SBMNH 348251, 345553, 348252, and 348253.

Of the 145 available specimens, 35 were opened and

examined for gross anatomy. Characters examined were

internal structure, shell dentition, and shape. Measure-

ments were made with vernier calipers and anatomical

observations were performed under a dissecting micro-

scope. Scanning electron micrography was performed at

SBMNH with a Zeiss EVO 40 XVP with a variable-

pressure secondary electron detector.

Page 106

THE NAUTILUS, Vol. 119, No. 3

Figure 1. A growth series of commensal Mysella pedroana, SBMNH 351473. Scale bar = 1 mm.

Living Mysella perdroana were placed in Petri dishes

with sea water and coarse sediment for comparison of

movement in different media. Specimen with brood

was stained with crystal violet in distilled water prior to

examination.

RESULTS

Mysella pedroana (Dall, 1898)

Description: SHELL (Figures 1—3): Shell morphology

variable from ovate to subtrigonal, thin, more elongate

anteriorly; umbones opisthogyrate; beaks range from

central to posterior; shell surface white with poorly

defined commarginal striae; periostracum thick and

yellow, variable in texture, rough and dehiscent to

smooth and adherent; prodissoconch line present;

maximum shell length 11.0 mm, mean height to length

ratio 77% (herein) (SD +4.8; range 11.0-1.0 mm) to

80% (Boyko and Mikkelsen, 2002).

Muscle scars apparent in larger specimens with

anterior adductor scar elongate and posterior ovate;

pallial line entire; hinge with prominent subumbonal

resilium; two lateral cardinal teeth on right valve with

longer anterior tooth and slight groove, posterior shorter;

teeth diverging into an inverted V; left valve with thin

grooves (lamellae) that interlock with the right valve.

Extensive variation shown in smaller specimens (length

1-3.4 mm), which may exhibit a posterior tubercular

tooth on the right valve with a similar posterior

tubercular tooth on the left valve.

Gross Anatomy: Mantle papillate, more so anteriorly;

ctenidium encompasses large area of internal space and

serves as a brood chamber; presence of eggs observed in

specimens from 1-11] mm in length; labial palps small.

OBSERVATIONS ON LIVE SPECIMENS: Activity was enhanced

following regular replacing of cold sea water. Clams

were observed crawling on surface of Petri dish with

shell positioned vertically, in the manner of a gastropod;

dug in sand with anterior end, process took 15-20 sec.

Detached M. pedroana took 3 min. to reattach to

setae on the ventral surface of I. pilosus. However,

when I. pilosus setae were easily accessible and M.

pedroana was placed next to them, attachment took 30—

45 sec.

Host INFORMATION AND PREVALENCE: Blepharipoda occi-

dentalis had a carapace length of 4.06-4.55 cm (n = 3)

and all three had Mysella pedroana present. Isocheles

pilosus had a carapace length of 1.9-2.57 cm (n = 6)

with three of the six crabs having bivalves present.

Figures 2-3. Mysella pedroana, hinge dentition. 2. From host Isocheles pilosus, SBMNH 351473, SEM. Scale bar = 200 um.

3. From host Blepharipoda occidentalis, SBMNH 351478, SEM. Scale bar = 1 mm.

S. M. Carpenter, 2005

Page 107

Figures 4-5. Mysella pedroana. 4. Host Isocheles pilosus showing attachment between the chela and carapace (arrows), SBMNH

345553, Scale bar = 2.2 mm. 5. On host Isocheles pilosus with byssus (arrow), SBMNH 351472. Scale bar = 0.8 mm.

Three B. occidentalis were examined with 100%

prevalence. Of the six preserved museum specimens of

I. pilosus, only one was infested. However, two live

I. pilosus were examined and fifty-nine M. pedroana

specimens were retrieved.

The live B. occidentalis examined had two M.

pedroana that were found on the external surface near

the antennae and on the second pereopod.

Host Isocheles pilosus: Mysella specimens on I. pilosus

were found in the branchial chambers, attached to the

chelae, the junction between the chela and carapace, the

ventral setae and branchial chambers (Figure 4). The

largest specimens were on the chelae (1 mm) with

smaller individuals on the ventral surface (0.7—1 mm)

and juveniles (<0.7 mm) anteriorly in the right and left

branchial chambers. Those on the ventral surface and in

the branchial chambers were attached with byssus

(Figure 5). The few on the chelae were observed

crawling or attached by byssal threads to the surface

spines of the chelae. Upon preservation in 70% ETOH

M. pedroana retained its byssal threads.

DISCUSSION

This study suggests that shell shape of Mysella pedroana

is more variable than previously appreciated. These

variable characteristics include texture of the periostra-

cum (rough and dehiscent to smooth and adherent),

dentition, shape of the shell. These differences have

been attributed to environmental conditions in other

molluscan shells (Wellington and Kuris, 1983; De Wolf

et al., 1998).

Shell variation probably led to the redescription of

Mysella pedroana as Rochefortia golischi by Dall in 1916

(Burch and Burch, 1944); these were later synonymized

by Scott (1987). However, this variation is also common

to many commensal species and makes it difficult to

identify them based on shell morphology (O Foighil and

Eernisse, 1988: Morton and Scott, 1989).

Due to shell variation and the small size at re-

productive maturity of M. pedroana, these smaller

bivalves on I. pilosus were thought to be a new species,

referred to as Mysella sp. H (Valentich-Scott and

Barwick, 2001). However, minimum size of brooding

adults of M. pedroana had been previously reported at

1 mm (Valentich-Scott and Barwick, 2001) and sper-

matogenesis has been confirmed in specimens of

1.2mm. Previous observations for individuals of M.

pedroana undergoing spermatogenesis were 4.7 mm in

length (pers. comm. Kevin Lafferty, 2004). Reproductive

maturity at small sizes has been reported also for

Pseudopythina macrophthalmensis at 2.0 mm (Jespersen

et al., 2001) and Mysella bidentata with egg production

at 1.7 mm (O Foighil et al., 1984).

The prevalence of Mysella pedroana was higher on B.

occidentalis than on I. pilosus. Those on B. occidentalis

were also larger in the gill chambers compared to only

juveniles found in the gills of I. pilosus. Further

examination of both hosts would be required to see if

there is a host preference.

Reattachment to a host is possible for the species.

Both Lafferty (pers. comm. 2004) and Valentich-Scott

(pers. comm. 2004) noted that individuals could drop

their byssus and disassociate from the host if perturbed.

Individuals of M. pedroana could use this procedure to

move from host to host or to become free-living.

The above observations on morphology and symbiont-

host relationships permit the identification of this

bivalve as Mysella pedroana. Previously reported host

specificity for commensal bivalves may be due to the

scarcity of studies or experimental observations (Boss,

1965b; Morton and Scott, 1989). This species was

previously thought to be host-specific (Boyko and

Mikkelsen, 2002), an assumption here shown not to be

valid.

Page 108

ACKNOWLEDGMENTS

I would like to thank to Paul Valentich-Scott for his

guidance, knowledge and for introducing me to the

world of bivalves. The Hearst Foundation Internship

through the Santa Barbara Museum of Natural History

made this research possible. Scanning electron micro-

scope facilities funded by NSF grant number

MRI0420726, Daniel Geiger assisted with SEM images.

Michael Caterino, Henry Chaney and Armand Kuris for

reviewing preliminary drafts. Shane Anderson provided

living crabs, Patricia Sadeghian contributed her knowl-

edge of crustaceans. Paula Mikkelsen and Kevin Lafferty

whom offered additional data. Kelvin Barwick contrib-

uted many specimens. Two anonymous reviewers

offered critical observations which greatly improved

the manuscript.

LITERATURE CITED

Boss, K. J. 1965a. Symbiotic erycinacean bivalves. Malacologia

3: 183-195.

Boss, K. J. 1965b. A new mollusk (Bivalvia, Erycinidae)

commensal on the stomapod crustacean Lysioquilla.

American Museum Novitates 2215: 1-11.

Boyko, C. B. and P. M. Mikkelsen. 2002. Anatomy and biology

of Mysella pedroana (Mollusca: Bivalvia: Galeommatoi-

dea), and its commensal relationship with Blepharipoda

occidentalis (Crustacea: Anomura: Albuneidae). Zoolo-

gischer Anzeiger 241: 149-160.

Burch, J. Q. and T. Burch. 1944. [Family Montacutidae].

Distributional list of the west American marine Mollusca

from San Diego, California, to the Polar Sea, Part I.

Pelecypoda. Conchological Club of Southern California,

Minutes 40: 14-16.

Dall, W. H. 1898. Synopsis of the Recent and Tertiary

Leptonacea of North America and the West Indies.

Proceedings of the United States National Museum 21:

873-897.

De Wolf, H., T. Backeljau, V. S. Dogen and R. Verhagen.

1998. Large-scale patterns of shell variation in Littorina

striata, a planktonic developing periwinkle from Macro-

nesia (Mollusca: Prosobranchia). Marine Biology 131:

309-317.

THE NAUTILUS, Vol. 119, No. 3

Fager, E. W. 1968. A sand-bottom epifaunal community of

invertebrates in shallow water. Limnology and Oceanog-

raphy 13(3): 448-464.

Gage, J. 1966a. Observations on the bivalves Montacuta

substriata and M. ferrunginosa, ‘commensals’ with spa-

tangoids. Journal of Marine Biology Association of the

United Kingdom 46: 49-70.

Gage, J. 1966b. The life-histories of the bivalves Montacuta

substriata and M. ferruginosa, ‘commensals’ with spatan-

goids. Journal of Marine Biology Association of the United

Kingdom 46: 499-511.

Jespersen, A., T. Kosuge and J. Lutzen. 2001. Sperm

dimorphism and spermatozeugmata in the commensal

bivalve Pseudopythina macrophthalmensis (Galeommatoi-

dea, Kelliidae). Zoomorphology 120: 177-189.

Lafferty, K. D. 1993. Ecology and parasites of the spiny sand

crab, Blepharipoda occidentalis. American Zoologist 33: 19A.

Morris, R. H., D. P. Abbott and E. C. Haderlie. 1980.

Intertidal Invertebrates of California. Stanford University

Press, Stanford, 584 pp.

Morton, B. and P. H. Scott. 1989. The Hong Kong

Galeommatacea (Mollusca: Bivalvia) and their hosts, with

descriptions of new species. Asian Marine Biology 6:

129-160.

O Foighil, D., D. McGrath, M. E. Connely, B. F. Keegan and

M. Costelloe. 1984. Population dynamics and reproduc-

tion of Mysella bidentata (Bivalvia: Galeommatacea) in

: Galway Bay, Irish west coast. Marine Biology 81: 283-291.

O Foighil, D. and D. J. Eemisse. 1988. Geographically

widespread, non-hybridizing sympatric strains of the

hermaphroditic, brooding clam Lasaea in the northeastern

Pacific Ocean. Biological Bulletin 175: 218-229.

Ricketts, E. F., J. Calvin and J. W. Hedgpeth. 1985. Between

Pacific Tides. 5‘ ed. Stanford University Press, Stanford,

336 pp.

Scott, P. H. 1987. A preliminary review of Mysella (Bilvalvia,

Montacutidae) from the northwestern Pacific. Western

Society of Malacologists 19: 13-14.

Valentich-Scott, P. and K. Barwick. 2001. Mysella sp. H

(Bivalvia, Lasaeidae). Southern California Association of

Marine Invertebrate Taxonomists Newsletter 20(2):

13-14.

Wellington, G. M. and A. M. Kuris. 1983. Growth and shell

variation in the tropical eastern Pacific intertidal gastro-

pod genus Purpura: ecological and evolutionary implica-

tions. Biological Bulletin 164: 518-535.

THE NAUTILUS 119(3):109-115, 2005

Page 109

Coralliophila trigoi (Gastropoda: Muricidae), a new species

from the northeastern Atlantic Ocean

Paolo Mariottini'

Dipartimento di Biologia

Universita di “Roma Tre”

Viale Marconi 446

00146 Roma

ITALY

[email protected]

Carlo Smriglio

00167 Roma

ITALY

Via di Valle Aurelia 134

[email protected]

Emilio Rolan

Museo de Historia Natural,

Campus Universitario Sur, 15782

Santiago de Compostela

SPAIN

[email protected]

ABSTRACT

Based on shell characters and with further support from mo-

lecular data, Coralliophila trigoi, a new species of gastropod of

the family Muricidea, is here described from the northeastern

Atlantic Ocean. The new taxon, consisting of several specimens

mainly collected along the Atlantic Spanish coast, has previ-

ously been misidentified in the literature as Coralliophila bas-

ilea (Dautzenberg and H. Fisher, 1896). Coralliophila trigoi

new species is conchologically similar to Coralliophila meyen-

dorffi (Caleara, 1845), and Coralliophila panormitana (Mon-

terosato, 1869), but it can be easily separated from them mainly

because it is differently sculptured. The new species is com-

pared with other members of the genus Coralliophila from the

same geographical area and Mediterranean Sea. Molecular se-

quencing of the internal transcribed spacer 2 region (ITS2) of

the nuclear rDNA and part of the mitochondrial gene for 12S

rDNA confirm the validity of the new species.

INTRODUCTION

The coralliophilines form a monophyletic group of neo-

gastropods that includes approximately 200-250 de-

scribed species grouped, based on their shell morphol-

ogy, in at least 7-10 “genera”, distributed worldwide in

temperate and tropical oceans. The subfamily Corallio-

philinae Chenu, 1859 (for the phylogenetic relationship

of this muricoidean groups, see Oliverio and Mariottini,

2001a) includes species invariably associated with cni-

darians, which are generally used as food by the gastro-

pods. Shell variability, absence of radula, absence of a

preserved protoconch (often eroded in adults and even

in young specimens), together with a limited knowledge

of the anatomy, represent constrains to the understand-

ing of the taxonomic status of this group of neogastro-

pods. Their classical systematics above the species level

is at present far from being stable (Clover, 1982; Bou-

chet and Warén, 1985; Kosuge and Suzuki, 1985; Oliv-

erio, 1989; Vaught; 1989; Oliverio, in press). Data from

1 Author for correspondence.

mitochondrial and nuclear genes (12S rDNA and ITS2

rDNA, respectively) have been recently utilized in the

proposal of a molecular framework for the phylogeny of

these muricids (Oliverio and Mariottini, 2001a; Oliverio,

Cervelli and Mariottini, 2002). Data from both sequence

and secondary structure show that Rapaninae Gray, 1853

(=Thaidinae Jousseaume, 1888) are their sister group

(Harasewych et al., 1997; Oliverio and Mariottini, 2001a;

Oliverio, Cervelli and Mariottini, 2002), indicating a

monophyletic radiation of the Coralliophilinae. The

state-of-the-art knowledge about feeding, anatomy, sex-

ual strategies, parental care, and protoconch of corallio-

philines was recently reviewed by Richter and Luque

(2002). The authors reported the available data on pro-

toconch and larval development of many coralliophilines

belonging to ten different genera, including Corallio-

phila H. and A. Adams, 1853. We had the opportunity

to examine several shells of a coralliophiline that we

could allocate to any of the Atlantic and Mediterranean

species of this subfamily. These shells, mostly collected

along the coast of Galicia, Spain, were previously mis-

identified in the literature as Coralliophila basilea

(Dautzenberg and H. Fisher, 1896) (Rolan, 1983; Rolan,

Lépez and Gutiérrez-Garcia, 1995). After comparisons

with other species, we realized that they represent an

undescribed species, possibly related to Coralliophila

meyendorffii (Calcara, 1845) and Coralliophila panor-

mitana (Monterosato, 1869).

In order to verify the taxonomic validity of Corallio-

phila trigoi, we carried out molecular sequencing of the

internal transcribed spacer 2 region (ITS2) of the nucle-

ar rDNA and of part of the mitochondrial gene for 12S

rDNA. Genomic DNA was extracted from the dissected

foot of two freshly collected individuals with standard

methods (SDS-proteinase K digestion, phenol/chloro-

form extraction, ethanol precipitation (Oliverio and Mar-

iottini, 2001b)). Mitochondrial rDNA was amplified

through the polymerase chain reaction (PCR) with prim-

ers 12S-I and 12S-III (Oliverio and Mariottini, 2001a).

Nuclear ribosomal ITS2 was amplified using the primers

Page 110

THE NAUTILUS, Vol. 119, No. 3

Table 1. Collecting data and DDBJ/EMBI/GenBank accession number for specimens assayed in molecular systematics.

Species/individuals

Accession number

Collecting locality and depth 12S ITS2

Coralliophila neritoidea

Coralliophila brevis

Coralliophila mejendorffii

Coralliophila panormitana

Coralliophila trigoi new species,

specim. #1

Coralliophila trigoi new species,

specim. #2

its-3d and its-4r complementary to conserved regions of

the ribosomal coding portions on the 5.8S and 28S

rRNAs (Oliverio and Mariottini, 2001b). PCR-amplified

products were directly sequenced by an automated se-

quencer. Nucleotide sequences were first aligned by

hand and the alignment progressively optimized accord-

ing to secondary structure homology. Phylogenetic anal-

yses were performed using PAUP* 4b10 (Swofford,

2002). GenBank accession numbers (12S and ITS2) of

the Coralliophila trigoi sequences are reported in Ta-

ble 1.

Institutional abbreviations used: MNCM, Museo

Nacional de Ciencias Naturales, Madrid, Spain; MZB,

Laboratorio di Malacologia, Museo di Zoologia

dell’ Universita di Bologna, Italy.

Abbreviations used for collections: CS-PM, Carlo

Smriglio and Paolo Mariottini (Rome, Italy); ER, Emilio

Rolan (Vigo, Spain); FS, Frank Swinnen (Lommel, Bel-

gium); JT, Juan Trigo (Brion, A Coruna, Spain); MO,

Marco Oliverio (Rome, Italy).

SYSTEMATICS

Superfamily Muricoidea Rafinesque, 1815

Family Muricidae Rafinesque, 1815

Subfamily Coralliophilinae Chenu, 1859

Genus Coralliophila H. and A. Adams, 1853

Type Species: Fusus neritoideus Lamarck, 1816,

Ency. Meth., pl. 435, figs. 2a—b. (=Purpura violacea Kie-

ner, 1836), by subsequent designation (Iredale, 1912).

Recent, Indo-Pacific.

Coralliophila trigoi new species

(Figures 1-8, 13-14, 17-21)

Description: Shell of large size, up to 35 mm length.

Protoconch usually eroded in adult specimens (proto-

conch observed in only one juvenile specimen, albeit

worn and lacking the embryonic stage). Protoconch in-

dicative of planktotrophic larval development, multispi-

ral, composed of protoconch I and II. Protoconch II of

about 24% whorls, with a diameter of about 800 wm,

showing two strong spiral keels, only one visible above

the suture of the first whorl, crossed by axial ribs forming

nodules at intersections. The protoconch-teleoconch de-

Taiwan, 23°10’ N, 120°05’ E, 5 m depth

La Maddalena Is. (Sardinia, Italy), 41°15’ N, 009°26’ E, 30 m depth

La Maddalena Is. (Sardinia, Italy), 41°15’ N, 009°26’ E, 3-7 m depth

Cape Circeo (Latium, Italy), 41°11’ N, 013°04' E, 70 m depth

Camarifias, Galicia, Spain, northeastern Atlantic Ocean, 15-50 m depth

AJ293679 AJ420258

AJ293676 AJ420256

AJ297517 AJ293661

AJ293681 AJ420259

AJ937305 AJ937307

AJ937306 AJ937308

marcation is well-defined, marked by a varix. Teleoconch

shape biconical, elongate ovoid-fusiform, solid, rather in-

flated. Spire relatively high, conical, 4-5 rather convex

whorls, shoulder rounded. Suture not very evident, par-

tially covered by the sculpture, which is formed of 15—

20 spiral cords, regularly ordered, rarely alternating with

smaller spiral cords. Spiral sculpture consisting of ribs

rounded in cross-section, all of similar width, densely

covered with imbricating, fine, and long lamellae. Axial

ribs 7—11, large, generally weak, crossing the spiral

cords. Siphonal canal short, narrow, open, moderately

curved. Aperture large, oval, representing about half of

the shell height, white or cream-white inside. Umbilicus

absent. Outer lip thin and crenulate. Shell color uni-

formly reddish- or pale-brown. Operculum oval, oblong,

horny, concentric, with lateral-terminal nucleus, reddish-

brown.

Type Material (Figures 1-8): Holotype (Figures 1-2),

30.6 X 20.4 mm, MNCN 15.05/46458; paratype A (Fig-

ures 3-4), 31.6 X 19.4 mm, MZB 31023; paratype B

(Figures 5-8), 27.3 xX 16.7 mm, CS-PM; paratype C,

27.4 X 18.2 mm, CS-PM; paratype D, 24.2 X 16.8 mm,

ER; paratype E, 26.8 X 16.6 mm, ER; paratypes A-E,

from type locality; paratype F, 30.9 < 18.5 mm, ER,

Malpica, Spain; paratype G, 23.5 X 16.3 mm, ER, Ca-

melle, Spain; paratype H, 24.8 x 17.3 mm, JT; paratype

I, 21.2 X 12.3 mm, JT; paratype L, 26.9 < 17.1 mm, JT;

paratype M, 21.4 X 12.3 mm, JT; paratype N, 31.2 x

18.0 mm, JT; paratype O, 18.5 X 12.8 mm, JT; paratype

P, 7.4 X 5.1 mm, CS-PM; paratypes H—P from Porto da

Baleeira, Sagres, Portugal; paratype Q, 30.2 < 18.1 mm,

ER, Lira A Coruna, Spain; paratype R, 26.4 X 17.2 mm,

ER, Malpica, Spain; paratype S, 27.3 X 16.2 mm, CS-

PM; paratype T, 25.1 X 16.2 mm, CS-PM; paratypes

Q-T from Lira A Corutia, Spain; paratype U, 39.2 x

23.2 mm, CS-PM, A Guarda, Spain; paratype V, 30.1 X

18.6 mm, CS-PM, type locality; paratype W, 29.8 X 17.4

mm, CS-PM; paratype Z, 25.8 X 16.2 mm, CS-PM;

paratypes W-Z from Malpica, Spain.

Type Locality: Camarifias, Galicia, Spain, northeast-

ern Atlantic Ocean, 15-50 m depth.

Distribution: Known from Galicia, Spain, to Algarve,

Portugal, in the Atlantic Ocean, and from Malaga and

Almeria (Alboran Sea, Spain) in the Mediterranean.

P. Mariottini et al., 2005

Page 111

Figures 1-8.

Habitat: Several live collected specimens were found

attached at the base of host cnidarians Calliactis paras-

itica (Couch) (a sea anemone).

Etymology: This species name is dedicated to our

friend Juan Trigo, who supplied some of the specimens

of the new species.

Molecular Sequencing: Given the often misleading

information conveyed by characters of shell morphology

in this group (Oliverio and Mariottini, 2001b), we veri-

Coralliophila trigoi new species. 1-2. Holotype, 30.6 < 20.4 mm, MNCN 15.05/46458. 3-4. Paratype A, 31.6 x

19.4 mm, MZB 31023. 5-8. Paratype B, 27.3 X 16.7 mm, CS-PM. From type locality, depth 15-50 m. Scale bars = 1 cm.

fied the validity of the new species using a molecular

approach. We sequenced the internal transcribed spacer

2 region (ITS2) of the nuclear rDNA and part of the

mitochondrial gene for 12S rDNA from individuals of

the new taxon and compared them with available se-

quences of C. meyendorffii, C. panormitana, and Cor-

alliophila brevis (Blainville, 1832), plus Coralliophila

neritoidea (Lamarck, 1816) as outgroup. Parsimony anal-

ysis of the aligned sequences of nuclear and mitochon-

drial DNA resulted in the tree reported in Figure 2

THE NAUTILUS, Vol. 119, No. 3

Figures 9-16.

(Calcara, 1845), 26.8 X 14.7 mm, FS, Punta del Carmen, Lanzarote Isl.,

The two related species C. meyendorffii and C. panor-

mitana, were more closely related to each other than to

the two specimens of the new species.

Other Material Examined: Coralliophila trigoi: 5

spec. from the type locality; 3 spec., Laxe; 1 spec. A

Guarda; 3 ee Camelle; 3 spec. Malpica, Galicia, pepe

ER; 2 spec: Sagres, Algarve, Portugal, CS-PM; 2 spec.

Almeria; 3 spec. Mar bella, Spain, MO; Coralliophila

meyendorffii: 10 spec. La Maddalena Isl., Sardinia Is-

land; 1 spec. San Pietro Isl., Sardinia Island; 6 spec. San-

ta Marinella; 3 spec. Ponza Isl.; 1 spec. Capo Palinuro;

3 spec. Le Castella; 5 spec. Elba Isl., Italy, CS-PM; 6

Shells of Coralliophila species. 9-10. Coralliophila squamosa (Bivona, 1838), 35

Galicia, Spain, 15-50 m depth. 11-12. Coieltepitte panormitana (Monterosato, 1869), 19.2 < 12.1 mm, CS-PM, Malaga, Spain, 40 m

depth. 13-14. Coralliophila trigoi new species, 27.2 X 18.1 mm, MO, en Spain, 30-50 m depth. 15-16. Coralliophila meyendorffii

9 X 20.9 mm, CS-PM, Camarinas,

Canary Islands, 20 m depth. Scale bars = 1 cm.

spec. Punta del Carmen, Lanzarote Isl., Canary Islands,

FS; Pee peroorntanees 5 spec. Marina di Ca-

merota, Italy; 1 spec Malaga, Spain; I sHee Portimao,

Portugal, CS-PM: Coralliophila squamosa: 23 spec. from

the type locality; 6 spec. Laxe; 8 spec. Camelle; 7 spec.

Malpica, Galicia, Spain, ER; 10 spec. Ria de Vigo, Ga-

licia, Spain, JT; 1 spec. Malaga, Spain; 1 spec. San Pietro

Isl., Sardinia Island; 1 spec. Ventotene Isl.; 1 spec. Ma-

rina di Camerata; 2 spec. Le Castella; 2 spec. Civitanova

Marche, Italy, CS-PM.

Remarks: Based on shell characters of the teleoconch

we take the conservative approach of conserving this

P. Mariottini et al., 2005

Page 113

Figures 17-26.

mm, CS- PM, Porto da Baleeira, Sagres, Portugal, 15-25 m depth.

mm, CS-PM, La Maddalena Isl., Sardinia, Italy, 1 m depth. ¢

Shells of Coralliophila species. 17-20. Coralliophila trigoi new species, 2

Portugal. 18, 19. Details of shell sculpture. 21. Coralliophila trigoi new species, detail of the ieravel whorls, paratype P, 7.4 X 5.1

22-25. Coralliophila meyendorffii (C

23, 24. Details of shell sculpture. 26. Coralliophila me yer ndorffti, detail

23.6 X 14.0 mm, CS-PM, Algarve,

Caleara, 1845), 28.6 * 16.0

of the larval whorls, specimen size 2.2 X 1.3 mm, CS-PM, La Maddalena Isl., Sardinia, Italy, 1 m depth. seats bars = 1 cm, except

for Figures 21, 26, scale bars = 200 wm.

species in the genus Coralliophila s. |., as traditionally

formulated (see also Oliverio, in press). The new species

was collected in Galicia together with the Atlantic-Med-

iterranean Coralliophila squamosa (Bivona, 1838). This

latter species is generally collected in the Mediterranean

Sea at depths ranging from 100 to 600 m (Figures 9—

10). The new taxon is ‘cleat ly conchologically distinguish-

able from all other eastern Atlantic aul Miadiiememenn

species of Coralliophilinae. Coralliophila trigoi shows a

certain resemblance with the Medtrenemcer Corallio-

phila panormitana (Monterosato, 1869), but the latter is

smaller, having a different sculpture that includes a larg-

er number of spiral cords (22-24), with smaller scales

(Figures 11-12).

ea trigoi is similar to Coralliophila meyen-

ae Calcara, 1845) (Figures 15-16, 22-26), but it dif

fers by its more rounded shape, by having the length/

width and length/aperture length ratios snl ei (1.60

and 1.68 vs. 1.72 and 1.86), by possessing a larger num-

ber of primary spiral cords (15-20 vs. 13-15), which are

narrower in width and differently sculptured, and by a

reddish- or pale-brown color (Coralliophila meyendorffii

is generally milky-white). Furthermore, the protoconch

I of Coralliophila trigoi shows a number of whorls (24%)

and a diameter (S00 um) different from the values on

Coralliophila meyendorffii (3% and 650-750 wm, re-

spectively; see Figures 21, 26). The main shell morpho-

logical dif (erences ibe een these two species are sum-

marized in Table Coralliophila meyendorffii is a lit-

toral, widely uate species, occurring in the Med-

iterranean Sea (Figures 22-26), along. the Atlantic

African coast and Canary Islands (Figures 15-16). It is

worth mentioning that Coralliophila meyendorffii preys

on a variety of amthoroena. ine nding the cnidarian Cal-

liactis parasitica (personal Ghasnvesone. Interestingly,

Coralliophila trigoi seems to be distributed along the

Atlantic coast ae Spain and Portugal, but vesiricted to

the Alboran Sea (Malaga, Almeria) within the Mediter-

ranean basin. The collecting depth is slightly deeper (50

m) than the bathymetric range (littoral) of Coralliophila

meyendorffi. Although the planktotrophic mode of de-

velopment (as snrdbiestied by the protoconch) of Corallio-

phila trigoi could bring about a wider geographic distri-

bution bern that amen known for the species, the

taxon has not yet been found in the Macaronesian Is-

lands and the West African coast.

Page 114

THE NAUTILUS, Vol. 119, No. 3

C. neritoidea

C. brevis

C. panormitana

100/97 C. meyendorffii

C. trigoi #1

100/100

C. trigoi #2

Figure 27. Cladogram of parsimony analysis for the studied

coralliophiline taxa. This topology was recovered under maxi-

mum parsimony analysis of the 12S + ITS2 dataset (either

including or excluding gap positions). Numbers at the branch

represent bootstrap support in MP analyses (gap included and

gaps as missing). Locality data and GenBank accession num-

bers (12S and ITS2) are reported in Table 1.

Coralliophila trigoi has been previously misidentified

as Coralliophila basilea (Rolén, 1983: 236, fig. 210; Ro-

lan, Lopez and Gutiérrez-Garcia, 1995: 30, fig.2), but it

is easily distinguishable from this taxon. Coralliophila

basilea has a more turreted shell and a coarser spiral

sculpture. Among the fossil coralliophiline records, the

only species that slightly resembles the new taxon is Cor-

alliophila burdigalensis (Tournouér, 1874), a species

from the Upper Oligocene and Lower Miocene of Aq-

uitaine (France); but the fossil species is smaller and has

a different shell outline and sculpture (Lozouet and

Renard, 1998: 173, figs. 2. 1-10).

Parsimony analysis of the aligned sequences of nuclear

and mitochondrial DNA resulted in the tree reported in

Figure 27. Accordingly, the two related species C. mey-

endorffii and C. panormitana were more closely related

to each other than to the two specimens of the new

species. The same results (not shown here) were ob-

tained including additional (yet shorter) sequences of C.

meyendorffi from other Mediterranean localities (Sar-

dinia, Sicily, and Southem Spain). This is a clear indi-

cation that the specimens of the new form constitute a

distinct, isolated gene-pool and strongly support our de-

cision to describe it as new.

ACKNOWLEDGMENTS

We would like express our deep gratitude to Mr. Juan

Trigo, Brion, A Coruna, Spain, for kindly sending us

Table 2. Comparison of shell characters between Corallio-

phila trigoi and C. meyendorffu.

Shell characters C. trigoi C. meyendorffui

Protoconch diameter 800 um 650-750 pm

Protoconch number of

whorls 25 3.5

Teleoconch primary spiral

cords 15-20 13-15

Teleoconch axial ribs lel 8-10

Length/width ratio 1.60 + 0.09 1.72 = 0.09

Length/aperture length

ratio 1.68 + 0.10 1.86 = 0.10

Size range 17.6-35.4 mm 19.5-34.3 mm

specimens of Coralliophila trigoi. Dr. Andrea Di Giulio

(Department of Biology, University of “Roma Tre’,

Rome, Italy) is acknowledged for SEM photographs,

which were carried out at the LIME (Inter-Department

Laboratory of Electron Microscopy, University of “Roma

Tre”). Sincere thanks are due to Dr. Antonio Bonfitto

(Zoological Museum, University of Bologna, Italy) for

generously providing bibliography. We are grateful to

Drs. Marco Oliverio and Maria Vittoria Modica (De-

partment of Animal and Human Biology, University of

“La Sapienza”, Rome, Italy) for help with molecular

work. M. Oliverio also provided valuable advices and dis-

cussion.

LITERATURE CITED

Bouchet, P. and A. Warén. 1985. Revision of the Northeast

Atlantic bathyal and abyssal Neogastropoda excluding Tur-

ridae (Mollusca, Gastropoda). Bollettino Malacologico,

Suppl. 1: 1-296.

Clover, P. 1982. Latiaxis catalog and illustrated check list of

the Coralliophilidae family. Privately printed, i—ii, 18 pls.

+ 18 unnumbered text pages.

Harasewych, M. G., S. L. Adamkewicz, J. A. Blake, D. Saudek,

T. Spriggs and C. J. Bult. 1997. Neogastropod phylogeny:

a molecular perspective. Journal of Molluscan Studies 63:

327-351.

Iredale, T. 1912. New generic names and new species of ma-

rine Mollusca. Proceedings of the Malacological Society

of London 10: 217-228.

Lamarck, J. B. P. A. de, M. 1816. Liste des objets représentés

dans les planches de cette livraison. “Vingt-troisiéme par-

tie: mollusques et polypes divers, par M. Lamarck. In:

Bruguiére, J. G., M. J. B. P. A. de Lamarck and B. de St.

Vincent. (1782-1832) Tableaux encyclopédique et méth-

odique des trois régnes de la nature”. 1-16, pls 391-488,

431 bis, 432 bis®.

Lozouet, P. and P. Renard. 1998. Les Coralliophilidae, Gastro-

poda de V'Oligocéne et du Miocéne inférieur d’Aquitaine

(sud-ouest de la France): systématique et coraux hétes.

Geobios 31 2: 171-184.

Kosuge, S. and M. Suzuki. 1985. Illustrated catalogue of La-

tiaxis and its related groups. Family Coralliophilidae. In-

stitute of Malacology of Tokyo, Special Publications 1: 1—

83.

P. Mariottini et al., 2005

Oliverio, M. 1989. Famiglia Coralliophilidae Chenu, 1896 in

Mediterraneo. La Conchiglia 246-249: 48-54.

Oliverio, M. In press. Coralliophilinae (Neogastropoda: Muri-

cidae) from the South West Pacific. In: P. Bouchet and V.

Heros (eds.) Tropical Deep-Sea Benthos. Memoires du

Muséum national d’Histoire naturelle, Paris.

Oliverio, M. and P. Mariottini. 200la. A molecular framework

for the phylogeny of Coralliophila and related muricoids.

Journal of Molluscan Studies 67: 215-224.

Oliverio, M. and P. Mariottini. 2001b. Contrasting morpholog-

ical and molecular variation in Coralliophila meyendorffii

(Muricidae, Coralliophilinae). Journal of Molluscan Stud-

ies 67: 243-246.

Oliverio, M., M. Cervelli and P. Mariottini. 2002. ITS2 rRNA

evolution and its use in the phylogeny of muricid neogas-

Page 115

tropods (Caenogastropoda, Muricoidea). Molecular Phy-

logenetics and Evolution 25: 63-69.

Richter, A. and A. A. Luque. 2002. Current knowledge on Cor-

alliophilidae (Gastropoda) and phylogenetic implication of

anatomical and reproductive characters. Bollettino Mala-

cologico, Suppl. 4: 5-18.

Rolan, E. 1983. Moluscos de la Ria de Vigo I. Gasterépodos.

Thalassas, Anexo 1: 1-383.

Rolan, E., D. Lépez and G. Gutiérrez Garcia. 1995. Nuevas

citas de moluscos de Galicia. Noticiario SEM 25: 20-21.

Swofford, D. L. 2002. PAUP*. Phylogenetic Analysis Using

Parsimony (*and Other Methods). Version 4 [1998], beta

4.0b10 [2002]. Sinauer Associates, Sunderland, Massachu-

setts.

Vaught, K. C. 1989. A classification of the living Mollusca.

American Malacologists Inc., Melboume, 143 pp.

THE NAUTILUS 119(3):116-117, 2005

Book Review

Page 116

Land Snails of British Columbia

Forsyth, Robert G. 2004. Land Snails of British

Columbia. Royal BC Museum Handbook. Royal BC

Museum, Victoria, 192 pp. ISBN 0-7726-5218-X. Black

and white illustrations, plus a section of color photo-

graphs.

Even as “cutting edge” systematics shifts its focus

from mor phological to molecular data sets, the need for

identification manuals based on real-world field marks

and accessible characters has never been greater. When

zoologists with landcare agencies or environmental

consultants ask for references on land snails, it hardly

helps to refer them to ponderous, outdated, and hard to

find tomes such as Pilsbry (1939-1948). So, manuals like

Land Snails of British Columbia are a welcome resource.

Land Snails of British Columbia is the latest in a Royal

BC Museum series of natural history books that began in

1942. It is the first work to describe and illustrate the 92

species of terrestrial mollusks known to occur in British

Columbia, Canada. Because all of the species treated

occur outside that province as well, its utility extends

beyond those borders. Identification aids include

a paragraph of physical description, usually brief, to

the point, and adequate; a sentence or two of

comparison with other, potentially confusing species;

clear illustrations (the Ifine- and-stipple drawings of many

of the smaller kinds are noteworthy); and keys to, first,

the genera of snails and slugs, and then to the species in

each multi-species genus. For slugs, where the di-

agnostic characters are often internal, excellent, pre-

donninerntley original, drawings of the distal genitalia are

provided. (The simple dissection needed to access these

features should be within the range of all to whom this

book is directed.) The language of the descriptions is

simple and intelligible, and a glossary of technical terms

is included. The. descriptive material is followed by

a summary of each species’ distribution, concise com-

ments on natural history, etymology of the generic and

specific name, remarks that, among other things, update

the species’ taxonomic history (e.g., since Pilsbry, 1939-

1948) or point out special conservation status, and selected

references pointing to the extensive and well researched

bibliography. All of this adds up to a highly functional

volume that will serve its intended ncaghenee well.

A six-page section of color photographs of living snails

and slugs, mainly by Kristiina Ovaska, is not only

attractive but also helpful, particularly in showing the

habitus of living slugs, something not always well

conveyed by drawings.

The information is amalgamated (the author's word)

from published literature, This own. observations, scien-

tific collections, and personal communications with

other workers. The distributions are based in part on

unpublished records from the Royal BC Museum and

other collections. Specific localities are not cited; three

online resources by the same author (Forsyth, 1999,

2005a, 2005b) provide more detail for specific regions.

A significant innovation is the introduction of Euconu-

lus (Euconulus) praticola (Reinhardt, 1883) 2 Pe North

American fauna. Its distinctness from E. ) fulvus

(Miiller, 1774) is well documented by cae of shell

and external anatomy. The former catchall “Zonitidae” is

correctly parsed into Pristilomatidae, Gastrodontidae, and

Daudebardiidae for regional genera, in keeping with

recent work by Hausdorf and others. Monadenia, iconic

of northwest American forests, is correctly assigned to

Bradybaenidae rather than the redundant and poorly

ar cued Monadeniidae of Nordsieck (1987) and Schileyko

(1996), In these and other ways, the author shows himself

to be well “booked up” on the current and evolving

literature of land snail systematics.

Overall, the taxonomy is simplified, appropriately for

a general interest manual. Synonyms are merely listed,

not referenced, as few potential users will need that

entry into the primary literature. Subspecies are treated

in the remarks, not in the major taxon headings or in the

taxonomic checklist (pp. 23-28). If experience in Cali-

fornia is any guide, upon further study some—perhaps

most—nominal subspecies will be shown to be species;

and examination of data not available to prior mono-

graphers may uncover local endemics, possibly in former

glacial refugia such as the Queen Charlotte Islands. For

the present, however, this handbook provides a reliable

source of biodiversity information. Author Forsyth is

fully aware of the dynamic character of taxonomic

analysis and astutely states (p. 2), “this book is a work in

progress.” By clearly showing the state of our knowledge

at this time, he has contributed greatly to that process.

LITERATURE CITED

Forsyth, R. G. 1999. Terrestrial Gastropods of the Columbia

Basin, British Columbia. Royal BC Museum, Victoria. http://

www. livinglandscapes.be.ca/cbasin/molluses/contents.html

[html version] or http://www. livinglandscapes.be.ca/cbasin/

molluscs/pdf/mollusc3.pdf [pdf version]. (Accessed August

20, 2005.)

Forsyth, R. G. 2005a. Tewestiaall Gastropods of the Peace

River Region — Northern Rockies of British Columbia.

Royal BC Museum, Victoria. http://livinglandscapes.be.

ca/prnr/prnr_snails/index.html [html version] or http://

livinglandscapes.be.ca/pmr/snails/prnr_snails.pdf [pdf ver-

sion]. (Accessed August 20, 2005.)

Forsyth, R. G. 2005b. Terrestrial Gastropods of the Upper

Fraser Basin of British Columbia. Living Landscapes,

Royal BC Museum, Victoria. http://livinglandscapes.be.ca/

Book Review, 2005

Page 117

upperfraserbasin/utb_snails/index.html [html version] or

http://livinglandscapes.be.ca/upperfraserbasin/snails/

ufb_snails.pdf [pdf version]. (Accessed August 20, 2005.)

Nordsieck, H. 1987. Revision des Systems der Helicoidea

(Gastropoda: Stylommatophora) Archiv fiir Mollusken-

kunde 118: 9-50.

Pilsbry, H. A. 1939-1948. Land Mollusca of North America

(north of Mexico). Academy of Natural Sciences of

Philadelphia, Monograph 3, 1 (1): i-xvii + 1-573 + i-ix

(1939); 1 (2): ievi + 574-994 + i-ix (1940): 2 (1): i-viii + 1—

520 + i-ix (1946): 2 (2): iexlviii + 521-1113 (1948).

Schileyko, A. A. 1996. Guamampa n.g. (Gastropoda,

Pulmonata), a bradybaenid land snail with monadeniid

characters. Bulletin du Muséum National d’Histoire

Naturelle, Section A, Zoologie, Biologie et Ecologie

Animales 18: 401-408.

Barry Roth

745 Cole Street

San Francisco, CA 94117 USA

barryroth@earthlink. net

Sponsored in part by the State of Florida, Department

of State, Division of Cultural Affairs, the Florida Arts

Council and the National Endowment for the Arts.

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& L, Volume 119, Number 4

LO ( December 30, 2005

Nat4 ISSN 0028-1344

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CONTENTS

Ellen E. Strong

Richard L. Squires

LouElla R. Saul

M. G. Harasewych

Yuri I. Kantor

Sven N. Nielsen

Patricia Miloslavich

Ana Karinna Carbonini

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Eliézer de Carvalho Rios

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Volume 119, Number 4

December 30, 2005

ISSN 0028-1344

A morphological reanalysis of Pleurocera acuta Rafinesque, 1831,

and Elimia livescens (Menke, 1830) (Gastropoda: Cerithioidea: Pleuroceridae) 119

New Late Cretaceous (Santonian and Campanian) gastropods from

C@alitomuayandyBajay Calitomiay Mex Conmne nen en ern ere 133

Daffymitra lindae, a new genus and species of Volutomitridae

(Neogastropoda) from the Bellingshausen Abyssal Plain ............... 149

Exilia alanbeui, a new species from the Neogene of central Chile:

the first record of Exilia (Gastropoda: Ptychatractidae) from South America 153

Spawn of Amphissa sp. and Cosmioconcha sp. (Caenogastropoda:

Columbellidae) from the Colombian Caribbean..................... 157

Pterorytis pacanana new species (Gastropoda: Muricidae): circumstantial

evidence for late Pliocene El Nifio events in southern Peru ............ 164

A new species of Falsimargarita (Gastropoda: Vetigastropoda: Trochidae)

Luiz Ricardo L. Simone fionm tae Sorin Adami OGM . oo occ ccocscacscnn occa nanoavoboe. 169

LEY RELIETIN 5 0-0-6 1dc0:i0-0fouesoe.ce 6. atin Aly B16: OREN ce LAH bar A Rote eee Ce Ne RM eg 0S a Be el inca jeans POT Ny eer Be Re 163

INORG so ace Biotetleeeo ta souac ta RUSS Breese tec er OEE Ths GOH UU een ROR ey Aa Foal Age Me ees 173, 174

ND Te 8 be

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THE NAUTILUS 119(4):119-132, 2005

Page 119

A morphological reanalysis of Plewrocera acuta Rafinesque,

1831, and Elimia livescens (Menke, 1830)

(Gastropoda: Cerithioidea: Pleuroceridae)

Ellen E. Strong

Department of Invertebrate Zoology

National Museum of Natural History

Smithsonian Institution

P.O. Box 37012-MRC 163

Washington, DC 20013-7012 USA

[email protected]

ABSTRACT

Pleurocera acuta and Elimia livescens have been the subject of

several anatomical and ecological studies and are two of the

most thoroughly documented species of North American

Pleuroceridae. Yet significant gaps still remain in our un-

derstanding of their structure. Consequently, the anatomy of

these two species is re-described, allowing a re-interpretation

of pallial oviduct homologies; features not previously portrayed

in the literature (midgut and kidney) are newly described.

These taxa are characterized by the presence of an ovipositor,

a kidney with a subdivided intemal lumen that invades the

pallial roof, a prostate with a highly folded anterior spermato-

phore-forming region, and a pallial oviduct with spermato-

phore bursa but lacking a seminal receptacle. This analysis

verifies the degree of similarity between the two species, but

a number of differences were identified including features of

the ovipositor, pallial oviduct, prostate, anterior esophagus,

midgut, kidney, pericardium and nervous system. Comparison

to other pleurocerids confirms that species distributed in

Western North America (Juga) and Asia (Hua, Semisulcospira)

share the presence of a seminal receptacle—a feature that is

lacking in all described Eastern North American species.

INTRODUCTION

The freshwater Pleuroceridae Fischer, 1885, comprises

one of the most species-rich assemblages of limnic

mollusks occurring in North America and Eastern Asia.

In North America, they are represented by eight genera

(Athearnia Morrison, 1971, Elimia H. and A. Adams,

1854, Io Lea, 1831, Juga H. and A. Adams, 1854,

Leptoxis Ratinesque, 1819, Lithasia Haldeman, 1840,

Pleurocera Rafinesque, 1818, and the extinct Gyrotoma

Shuttleworth, 1845) and estimates of 159 species

currently considered valid; of these, 34 are extinct and

59 are listed as critically imperiled (G1) or imperiled

(G2) (Johnson et al., 2005).

The current concept of the family Pleuroceridae (e.g.

Bouchet and Rocroi, 2005) can be traced to the works of

Thiele (1928, 1929) who recognized 6 subfamilies within

the heterogeneous “Melaniidae” (an invalid name for

Thiaridae Gill, 1871), including the Pleurocerinae. An

alternative, highly polyphyletic view promoted by

Morrison (1954) caused great confusion for more than

four decades conceming the extension and indepen-

dence of the Pleuroceridae and Pachychilidae P. Fischer

and Crosse, 1892 (e.g. Ponder and Warén, 1988). Recent

work on the systematics of limnic lineages within the

Cerithioidea Fleming, 1822 (e.g. Glaubrecht, 1996,

1999: Kohler et al., 2004) has resolved some of this

confusion and supports the distinctiveness of these

families on both morphological and molecular grounds.

However, molecular data (Lydeard et al., 2002) do not

support monophyly of the Pleuroceridae as currently

defined, but suggest that a clade of eastern North

American species are more closely related to Melanopsis

(Melanopsidae H. Adams and A. Adams, 1854) than to

a clade of western North American (Juga) and eastern

Asian pleurocerids (Semisulcospira Boettger, 1886, Hua

Chen, 1943).

As noted by Woodard (1934), anatomical data are

critical in refining the phylogenetic relationships and

classification of these species. However, all taxonomic

treatments of the family have been based primarily on

conchological characters in the absence of a broad

comparative understanding of morphology. Although the

subject of numerous ecological studies (e.g. Dillon,

2000) and comparably many descriptions exist for the

radula, operculum and life history of North American

pleurocerids, surprisingly little is known about their

basic biology and anatomy. Scant information on the

anatomy (besides radula and operculum) has been

provided for Elimia laqueata (Say, 1829) (Woodard,

1934), E. potosiensis (Lea, 1841) (Jones and Branson,

1964) and Pleurocera canaliculata (Say, 1821) (Magru-

der, 1935b); among western North American forms, only

reproductive anatomy is known for several species

(Prozorova and Raschepkina, 2004). In addition, the

Page 120

THE NAUTILUS, Vol. 119, No. 4

classically cited work of Dazo (1965) on the natural

history, ecology, distribution and anatomy of Plewrocera

acuta Rafinesque, 1831 and Elimia livescens (Menke,

1830) has long stood as the most comprehensive

morphological study of any pleurocerid gastropod. As

such, that study has formed the basis for hypotheses of

homology in recent higher-order phylogenetic studies

based on morphology (e.g. Houbrick, 1988; Glaubrecht,

1996). Yet, Dazo’s study was completed long before our

present understanding of the structure, function and

homologies of cerithioidean reproductive anatomy was

in place, particularly through the work of R. S. Houbrick

on marine species. We also have a much more thorough

understanding of midgut structure and its utility in

revealing phylogenetic affinities of cerithioideans (Glau-

brecht and Strong, 1999; Strong and Glaubrecht, 1999,

unpublished data).

Given the persistent paucity of anatomical data

available for the family, the goal of this contribution is

to reevaluate the morphology and putative homologies of

the two species described by Dazo, and to place them

within the emerging framework now available for limnic

cerithioideans. This is critical for ongoing morphological

and phylogenetic studies of cerithioidean gastropods,

and in particular for clarifying the monophyly and

systematic affinity of the family Pleuroceridae.

MATERIALS AND METHODS

This study is based on collections of individuals of

Pleurocera acuta and Elimia livescens living sympatri-

cally in the Mukwonago River at the outflow from Lower

Phantom Lake, Mukwonago, Waukesha County, Wis-

consin (42°51.402 N; 88°19.767 W). Populations were

sampled in May and June; only reproductively mature

individuals were used for observations of reproductive

anatomy. Individual specimens were cracked, preserved

in 95% ethanol and were not relaxed. Voucher material

is deposited in the National Museum of Natural History

in Washington (USNM).

As stated above, Dazo (1965) provided a rather

detailed account of external features (operculum, shell,

ovipositor, color patterns of the head-foot) as well as of

the radula and all internal organ systems (alimentary,

nervous, excretory, respiratory, vascular, and reproduc-

tive systems). Baker (1928) also provided information on

the operculum, jaw, radulae, and external anatomy of the

two species; additional scattered accounts relating to the

radula, external anatomy and life history are also

available (e.g. Jewell, 1931; Howe, 1938; Goodrich,

1945). As such, the following reanalysis emphasizes

anatomy of the soft parts. As the two species are

overwhelmingly similar, a thorough account is provided

for Pleurocera acuta — the type species of the type genus

for the family; only details that differ are noted for

Elimia livescens. However, the internal kidney structure

of P. acuta, while displaying the same configuration as E.

livescens, is too occluded with excretory tissue to allow

a clear representation. As such, only the internal

structure for the latter species is illustrated.

Specimens were examined using a Leica MZ 12,5

binocular microscope with camera lucida; visualization

of structures was enhanced through the use of aqueous

toluidine blue. Typically four to five specimens were

examined for each organ system investigated, especially

for those systems showing high levels of individual and/

or seasonal variation (i.e. reproductive system); a mini-

mum of two individuals were examined for particularly

complex structures (i.e. midgut, nerves). Descriptions of

midgut morphology are given with the stomach opened

dorsally and the style sac uppermost. Unless otherwise

indicated, an incision is made along the extreme right,

upper and lower margins, and the roof deflected laterally

to the left. Terminology follows Strong and Glaubrecht

(2002, 2003).

RESULTS

Pleurocera acuta Rafinesque, 1831

Material Examined: Wisconsin: Mukwonago River:

USNM 1081522, 1081524.

External Anatomy: Operculum sub-ovate, corneous,

dark reddish-brown in color, with 3.5 whorls; paucispiral

with large, eccentric nucleus of approximately 2.75 to 3

whorls (Figure 1). Final whorl moderately inflated.

Nucleus occupying slightly under 1/2 (~43%) of total

length.

Foot ovate with narrow propodium; anterior pedal

gland opening along anterior margin (Figure 2, ap).

Ovipositor (ovp) located on side of neck below right

cephalic tentacle. Ciliated egg groove extending short

distance up side of neck from ovipositor, shallowing past

mantle margin; groove fading near anal aperture. Two,

partially juxtaposed, parallel folds, extending into

ovipositor pore from aperture (*); one fold at upper

posterior edge, second at lower anterior edge of

ovipositor (when viewed laterally), forming obliquely

flattened H-shaped lumen. Folds unequal in size;

anterior fold along floor significantly larger than

posterior fold along roof. Ovipositor pore expanding

medially into head-foot, then curving and narrowing

slightly posteriorly. Two vertical limbs of “H” unequal

such that posterior limb forming rather narrow channel

along posterior wall. Anterior limb inflated and expand-

ing medially into foot, forming large flattened, sub-

triangular chamber. Folds diminishing toward blind tip

of pore. Short grooved tract extending ventrally from

ovipositor aperture toward foot sole, but not reaching

edge of foot.

Most individuals with straight osphradium, but some

with curved anterior tip (Figure 3, os). Hypobranchial

gland highly developed with pendulous, transverse folds

(Figure 4, hg).

Alimentary System: Forecur. Buccal mass short and

stout, extending to base of cephalic tentacles (Figure 3,

E. E. Strong, 2005 Page 121

Figures 1-4. Anatomy of Plewrocera acuta. 1. Operculum. 2. Ovipositor and egg groove. Right lateral view of side of foot. Asterisks

(*) indicate folds extending into ovipositor. 3. Mantle cavity and anatomy of cephalic hemocoel. Dorsal view, anterior is below.

Hypobranchial gland removed for clarity. 4. External view of organs in visceral mass. Dotted line indicates extent of pericardium

under main kidney chamber. Abbreviations: ap, anterior pedal gland; b, bladder; bm, buccal mass; em, columellar muscle; ct,

ctenidium; e, esophagus; hg, hypobranchial gland; int, intestine; kd, main kidney chamber; me, mantle edge; nr, circum-esophageal

nerve ring; op, operculum; os, osphradium; ov, ovary; ovp, ovipositor; per, pericardium; po, pallial oviduct; ps, peri-intestinal sinus;

r, rectum; rt, buccal mass retractor muscle; sg, salivary gland; sn, snout; sp, supra-esophageal ganglion; ss, style sac; sto, stomach.

Scale bars = 1 mm.

Page 1122

THE NAUTILUS, Vol. 119, No. 4

bm). Odontophore occupying majority of buccal cavity

with small, glandular subradular organ protruding before

radula. Small jaws present at anterior ends of dorsal

folds; epithelium of buccal cavity overlying dorsal folds

glandular (stippled region). Shallow, non-glandular

buccal pouches extending underneath dorsal folds

adjacent to buccal ganglia at rear of buccal cavity.

Radular sac short, curving upward behind base of buccal

mass. Robust buccal retractors (rt) inserting onto lateral

walls of cephalic hemocoel adjacent to cerebral ganglia

(nr). Short, glandular mid-ventral fold forming small flap

just behind odontophore in anterior esophagus, flanked

laterally by two ventro-lateral folds. Ventro-lateral folds

converging short distance behind mid-ventral fold and

continuing through mid-esophagus (e). Mid-esophagus

long, bearing paired longitudinal ventral and dorsal

folds. Epithelium between dorsal and ventral folds

weakly glandular and irregularly striated; septate esoph-

ageal gland lacking. Dorsal and ventral folds converging

and fusing at distal end of mid-esophagus before

continuing into posterior esophagus and_ subdividing

into numerous folds equal in height. Long, tubular

salivary glands (sg) opening dorso-laterally to buccal

cavity alongside odontophore, passing through circum-

esophageal nerve ring (nr), almost reaching posterior

esophagus.

Mipcut. Esophagus opening under ledge on left side of

midgut floor (Figure 5, e). Marginal fold (mf) extending

anteriorly from esophageal aperture alongside major

typhlosole (t1), then turning posteriorly bordering right

margin of sorting area (sa). Groove present along

midline of marginal fold (mf) for much of its length;

groove fading proximally and distally. Sorting area

elongate- triangular, tapering posteriorly; posterior tip

curving slightly to the left around wedge-shaped sorting

area pad (sap). Accessory marginal fold (amf) emerging

from esophageal aperture, paralleling marginal fold and

curving around posterior tip of sorting area; fold

bifurcating at posterior end of gastric chamber to form

two folds. Fine parallel striations extending anteriorly

from esophagus up face of major typhlosole. Midgut roof

to the left of sorting area coarsely folded and cuticular-

ized (eu). Gastric shield (gs) small and delicate, strongly

concave, with narrow, tubular posterior end and more

flaring, flattened anterior end; shield continuous with

cuticle of stomach roof and crystalline style pocket (p).

Glandular pad (gp) large and broadly rounded. Cres-

centic ridge (er), bounding deep crescentic groove,

extending from esophageal aperture and fusing to right

side of glandular pad. Paired digestive gland ducts (dd)

opening to deep pocket near proximal tip of crescentic

ridge. Shallow caecum (ec) extending ventrally under

glandular pad behind gastric shield. Single longitudinal

fold (cf) extending from caecum (e) around posterior

end of gastric chamber. Prominent fold (u) extending

from right side of style sac lip, along floor of crystalline

style pocket, to base of major typhlosole; fold bounding

u-shaped depression below lip of style sac (ss). Style sac

and intestinal groove communicating along entire length.

Crystalline style present.

Hinpcur. Proximal intestine (Figure 4, imt) passing

below distal tip of style sac (ss), then extending

posteriorly alongside style sac to main gastric chamber

(sto). Intestine curving anteriorly, with broad loop

overlying proximal style sac. Intestine extending under

posterior end of main kidney chamber (kd), entering

pallial roof alongside bladder (b) and pallial gonoduct

(Figure 3, po), continuing forward to papillate anus near

mantle margin (r).

Reno-pericardial System: Kidney comprising three

interconnected chambers (Figure 4, 17, kd, b). Main

chamber (kd) lying along dorsal surface of body whorl,

anteriorly surrounding pericardium (Figure 4, per),

crossing axis of body from right to left and extending

short distance into pallial roof at base of mantle cavity.

Chamber occluded anteriorly (within pallial roof) with

excretory tubules. Posteriorly, main chamber with small,

narrow lumen, dorsally enclosing intestine. Second

chamber (see exposed chamber in Figure 17) extending

between pericardial chamber to right body wall below

intestine, forming small bladder (Figure 4, b). Chamber

mostly occluded by vertical sheets of excretory tissue

radiating from afferent renal vessel (see Figure 17, arv),

and communicating to mantle cavity via large nephro-

pore (np). Sheets of excretory tissue branching and

anastomosing, and fusing to right lateral floor, roof and

walls; vertical sheets highly branched anteriorly and

forming comparatively dense honeycomb of excretory

tissue. Bladder penetrating connective tissue along right

side of body, short distance into mantle roof. Excretory

tissue separating small ventral chamber below, within

pallial portion (dotted line). Size of ventral chamber, as

well as branching pattern and number of excretory

sheets of tissue variable between individuals. Small

aperture just behind afferent renal vessel connecting

main chamber and bladder (arrow). Nephridial gland

absent.

Pericardium voluminous (Figure 4, per), extending to

right side of body (dotted line).

Nervous System: Circum-esophageal nerve ring

(Figure 3, nr) lying immediately behind buccal mass,

at base of cephalic tentacles. Cerebral ganglia (Figure 6,

ce) connected by short, stout commissure, each pro-

ducing seven nerves (optic, statocyst, tentacular, and

four labial nerves). Buccal connectives short, innervating

buccal ganglia lying ventro-laterally at base of buccal

cavity immediately behind buccal retractor muscles.

Pleural ganglia (pl) lying behind and below cerebral

ganglia connected to cerebral ganglia by short, thick

connectives. Pedal ganglia (pe) with two prominent

anterior nerves and five smaller accessory nerves. Small

statocysts (st) with approximately 10-15 statoconia

present dorsally alongside pedal ganglia behind pedal

connectives. Sub-esophageal ganglion (sb) joined to left

pleural ganglion by thickened connective (co); connec-

E. E. Strong, 2005

Page 123

Figures 5-6. Anatomy of Plewrocera acuta. 5. Midgut anatomy. Dorsal view, anterior is uppermost. 6. Circum-esophageal nerve

ring. Frontal view on the left, right lateral view on the right. Abbreviations: amf, accessory marginal fold; e, caecum; ef, caecal fold;

ce, cerebral ganglion; co, thickened connective between left pleural and sub-esophageal ganglia; er, crescentic ridge; cu,

cuticularized region of stomach roof; dd, duct of digestive gland; e, esophageal aperture; gp, glandular pad; gs, gastric shield; int,

intestine; mf, marginal fold; p, crystalline style pocket; pe, pedal ganglion; pl, pleural ganglion; pn, pallial nerve from left pleural

ganglion; sa, sorting area; sap, sorting area pad; sb, sub-esophageal ganglion; sp, connective to supra-esophageal ganglion; ss, lip of

style sac; st, statocyst; tl, major typhlosole; u, u-shaped fold; z, zygoneury. Scale bars = 1 mm.

tive producing 1-3 small nerves (n=2). Zygoneury (z)

formed between sub-esophageal and right pleural

ganglia. In addition to zygoneury (z), sub-esophageal

ganglion (sb) producing one other prominent nerve and

connectives to the left pleural (pl) and visceral ganglia.

Long connective uniting right pleural and supra-

esophageal ganglia (Figures 3, 6, sp), the latter lying

on left side mantle floor near midline of osphradium.

Dialyneury formed between pallial nerve of left pleural

ganglion and osphradial nerve of supra-esophageal

ganglion at junction of mantle roof and floor. Single

visceral ganglion present between pericardium and

kidney at base of mantle cavity, above posterior

esophagus on the right. Ganglion producing two

prominent nerves.

Reproductive System: Mate. Narrow vas deferens

(Figure 8, vd) emerging ventrally from testes, continuing

forward along ventral midline of whorl. Short, distal

portion of vas deferens thickened and forming straight

seminal vesicle. Vas deferens narrowing and curving

dorsally to enter posterior end of prostate (Figure 7, pr)

at base of mantle cavity. Prostate glandular, forming

flattened tube, opening to mantle cavity through broad

slit along entire length except for a short fused segment

at base of mantle cavity (arrows). Lateral lamina

epithelium thin along aperture; short distance into

lumen, glandular tissue forming flattened longitudinal

shelf. Glandular tissue diminishing in thickness at

anterior and posterior ends of gonoduct. Shelf undercut

along much of its length by flattened sinus (Figure 8,

dotted line). Glandular tissue forming central rounded

mass flanked by shallow trough along gonoductal groove.

Medial lamina unevenly glandular, with irregular troughs

and ridges; glandular tissue diminishing in thickness

anteriorly and posteriorly (Figure 9). Epithelium of

medial lamina posterior 1/5 smooth and concave, partially

separated from anterior region by prominent curving

ridge; ridge embracing corresponding rounded glandular

mass in lateral lamina. Ridge continuous anteriorly with

curving trough formed by opposing ridges of tissue.

Trough fading anteriorly, and becoming flanked by short,

shallow flap running along gonoductal groove.

FemaLe. Gonad (Figure 4, ov) dorsally surrounding

digestive gland (except for a narrow ventral strip) from

tip of visceral mass to posterior end of midgut (sto).

Oviduct emerging ventrally from ovary. Renal oviduct

(Figures 10, 11, ovi) deflected dorsally behind mantle

cavity before entering base of glandular pallial oviduct.

Pallial oviduct, with proximal albumen (ag) and distal

capsule glands (eg). Albumen and capsule glands

forming narrow bands with opposing flattened surfaces

bounding gonoductal groove (Figure 11, gg); non-

glandular portions of medial and lateral laminae formed

by thickened connective tissue (ct). Proximal segment of

albumen gland at base of mantle cavity under pallial

kidney extension rather short and straight (Figures 10,

Page 124

THE NAUTILUS, Vol. 119, No. 4

Figures 7-11. Reproductive anatomy of Plewrocera acuta. 7. External, left lateral view of prostate. Anterior is to the left. Arrows

indicate extent of opening to gonoductal groove. 8. Internal aspect of prostate lateral lamina. Dotted line designates extent of sinus

under glandular shelf. 9. Internal aspect of prostate medial lamina. Anterior is to the right. Note parallel folds and groove at anterior

end, representing presumptive spermatophore forming region. 10. External, left lateral view of pallial oviduct. Anterior is to the left.

Arrows indicate extent of opening to gonoductal groove. 11. External, right lateral view of pallial oviduct. Anterior is to the right.

Abbreviations: ag, albumen gland; eg, capsule gland; ct, connective tissue; gg, gonoductal groove; ovi, renal oviduct; pr, prostate;

sg, sperm gutter; spb, spermatophore bursa; vd, vas deferens. Scale bars = 1 mm.

11, ag). Anterior to pallial kidney chamber, albumen

gland curving under distal tip of bursa to the right, then

arcing dorsally. Capsule gland comprising approximately

anterior 1/4 of pallial oviduct; externally capsule gland

not inflated (Figure 10, eg). Broad aperture along entire

length of pallial oviduct except for a short fused segment

at base of mantle cavity (Figure 10, arrows). Above

aperture, sperm gutter (sg) opening in medial lamina at

anterior tip of pallial oviduct and deepening posteriorly;

gutter leading to short, blind spermatophore bursa

(spb). Seminal receptacle absent.

Elimia livescens (Menke, 1830)

Material Examined: Wisconsin: Mukwonago River:

USNM 1081521, 1081523.

External Anatomy: Operculum ovate, corneous, dark

reddish-brown in color, with three whorls; paucispiral

with small, basal nucleus of approximately 2.5 whorls

(Figure 12). Last whorl expanding rapidly. Nucleus

comprising approximately 1/5 of total length.

Foot broad and rounded, with wide propodium and

long anterior pedal gland along anterior margin

(Figure 13, ap). Ciliated egg groove extending short

distance up side of neck from ovipositor; groove fading

near base of tentacle. Parallel folds extending into

ovipositor pore from aperture (*) roughly equal in size.

Grooved tract extending ventrally from ovipositor to sole

of foot at junction of propodium and mesopodium, just

behind termination of anterior pedal gland.

Osphradium with curved anterior tip; some individu-

als with mostly straight osphradium (Figure 14, os).

Hypobranchial gland well developed with deep, trans-

verse folds (Figures 14, 15, hg).

Alimentary System: FoREGUT. Epithelium between

dorsal and ventral folds glandular and irregularly to

transversely striated. Dorsal and ventral folds diminish-

E. E. Strong, 2005 Page 125

Figures 12-15. Anatomy of Elimia livescens. 12. Operculum. 13. Ovipositor and egg groove. Right lateral view of side of foot.

Asterisks (*) indicate folds extending into ovipositor. 14. Mantle cavity and anatomy of cephalic hemocoel. Dorsal view, anterior is

below. 15. External view of organs in visceral mass. Dotted line indicates extent of pericardium under main kidney chamber.

Abbreviations: ap, anterior pedal gland; b, bladder; bg, buccal ganglion; ct, ctenidium; e, esophagus; hg, hypobranchial gland; int,

intestine; kd, main kidney chamber; me, mantle edge; nr, circum-esophageal nerve ring; op, operculum; os, osphradium; ov, ovary;

Ovp, ovipositor; per, pericardium; po, pallial oviduct; r, rectum; rt, buccal mass retractor muscle; sg, salivary gland; sp, supra-

esophageal ganglion; ss, style sac; sto, stomach; t, cephalic tentacle. Scale bars = 1 mm.

Page 126

THE NAUTILUS, Vol. 119, No. 4

Figures 16-17. Anatomy of Elimia livescens. 16. Midgut anatomy. Dorsal view, anterior is uppermost. 17. Kidney anatomy.

Internal view of bladder. Lateral view, anterior is to the right. Right wall of bladder and adhering sheets of excretory tissue removed

to reveal interior; cross-hatching indicates cross-section through sheets of tissue. Arrow indicates opening in septum allowing

communication between bladder and main kidney chamber. Dotted line anteriorly indicates extent of ventral chamber.

Abbreviations: amf, accessory marginal fold; arv, afferent renal vessel; c, caecum; ef, caecal fold; er, crescentic ridge; cu,

cuticularized region of stomach roof; dd, duct of digestive gland; e, esophageal aperture; gp, glandular pad; gs, gastric shield; int,

intestine; kd, main kidney chamber; mf, marginal fold: np, nephropore; p, crystalline style pocket; sa, sorting area; sap, sorting area

pad; ss, lip of style sac; tl, major typhlosole; u, u-shaped fold. Scale bars = 1 mm.

ing at distal end of mid-esophagus (Figure 14, e) but

continuous into posterior esophagus. Posterior esopha-

gus narrow, bearing numerous folds of equal height.

Long, tubular salivary glands (sg) just reaching transition

to posterior esophagus.

Mipcut. Groove along midline of marginal fold lacking

(Figure 16, mf). Single, weak, caecal fold (ef) along right

side of midgut behind gastric shield, opposite caecum.

Hinpcut. Proximal intestine (Figure 15, int) passing

below distal tip of style sac (ss), then extending

posteriorly alongside style sac almost reaching main

gastric chamber (sto). Intestine curving anteriorly with

loop partially overlying proximal style sac. Intestine

extending under posterior end of main kidney chamber

(kd), entering pallial roof alongside bladder (b) and

pallial gonoduct (po), continuing forward to papillate

anus near mantle margin (Figure 14, r).

Reno-pericardial System: _ Bladder (Figure 15, b) largely

occluded by vertical sheets of excretory tissue radiating

from afferent renal vessel (Figure 17, arv) and fusing to

right lateral floor, roof and walls. Sheets of excretory tissue

loosely and regularly branching and anastomosing;

posterior sheets less branched than those anteriorly.

Pericardium rather narrow and short (Figure 15, per),

extending to intestinal loop (dotted line).

Nervous System: Buccal ganglia (Figure 14, bg) lying

dorso-laterally at base of buccal mass between buccal

retractor muscles (rt) and salivary glands (sg). Thick-

ened connective (Figure 18, co) between left pleural and

sub-esophageal ganglia producing 1-2 nerves (n=2). In

addition to zygoneury (z) and connectives to left pleural

and visceral ganglia, sub-esophageal ganglion producing

2-3 prominent nerves (n=2). Small statocysts (st) with

approximately 20-30 statoconia.

Reproductive System: Mate. Flattened longitudinal

shelf of glandular tissue within lateral lamina diminish-

ing in thickness at anterior and posterior ends of

prostate (Figures 19, 20, pr). Deep longitudinal cleft

opening in glandular shelf along gonoductal groove at

midpoint of gonoduct and extending anteriorly; cleft

closing a short distance back from anterior tip of

gonoduct (Figure 20, cl). Medial lamina thinly and

rather evenly glandular along its length; glandular

tissue slightly diminishing anteriorly (Figure 21). Epi-

thelium of medial lamina posterior third smooth

and strongly concave, separated from anterior 2/3 by

prominent curving ridge; ridge embracing correspond-

ing groove in glands of lateral lamina at proximal end of

cleft. Epithelium of medial lamina anterior 2/3 irregu-

larly and variably crossed by oblique and longitudinal

ridges.

E. E. Strong, 2005

Figure 18. Circum-esophageal nerve ring of Elimia lives-

cens. Frontal view on the left, right lateral view on the right.

Abbreviations: ce, cerebral ganglion; co, thickened connective

between left pleural and sub-esophageal ganglia; pe, pedal

ganglion; pl, pleural ganglion; sb, sub-esophageal ganglion; sp,

connective to supra-esophageal ganglion; st, statocyst; z,

zygoneury. Scale bar = 1 mm.

FEMALE. Renal oviduct (Figures 22, 23, ovi) deflected

dorsally behind mantle cavity before entering base of

glandular pallial oviduct. Proximal albumen gland rather

long and initially curved, then forming straight segment

along base of mantle cavity under pallial kidney

extension. Capsule gland comprising approximately

anterior 1/3 of pallial oviduct; externally capsule gland

markedly inflated. Above aperture, short distance back

from anterior tip of oviduct (~1/5 of length), sperm

gutter (sg) opening in medial lamina and deepening

posteriorly; gutter leading to short, blind spermatophore

bursa (spb).

DISCUSSION

Whatever may be discovered in the future regarding the

structure and relationships of species currently placed

within the Pleuroceridae, given that Plewrocera acuta is

the type species of the type genus for the family, this

description will necessarily remain as the standard for

the application of the name.

COMPARISON OF PLEUROCERA ACUTA AND ELIMIA LIVESCENS

Baker (1928) commented on the considerable uniformity

of structure in the genitalia of these two species and that

Page 12

the soft parts do not seem to show the same degree of

differentiation as the shells. Although this statement was

based on external observations, Dazo (1965) similarly

noted that, except for differences in size, the internal

anatomy of the two species is quite similar or often

identical in all organ systems; the most significant

differences were those relating to size and shape of the

operculum, snout, tentacles, foot, and radula.

Yet, notions of similarity and how similar two entities

must be to be characterized as “identical” are subjective

concepts. The thorough documentation of these two

species has been provided to allow a more objective

means of assessing the degree of similarity between the

two. Of course, any anatomical rendering will maintain

some element of subjectivity.

With this in mind, the present analysis confirms that

Pleurocera acuta and Elimia livescens are remarkably

similar, both in overall organization and in many details.

This level of similarity is perhaps not unexpected given

the sister group relationship between the two genera

supported in one molecular analysis (Holznagel and

Lydeard, 2000). However, it should be noted that

monophyly of these genera has not been demonstrated

unambiguously (e.g. Sides, 2005), but awaits confirma-

tion within a more comprehensive phylogenetic frame-

work. Thus, the present results may indicate a closer

systematic affinity than currently appreciated.

Yet, a number of differences between the two species

are apparent. Externally, the two differ in development

of the hypobranchial gland, but this is difficult to

quantify. They also differ in the position of the ovipositor

and its relationship to the foot sole. This is consistent

with described differences in the mode of egg capsule

transfer to the substrate; in Pleuwrocera acuta, only the

everted walls of the ovipositor function in oviposition

(van Cleave, 1932), but in Elimia laqueata—a species

with an ovipositor configuration identical to E. livescens

(see below)—both the everted lips of the ovipositor and

the margins of the finely grooved tract guide the ova to

the substrate (Woodard, 1934).

The most significant differences in midgut structure

are length of caecal fold, and the presence of a groove

along the marginal fold; however, these variations may

be attributable to preservation artifacts. The significance

of other minor differences (size and/or shape of caecum,

glandular pad, major typhlosole, gastric shield, crescen-

tic ridge, field of parallel striations above esophageal

aperture) can only be ascertained once a broader

sampling of species from both genera have been

examined. But these may also be attributable to

preservation artifacts and/or intra-specific variation.

Internal structure of the kidney differs only in that the

vertical sheets of excretory tissue are more highly and

densely branched within the bladder in Pleurocera

acuta. It was observed that the amount of such excretory

tissue varied between individuals in a species and with

maturity. Although a highly qualitative character, the

degree of difference between the two species surpasses

that of intra-specific variation.

Page 128

THE NAUTILUS, Vol. 119, No. 4

Z °

Ovl

Figures 19-23. Reproductive anatomy of Elimia livescens. 19. External, left lateral view of prostate. Anterior is to the left. Arrows

indicate extent of opening to gonoductal groove. 20. Internal aspect of prostate lateral lamina. Note deep cleft at anterior end (el),

representing presumptive spermatophore forming region. 21. Internal aspect of prostate medial lamina. Anterior is to the right. 22.

External, left lateral view of pallial oviduct. Anterior is to the left. Arrows indicate extent of opening to gonoductal groove. 23.

External, right lateral view of pallial oviduct. Anterior is to the right. Abbreviations: ag, albumen gland; eg, capsule gland; el, deep

cleft; et, connective tissue; gg, gonoductal groove; ovi, renal oviduct; sg, sperm gutter; spb, spermatophore bursa; vd, vas deferens.

Scale bars = 1 mm.

The configuration of the nerve ring and visceral loop is

largely identical between the two. One difference is the

number of nerves issuing from the sub-esophageal/left

pleural connective and from the sub-esophageal gangli-

on. However, given the observed intra-specific variation,

these differences likely fall within the range of individual

variation.

In contrast to the results presented here, Dazo (1965)

reported 9 cerebral nerves, only a single nerve from the

visceral ganglion, and an inconstant number of accessory

pedal nerves; however, Dazo did confirm the unique

thickened left pleural/sub-esophageal connective. Dazo

also commented on the unlikely generality of Rose-

water's (1961) findings that pleurocerids differ primarily

in the lengths of the cerebral commissure and left

pleural/sub-esophageal connective (n=6 for 9 species).

Indeed, these lengths were found to be sometimes

conspicuously different between individuals examined in

the present study.

Males of the two species differed in the pattern of

folds within the anterior region of the prostate. Regard-

less, the anterior region is inferred to be the site of

spermatophore formation rather than the comparatively

smoother posterior region. This conclusion seems

justified given the similarity in the configuration of the

folds as compared to overall form and shape of the

spermatophore (Jewell, 1931; Dazo, 1965). Limnic

cerithioideans in the family Paludomidae Stoliczka,

1868, have separated the glands in this anterior region

to form a hollow tube that has been implicated in

spermatophore formation (Glaubrecht and Strong,

2004). That discovery further supports the notion

that sperm packets are produced anteriorly and suggests

that this function may be homologous in different

lineages.

For a summary of these and other morphological

differences, see Table 1.

COMPARISON TO OTHER PLEUROCERIDS

With minor exception, published accounts agree on the

main patterns of pleurocerid anatomy. Thus, pleurocer-

ids have long been known to be dioecious and oviparous

(except Semisulcospira) with an ovipositor involved in

the deposition of the egg capsules (Stimpson, 1864).

Members of the family are also aphallate, with open

gonoducts, a gonad that dorsally surrounds the digestive

gland, and produce crescent-shaped spermatophores

(e.g. Stimpson, 1864; Jewell, 1931; Woodard, 1934, 1935;

Magruder, 1935b; Jones and Branson, 1964; Dazo,

1965). Like other cerithioideans, pleurocerids possess

E. E. Strong, 2005

Page 129

Table 1. Summary of morphological differences between Pleurocera acuta and Elimia livescens.

Pleurocera acuta

External Anatomy:

Propodium

Ovipositor ventral groove

Curved anterior tip of osphradium

Narrow

Alimentary System:

Ventral folds at posterior end of mid- Fused

esophagus

Groove along marginal fold Present

Length of caecal fold Long

Hindgut loop

Reno-Pericardial System:

Bladder excretory tubules

Pericardium

Nervous System:

Buccal ganglia Ventro-lateral

Statoconia ~10-15

Nerves from sub-esophageal/left pleural 1-3

connective

Nerves from sub-esophageal ganglion

zygoneury)

Reproductive System:

Spermatophore-forming region

Sperm gutter

Junction of renal and pallial oviduct

Proximal albumen gland

two types of glands within the pallial oviduct — a feature

not previously documented among eastern North

American species.

The gut is characterized by the presence of tubular

salivary glands, a crystalline style and a style sac in

restricted communication with the proximal intestine

(Magruder, 1935a, b; Itagaki, 1960; Dazo, 1965). The bi-

lobed nature of the kidney has been noted before

(Magruder, 1935b; Itagaki, 1960), but the internal sub-

division of the organ had not been previously documented.

The nervous system is consistent with many other

cerithioideans (left dialyneurous, long connective between

the right pleural and supra-esophageal ganglia, single

visceral ganglion) (e.g. Strong, 2003; Strong and Glau-

brecht, 2002, 2003), but is distinguished by the presence

of an enlarged connective between the left pleural and

sub-esophageal ganglia and a zygoneurous connection on

the right—the latter a highly homoplastic character in the

Cerithioidea (see review in Houbrick, 1988). However, the

presence/absence of zygoneury among pleurocerids re-

quires confirmation as it has been depicted as dialyneury

in several other species (Magruder, 1935b; Itagaki, 1960).

Only the former study confirmed the presence of the

enlarged left pleural and sub-esophageal connective.

Additionally, published descriptions (Magruder, 1935b;

Does not extend to foot margin

Sometimes present

Extends to main gastric chamber

Densely and highly branched

Extends to right body wall

2 connectives and 2 nerves (including

Renal oviduct curves dorsally to straight

segment of albumen gland

Short, straight segment between renal

oviduct and posterior end of bursa

Elimia livescens

Broad

Extends to foot margin

Often present

Unfused

Absent

Short

Does not reach main gastric chamber

Loosely and regularly branched

Extends to intestinal loop

Dorsal

~20-30

12:

2 connectives and 3—4 nerves

(including zygoneury)

Parallel folds and trough in medial lamina Deep cleft in lateral lamina

Extends to anterior tip of oviduct

Does not extend to anterior tip of

pallial oviduct

Renal oviduct ventrally joins curved

portion of albumen gland

Initially curved, with long, straight

segment to posterior end of

spermatophore bursa

Itagaki, 1960; Dazo, 1965; present study) disagree on the

number of nerves produced by various ganglia, but as

noted above, this can be highly variable even within

species. However, the number of statoconia reported by

Magruder (30-40; 1935b) is significantly more than the

number reported herein, possibly exceeding the level of

intra-specific variation and, thus may be an informative

phylogenetic character.

Several significant discrepancies among previous

descriptions of pleurocerid anatomy are now resolved.

Woodard (1934, 1935) observed a so-called “cytophore

organ” at the base of the mantle cavity, apparently

confluent with the sperm duct. Similarly, Dazo (1965)

reported the presence of a cytophore organ in males of

both Pleurocera acuta and Elimia livescens. Based on the

present analysis and Woodard’s description of the

internal structure and position of this organ, it is clear

that Woodard misidentified the kidney bladder as a part

of the reproductive tract. However, it is not clear why

the cytophore organ was described as lacking in females.

The intimate connection between the bladder and

proximal pallial gonoduct was correctly depicted in

Hua by Prozorova (1990).

Dazo (1965), as well as several other workers

(Woodard, 1934; Jones and Branson, 1964), have

Page 130

Table 2.

THE NAUTILUS, Vol. 119, No. 4

Summary of morphological differences between Pleurocera acuta and Elimia livescens compared to other limnic

gastropods classified in the Pleuroceridae and Melanopsidae. Details from Itagaki, 1960; Bilgin, 1973; Houbrick, 1988; Nakano and

Nishiwaki, 1989; Glaubrecht, 1996; Strong and Glaubrecht, unpubl. data.

Pleuraceura acuta

Elimia livescens

External Anatomy:

Ovipositor pore Simple, weakly glandular

Juga

Simple, weakly

Semisulcospira Melanopsidae

Not applicable Complex, highly glandular

glandular

Alimentary System:

Salivary glands Tubular ? Tubular Tubular/branched

Salivary gland position Pass through nerve ring ? Pass through nerve _- Pass through/anterior to

ring nerve ring

Esophageal gland Absent ? Absent? Present

Digestive gland ducts 2 >? 2 il

Caecum Small >? Small Deep and spiral

Reno-Pericardial System:

Bladder Small, pallial P i Small, pallial

Nervous System:

Dialyneury, Zygoneury Zygoneury ie Dialyneury? Zygoneury

Reproductive System:

Seminal vesicle Straight ? Straight Coiled

Pallial oviduct Open Open Closed Open

Seminal receptacle Absent Present Present Present

Reproductive strategy Oviparous Oviparous Viviparous Oviparous

reported the presence of a seminal receptacle, but no

mention of a spermatophore bursa was made. The

present study has confirmed that the sperm storage

structure in Plewrocera acuta and Elimia livescens is

a bursa based on the presence of unorientated sperm.

Examination of specimens of E. laqueata (USNM

1081558) confirmed that the structure reported as

a seminal receptacle is indeed a bursa—no seminal

receptacle is present; judging from the description of

Jones and Branson (1964), the same holds true for E.

potosiensis. The pallial position of the bladder was also

confirmed in E. laqueata (pers. obs.). It is interesting to

note that the sperm gutter extends farther anteriorly in

E. laqueata than in E. livescens. Additionally, the straight

segment of the albumen gland between the tip of the

bursa and the junction of the renal oviduct is lacking in

E. laqueata. Instead, the albumen gland arcs dorsally to

join the renal oviduct just behind the tip of the bursa. In

all other respects, the overall structure of the pallial

oviduct in E. laqueata is consistent with features

described here, as well as in the close association

between the ovipositor and the junction of the

propodium and mesopodium.

A final discrepancy is the presumed site of spermato-

phore formation. Woodard (1934, 1935) described the

distal prostate as smooth and the highly folded proximal

portion as the site of spermatophore formation. Jones

and Branson (1964) did not distinguish a spermatophore-

forming region in Elimia potosiensis. In the present

study, it is the highly folded distal region that is inferred

to be the site of spermatophore formation.

SYSTEMATICS OF PLEUROCERIDAE

As mentioned above, although confusion has long

existed, the distinctiveness of the Pachychilidae from

other limnic lineages including the Pleuroceridae has

now been clarified based on morphological and molec-

ular data (e.g. Glaubrecht, 1996, 1999; Lydeard et al.,

2002; Kohler et al., 2004). However, the paraphyly of

eastern and western North American and Asian pleur-

ocerids with respect to the Melanopsidae based on

molecular data (Lydeard et al., 2002) remains at issue.

The analysis of Houbrick (1988) did not include

sufficient taxon sampling to adequately assess mono-

phyly of the two families, but a sister-group relationship

between the two was supported.

Although an in depth analysis of monophyly and

affinity of the two families is beyond the scope of this

study, several morphological features may be informa-

tive in clarifying these relationships. As noted by

Prozorova (1990) the reproductive anatomy of eastern

North American pleurocerids differs from species in

western North America (Juga) and Asia (Semisulcospira)

in that both Juga and Semisulcospira possess a seminal

receptacle in addition to a spermatophore bursa. The

latter genus has modified the pallial oviduct into a closed

brood pouch (Itagaki, 1960; Nakano and Nishiwaki,

1989; Prozorova, 1990; Rashchepkina, 2000; Prozorova

and Raschepkina, 2001, 2004).

In addition to the synapomorphies recovered in the

analysis of Houbrick (weakly developed hypobranchial

gland, zygoneury, long left pleural/sub-esophageal con-

E. E. Strong, 2005

Page 131

nective), midgut anatomy is broadly congruent in the

two families, differing in several significant respects from

that of the other limnic lineages (e.g. Paludomidae,

Pachychilidae, Thiaridae) (Bilgin, 1973; KGhler and

Glaubrecht, 2001; Strong and Glaubrecht, 2002, 2003,

unpubl. data). They also share similarities in reno-

pericardial (presence of a bladder) and reproductive

anatomy (open pallial gonoducts, presence of a seminal

receptacle)—the latter, in particular, are undoubtedly

symplesiomorphic. But other aspects of the anatomy are

consistent within each family and clearly differentiate

the two when sufficient information is available. Thus, in

addition to features of the radula and shell, melanopsids

may be distinguished by the presence of an esophageal

gland, salivary glands that lie anterior to the nerve ring

(although variable in the family), a single digestive gland

duct and spiral caecum in the midgut, and a coiled

seminal vesicle. Of course, the extent to which these

features represent shared derived or homoplastic

features remains to be discovered in the context of

a phylogenetic analysis.

CONCLUSIONS

The present study has provided the first detailed

description of the midgut and kidney for any pleurocerid

snail, and has clarified the internal structure and

homologies of the pallial gonoducts of eastern North

American forms. This comparative analysis has confirmed

the high degree of morphological similarity between

Pleurocera acuta and Elimia livescens, but has also

revealed a number of differences in detail; the extent to

which these features support monophyly of the genera

remains to be established. The fact that the presence/

absence of a seminal receptacle distinguishes eastern and

western North American/Asian pleurocerids is confirmed.

Pleurocerids and melanopsids are broadly similar in

features of the midgut and share a similar configuration of

the pallial oviduct, but can be distinguished by anatomical

characters of the alimentary (salivary glands, esophageal

gland, digestive gland ducts, caecum) and reproductive

(seminal vesicle) systems. However, comprehensive

anatomical treatments of western North American and

Asian pleurocerids are needed to fully assess the

morphological distinctiveness of the two families. The

clarification of the distribution of these features, within

the context of a phylogenetic analysis, should aid in

refining the monophyly of the Pleuroceridae and their

affinity to other freshwater lineages.

ACKNOWLEDGMENTS

I thank Charles Lydeard for organizing a gastropod

morphology workshop at University of Alabama, Tusca-

loosa, in September 2003 that inspired this study. I also

thank Rex Hanger (University of Wisconsin-Whitewater)

for assistance with collecting the samples of Plewrocera

acuta and Elimia livescens, Paul Johnson (Alabama

Department of Conservation and Natural Resources) for

supplying comparative material of Elimia laqueata, and

Jonathan Slaght (University of Minnesota) for providing

translations of several Russian texts. I am indebted to

Marilyn Schotte (USNM) for inking the anatomical

drawings. Arthur Bogan (North Carolina State Museum

of Natural Sciences), Philippe Bouchet (Muséum

national d'Histoire naturelle, Paris) and John Wise

(College of Charleston) provided valuable comments

that improved the quality of the manuscript.

LITERATURE CITED

Baker, F. C. 1928. The fresh water Mollusca of Wisconsin. Part

I. Gastropoda. Bulletin of the Wisconsin Geological and

Natural History Survey, Wisconsin, 70: 1-507, i-xx, 28 pls.

Bilgin, F. H. 1973. Studies on the functional anatomy of

Melanopsis praemorsa (L.) and Zemelanopsis trifasciata

(Gray). Proceedings of the Malacological Society, London,

40; 379-393.

Bouchet, P., and J.-P. Rocroi (eds.) (2005). Classification and

nomenclator of gastropod families. With classification by

J. Fryda, B. Hausdorf, W. Ponder, A. Valdés and A.

Warén. Malacologia 47: 1-397.

Dazo, B. C. 1965. The morphology and natural history of

Pleurocera acuta and Goniobasis livescens (Gastropoda:

Cerithiacea: Pleuroceridae). Malacologia 3: 1-80.

Dillon, R. T. 2000. The ecology of freshwater mollusks

Cambridge University Press, Cambridge.

Glaubrecht, M. 1996. Evolutionsékologie und Systematik

am Beispiel von Si®- und Brackwasserschnecken

(Mollusca: Caenogastropoda: Cerithioidea): Ontogenese-

Strategien, palaontologische Befunde und Historische

Zoogeographie. Backhuys Publishers, Leiden, 499 pp.,

25 pls.

Glaubrecht, M. 1999. Systematics and the evolution of

viviparity in tropical freshwater gastropods (Cerithioidea:

Thiaridae sensu lato) - an overview. Courier Forschungs-

Institut Senckenberg 215: 91-96.

Glaubrecht, M. and T. v. Rintelen. 2003. Systematics,

molecular genetics and historical zoogeography of the

viviparous freshwater gastropod Pseudopotamis (Cer-

ithioidea, Pachychilidae): a relic on the Torres Strait

Islands, Australia. Zoologica Scripta 32: 415-435.

Glaubrecht, M. and E. E. Strong. 1999. Midgut Morphology

and Implications for Cerithioidean Phylogeny (Mollusca:

Gastropoda). Abstracts. XVIII Meeting of the Willi

Hennig Society, Géttingen, 1999, p. 24.

Glaubrecht, M. and E. E. Strong. 2004. Spermatophores of

thalassoid gastropods (Paludomidae) in Lake Tanganyika,

East Africa, with a survey of their occurrence in

Cerithioidea: functional and phylogenetic implications.

Invertebrate Biology 123: 218-236.

Goodrich, C. 1945. Goniobasis livescens of Michigan. Mis-

cellaneous Publications, Museum of Zoology, University

of Michigan 64: 5-36.

Holznagel, W. E. and C. Lydeard. 2000. A molecular

phylogeny of North American Pleuroceridae (Gastropoda:

Cerithioidea) based on mitochondrial 16S rDNA se-

quences. Journal of Molluscan Studies 66: 233-257.

Houbrick, R. S. 1988. Cerithioidean phylogeny. In: W. F.

Ponder (ed.) Prosobranch Phylogeny. Malacological

Review, Supplement 4: 88-128.

Page 132

Howe, S. W. 1938. A study of the radulae of snails of the

family Pleuroceridae. American Midland Naturalist 20:

549-561.

Itagaki, H. 1960. Anatomy of Semisulcospira bensoni, a fresh-

~ water gastropod. Venus 21: 41-51.

Jewell, D. D. 1931. Observations on reproduction in the snail

Goniobasis. The Nautilus 44: 115-119.

Johnson, P. D., A. E. Bogan, C. E. Lydeard, K. M. Brown and

J. E. Cordeiro. 2005. Development of an initial conser-

vation assessment for North American freshwater gastro-

pods. Freshwater Mollusk Conservation Society, 4th

Biennial Symposium. Meeting Program and Abstracts,

p. 30.

Jones, W. C. and B. A. Branson. 1964. The radula, genital

system, and external morphology in Mudalia potosiensis

(Lea) 1841 (Gastropoda: Prosobranchiata: Pleuroceridae)

with life history notes. Transactions of the American

Microscopic Society 83: 41-62.

Kohler, F. and M. Glaubrecht. 2001. Toward a systematic

revision of the Southeast Asian freshwater paeched

Brotia H. Adams, 1866 (Cerithioidea: Pachychilidae): a:

account of species from around the South China Ses

Journal of Molluscan Studies 67: 283-321.

Kohler, F., T. v. Rintelen, A. Meyer and M. Glaubrecht. 2004.

Multiple origin of viviparity in Southeast Asian gastropods

(Cerithioidea: Pachychilidae) and its evolutionary implica-

tions. Evolution 58: 2215-2226.

Lydeard, C., W. E. Holznagel, M. Glaubrecht and W. F.

Ponder. 2002. Molecular phylogeny and evidence for

multiple origins of freshwater gastropods of the circum-

global, diverse superfamily Cerithioidea (Mollusca: Cae-

nogastropoda). Molecular Phylogenetics and Evolution

22: 399-406.

Magruder, S. R. 1935a. Record of a crystalline style in two

fresh water gastropods. The Nautilus 48: 101-102.

Magruder, S. R. 1935b. The anatomy of the fresh water

prosobranchiate gastropod, Plewrocera canaliculatum

undulatum (Say). American Midland Naturalist 16:

883-912.

Mihalcik, E. L. and F. G. Thompson. 2002. A taxonomic

revision of the freshwater snails referred to as Elimia

curvicostata, and related species. Walkerana 13: 1-108.

Minton, R. L. and C. Lydeard. 2003. Phylogeny, taxonomy,

genetics and global heritage ranks of an imperiled,

freshwater snail genus Lithasia (Pleuroceridae). Molecu-

lar Ecology 12: 75-87.

Morrison, J. P. E. 1954. The relationships of Old and New

World Melanians. Proceedings of the United States

National Museum 103: 357-394.

Nakano, D. and S. Nishiwaki. 1989. Anatomical and histolog-

ical studies on the reproductive system of Semisulcospira

libertina (Prosobranchia: Pleuroceridae). Venus 48:

263-273.

Ponder, W. F. and A. Warén. 1988. Classification of the

Caenogastropoda and Heterostropha — a list of the family

group names and higher taxa. In: W. F. Ponder (ed.)

Prosobranch Phylogeny. Malacological Review, Supple-

ment 4: 288-326.

Prozorova, L. A. 1990. On the biology reproduction of molluscs

Pachychilidae (Gastropoda, Cerithiiformes). Zoologiches-

kii Zhurnal 69: 24-37.

THE NAUTILUS, Vol. 119, No. 4

Prozorova, L. A. and A. V. Raschepkina. 2001. Comparative

anatomy of reproductive system of the Juga-like gastro-

pods (Gastropoda, Cerithioidea) from South Korea and

Primorye Territory. The Bulletin of the Russian Far East

Malacological Society 5: 62-70. [In Russian. ]

Prozorova, L. A. and A. V. Raschepkina. 2004. Reproductive

anatomy of some genera of North American Pleuroceridae

(Gastropoda: Cerithiiformes: Cerithioidea). The Bulletin

of the Russian Far East Malacological Society 8: 87-94.

[In Russian. ]

Rashchepkina, A. V. 2000. Anatomy of the pallial oviduct of the

genus Hua Chen (Pachychilidae, Cerithioidea). The

Bulletin of the Russian Far East Malacological Society 4:

99-100. [In Russian. ]

Rosewater, J. 1961. Preliminary observations on the nervous

systems of some Pleuroceridae. Program and Abstracts.

American Malacological Union, 28th Annual Meeting,

June 19-23.

Sides, J. D. 2005. The systematics of freshwater snails of the

genus Pleurocera (Gastropoda: Pleuroceridae) from the

Mobile River basin. Unpublished Ph.D thesis, University

of Alabama — Tuscaloosa, 222 pp.

Stimpson, W. 1864. On the structural characters of the so-

called Melanians of North America. The American Journal

of Science and Arts, Second Series 38: 41-53.

Strong, E. E. 2003. Refining molluscan characters: morphol-

ogy, character coding and the phylogeny of the Caenogas-

tropoda (Gastropoda). Zoological Journal of the Linnean

Society 137: 447-554.

Strong, E. E. and M. Glaubrecht. 1999. Tapping the un-

explored: midgut morphology of cerithioidean gastropods

(Caenogastropoda)—preliminary results and implications

for homology and phylogeny. Program and Abstracts.

American Malacological Society, 65th Annual Meeting,

Pittsburgh, PA, July 4-9, 1999, pp 53-54.

Strong, E. E. and M. Glaubrecht. 2002. Evidence for

convergent evolution of brooding in a unique gastropod

from Lake Tanganyika: anatomy and affinity of Tanganyj-

cia rufofilosa (Smith, 1880) (Caenogastropoda, Cerithioi-

dea, Paludomidae). Zoologica Scripta 31: 167-184.

Strong, E. E. and M. Glaubrecht. 2003. Anatomy and

systematic affinity of Stanleya neritinoides (Smith, 1880),

an enigmatic member of the thalassoid gastropod fauna

from Lake Tanganyika, East Africa (Cerithioidea, Paludo-

midae). Acta Zoologica 84: 249-265.

Strong, E. E., M. Glaubrecht, C. Lydeard and W. F. Ponder.

9002. A total evidence phylogeny of the Cerithioidea. In:

R. T. Dillon (ed.) Program and Abstracts, 6Sth American

Malacological Society. American Malacological Society,

Inc., Charleston, p. 102.

van Cleave, H. J. 1932. Studies on snails of the genus

Pleurocera. 1. The eggs and egg laying habits. The

Nautilus 46: 29-34.

Woodard, T. M. 1934. Anatomy of the reproductive system of

Goniobasis laqueata (Say). Journal of the Tennessee

Academy of Science 9: 243-259.

Woodard, T. M. 1935. Spermic dimorphism in Goniobasis

laqueata (Say). Journal of Morphology 57: 1-23.

Woodard, T. M. 1940. The function of the apyrene spermatozoa

of Goniobasis laqueata. 1. The behavior of the apyrene and

eupyrene spermatozoa under natural and artificial condi-

tions. The Journal of Experimental Zoology 85: 103-123.

THE NAUTILUS 119(4):133-148, 2005

New Late Cretaceous (Santonian and Campanian) gastropods

from California and Baja California, Mexico

Richard L. Squires

Department of Geological Sciences

Califormia State University

Northridge, CA 91330-8266 USA

[email protected]

County

LouElla R. Saul

Invertebrate Paleontology Section

Natural History Museum of Los Angeles

900 Exposition Boulevard

Los Angeles, CA 90007 USA

[email protected]

ABSTRACT

Three new genera and six new species of shallow-marine Late

Cretaceous gastropods are reported from various formations in

California and from one formation in Baja Califormia, Mexico.

Tegula jeanae new species, of early Campanian age, is the

earliest known species of this trochid genus. Nerita (sub-

genus?) orovillensis new species is the second known Early

Campanian neritid from California. The cerithioid Bullamir-

ifica new genus is represented by three species: Bullamirifica

verruca new species of Coniacian age; Bullamirifica elegans

new species of early Campanian age; and Bullamirifica ainiktos

(Dailey and Popenoe, 1966) of middle to late Campanian age.

The latter species has the most widespread distribution, with

occurrences in southern California and northern Baja Cali-

fornia. Minytropis melilota new genus and species of Santonian

age, and Paxitropis dicriota new genus and species of Late

Santonian to early Campanian age are high-spired trichotro-

pids. As presently known, Bullamirifica, Minytropis, and

Paxitropis were endemic to the study area.

INTRODUCTION

This study is based largely on specimens collected by

Eric Goéhre of Oroville, California. Over the years, he

has amassed a sizeable collection of shallow-marine

mollusks from the lower Campanian Pentz Road

member of the Chico Formation near Pentz, Butte

County, northern California (Figure 1). His collection

has yielded several new species of gastropods, and some

of these were described by Groves (2004) and Squires

and Saul (2004). In part, this present study concerns

three additional new species and a new genus of

gastropods found in his collection. They are the trochid

Tegula jeanae new species, the neritid Nerita (sub-

genus?) orovillensis new species, and the cerithioid

Bullamirifica elegans new genus and species.

Inspection of the literature, as well as examination of

the collections at the Natural History Museum of Los

Angeles County, allowed us to incorporate two additional

species into Bullamirifica. These are Bullamirifica

verruca new genus and species from the Coniacian

Member IV of the Redding Formation in the Oak Run

area, northem Califomia, and Bullamirifica anikitos

Dailey and Popenoe (1966) new combination from the

middle Campanian Pigeon Point Formation southwest of

San Francisco, northern California; the middle upper

Campanian Punta Baja Formation, Baja Califomia,

Mexico; and the upper Campanian Jalama Formation,

southern California (Figure 1). “Cimolithium miya-

koense” (Nagao, 1934) and “Vicarya (Shoshiroia) yabei”

Kamada, 1960, reported by Perrilliat-Montoya (1968)

from Baja Califormia, Mexico (see Figure 1, formation 6),

are judged by us to be synonyms of Bullamirifica ainiktos.

Also included in this present study are new tricho-

tropid gastropods found in the collections at the Natural

History Museum of Los Angeles County. They are

Minytropis melilota new genus and species from the

Santonian part of the Redding and Chico formations of

northern California, and Paxitropis dicriota, new genus

and species from the of upper Santonian part of the

Redding Formation, northern California; the lower

Campanian part of the Chico Formation; and the lower

Campanian part of the upper Holz Member of the Ladd

Formation, southern California (Figure 1).

The geologic age of each new species described in this

paper is shown in Figure 2. The entire interval of time

that encompasses all “anes species is Coniacian to late

Campanian, or about 19 million years. The new species

are locally common, except for Tegula jeanae, Nerita

(subgenus?) orovillensis, and B. verruca.

The classification system used here generally follows

that of Hickman and McLean (1990) for the tegulines,

Ponder (1988) for the trichotropids, and Ponder and

Warén (1988) for the other taxa.

Study localities are listed in Appendix 1. Abbreviations

used in the text are: CAS: California Academy of Sciences,

Page 134

1-Redding Formation

2-Chico Formation

3-Pigeon Point

Formation

4-Jalama Formation

5-Ladd Formation

6-Punta Baja

Formation

Figure 1. Location of formations bearing the new taxa.

San Francisco; IGM: México Museo del Paleontologia del

Instituto de Geologa; LACMIP: Natural History Museum

of Los Angeles County, Invertebrate Paleontology

Section; UCLA: University of California, Los Angeles

(collections now housed at LACMIP); UCMP: University

of California Museum of Paleontology (Berkeley); USGS:

United States Geological Survey.

STRATIGRAPHY

Except for the Punta Baja Formation, which is discussed

below, the ages and depositional environments of all the

formations and members containing the new taxa

discussed in this paper can be found in the following

papers: Member IV of the Redding Formation, Squires

and Saul (2003a); Musty Buck Member of the Chico

Formation, Saul and Squires (2003); Pentz Road

member (informal) of the Chico Formation, Squires

THE NAUTILUS, Vol. 119, No. 4

AGE (m.y.) 80

LL}

UPPER CRETACEOUS

Campanian

Coniacian

°o

polarity

chrons

Tegula Fee:

orovillensis

Nerita

verruca

Bullamirifica abs

! ainiktos

Minytropis

Figure 2. Chronostratigraphic positions of the new taxa.

Ages of stage boundaries and magnetostratigraphy from

Gradstein et al. (2004, fig. 19.1).

and Saul (1997); Ten Mile Member of the Chico

Formation, Squires and Saul (2003b); upper Holz Shale

of the Ladd Formation, Squires and Saul (2001); Pigeon

Point Formation, Elder and Saul (1993) and Squires and

Saul (2003b); and Jalama Formation, Squires and Saul

(2003b). The locales of these formations are shown in

Figure 1. Stratigraphic information mentioned below

concerns additional pertinent biostratigraphic details.

The age of the Jalama Formation used here is slightly

younger than used in our previous papers because we

had to adjust its chronostratigraphic position based on

the latest published (Gradstein et al., 2004) absolute-

time and global-paleomagnetic data correlations.

Punta Baja FORMATION

Perrilliat- Montoya (1968) reported specimens of gastro-

pods, herein assigned to Bullamirifica ainiktos, from the

“Rosario Formation” at Punta Baja, near El Rosario,

northern Baja California, Mexico. The 5-140 m thick

Punta Baja Formation (Figure 1) overlies fluvial deposits

of the La Bocana Roja Formation, and the angular

unconformity between these two formations is canyon-

shaped (Boehlke and Abbott, 1986). This canyon is filled

with conglomerate, sandstone, and siltstone reported by

Kilmer (1963) to have been deposited in shallow-marine

depths not exceeding 60 meters. Boehlke and Abbott

(1986) have a differing viewpoint and reported that the

deposits represent turbidites that accumulated in bathyal

depths. They reported, furthermore, that shallow-marine

mollusks are common, but Kilmer’s collection at UCMP

does not contain very many specimens. The Punta Baja

Formation is unconformably overlain by terrestrial

deposits of the La Escarpa Member of the El Gallo

R. L. Squires and L. R. Saul, 2005

Page 135

Formation, which, in turn, is overlain by the Rosario

Formation.

Based on molluscean fossils collected by F. H. Kilmer,

Saul (1983: 21-22, fig. 9) reported the ammonite

Metaplacenticeras cf. pacificum (Smith, 1900) and the

gastropod Turritella chicoensis pescaderoensis Arnold,

1908, from the siltstone in the Punta Baja Formation.

Although these two mollusks were reported by Saul

(1983: 65-66) to be of late Campanian age, more recent

biostratigraphic studies (Elder and Saul, 1996: fig. 1)

depicted both of these taxa as ranging in age from late

middle Campanian to earliest late Campanian. Adjust-

ments for the most recently published (Gradstein et al.,

2004) absolute-time and global-paleomagnetic data

correlations place these ammonite and turritellid zones

in the middle late Campanian. Recent examination by

the junior author of additional Punta Baja Formation

mollusks revealed three specimens of the bivalve Calva.

The best preserved specimen is from LACMIP loc.

12582 and is Calva (Egelicalva) crassa Saul and

Popenoe, 1992, whose geologic range is early late

Campanian to early Maastrichtian elsewhere on the

Pacific slope of North America (Saul and Popenoe,

1992). The other two Calva specimens are worn and

broken, from UCMP loc. B-3388. These two specimens

are similar to Calva (Calva) peninsularis (Anderson and

Hanna, 1935), whose geologic range is latest Campanian

to early Maastrichtian elsewhere on the Pacific slope of

North America (Saul and Popenoe, 1992).

Boehlke and Abbott (1986: fig. 4) assigned the age of

the Punta Baja Formation to the early Campanian based

entirely on calcareous nannofossils. They also reported

that the benthic foraminifera in this formation corre-

spond to the F2-lower E foraminifera zones of Goudkoff

(1945), but they did not rely on the foraminifera for their

age call. Almgren (1986: table 2) reported that the F2-

lower E foraminifera zones are essentially correlative to

the early Campanian to late Campanian. It is important

to mention that the Alcalde Shale in the Coalinga area

along the west side of the San Joaquin Valley, central

California, is correlative to the E zone (Almgren, 1986:

table 3). As depicted in Saul (1983: fig. 10), the Alcalde

Shale contains Metaplacenticeras cf. M. pacificum, and

Almgren (1986) assigned the Alcalde Shale to the early

late Campanian.

In summary, the Metaplacenticeras, Turritella, Calva,

and benthic foraminifera data strongly support a middle

late Campanian age for the Punta Baja Formation. The

calcareous nannofossils, however, support an early

Campanian age. It seems probable that the older

calcareous nannofossils are reworked, and this would

be consistent with the depositional environment of the

formation.

SYSTEMATIC PALEONTOLOGY

Superfamily Trochoidea Rafinesque, 1815

Family Trochidae Rafinesque, 1815

Subfamily Tegulinae Kuroda, Habe and Oyama, 1971

Genus Tegula Lesson, 1835

Type Species: Tegula elegans Lesson, 1835, by

monotypy; Recent, west coast of Central America to

the Gulf of California, Mexico.

Discussion: Although Wenz (1938), Keen (1960), and

Davies (1971) reported the geologic range of Tegula to

be Miocene to Recent, Bandel and Stinnesbeck (2000)

reported a species of Tegula of Late Cretaceous

(Maastrichtian) age from central Chile. Kiel and Bandel

(2001) reported a tentatively identified Tegula from

upper Campanian strata in northern Spain. The early

Campanian new species described below represents the

confirmed earliest record we know of for Tegula. For the

Pacific slope of North America, the previous earliest

record of Tegula was given by Addicott (1973: 17, pl. 8,

figs. 2, 4), who reported it from the Wygal Sandstone

Member of the Temblor Formation, southwestern

margin of the San Joaquin Valley, Kern County, central

California. Squires (2003: table 2.1, fig. 2.1) placed this

member in the lower Oligocene Matlockian Stage.

Tegula jeanae new species

(Figures 3-5)

Diagnosis: A Tegula with low to moderate spire.

Whorls convex, smooth, and bearing one spiral groove

on posterior third of last whorl. Anomphalous. Last

whorl with raised lip along basal edge, base sunken

between this lip and columellar lip, which bears at least

one denticle and one much smaller denticle adapically.

Description: Shell medium (up to 16.6 mm height

and 21.7 mm diameter, same specimen). Turbiniform

with spire low to moderately elevated. Protoconch

unknown. Teleoconch consisting of three whorls. Suture

impressed, slightly channeled. Whorls convex, some-

times slightly concave short distance anterior of suture;

blunt angulation anterior of medial part of last whorl.

Whorls smooth; spiral groove posterior of medial part

of last whorl. Aperture oblique, peristome discontinu-

ous. Anomphalous, umbilical area covered by broad

callus. Base wide and smooth, peripheral (abaxial)

margin coincident with raised lip along edge of last

whorl; area depressed between this lip and columella.

Outer lip strongly prosocline. Columellar lip with at least

one oblique denticle; much weaker second denticle

sometimes present immediately posterior of main

denticle. Growth lines strongly prosocline, forming wide

bands.

Holotype: LACMIP 13322,

22 mm in diameter.

Paratype: LACMIP 13323.

Type Locality: LACMIP loc. 24337.

18.7mm _ in _ height,

Geologic Age: Early Campanian.

Page 136

THE NAUTILUS, Vol. 119, No. 4

Figures 3-8. New tegulid and neritid gastropods. Specimens coated with ammonium chloride. 3-5. Tegula jeanae new species,

LACMIP loc. 24337. 3. Paratype LACMIP 13323, apertural view, height 13.9 mm, diameter 17.9 mm. 4-5. Holotype 13322, height

18.7 mm, diameter 22 mm. 6-7. Nerita (subgenus?) orovillensis new species. 6. Holotype LACMIP 13324, apertural view, height

11.6 mm, diameter 16.8 mm. 7-8. Paratype LACMIP 13325, total shell height 10 mm, diameter 13.7 mm. 7. Abapertural view. 8.

Apical view.

Distribution: Chico Formation, Pentz Road member

(informal), near Pentz, Butte County, northern Califor-

nia.

Etymology: Named for Jean Géhre, mother of Eric

Géhre, who collected and donated the type material to

LACMIP.

Discussion: This new species is known from two

specimens, both showing good preservation. The new

species is remarkably similar to Tegula (Chlorostoma)

funebralis (Adams, 1855), from Pliocene and Pleistocene

strata of southern California (Grant and Gale, 1931) and

from the Recent of Vancouver Island, British Columbia,

to central Baja California, Mexico (McLean, 1978). The

new species differs from T. (C.) funebralis by having

a smooth shell rather than being ornamented by weak

spiral ribs. The similarity between the new species and

T. (C.) funebralis is even stronger if the specimens of the

latter are worn.

Tegula ovallei (Philippi, 1887: pl. 12, fig. 4; Bandel

and Stinnesbeck, 2000: 761, pl. 1, B), the only other

positively identified Cretaceous Tegula that we know of,

is from Maastrichtian strata in central Chile. The new

species differs from T. ovallei by having a smooth shell

rather than being ornamented by granulated spiral

ridges.

Kiel and Bandel (2001: 139, pl. 1, fig. 1) reported

a tentatively identified Tegula? simplex (Quintero and

Revilla, 1966: 49, pl. 8, fig. 3) from upper Campanian

strata in northern Spain. The new species differs greatly

from T.? simplex by having a less elevated spire, smooth

shell, blunt rather than a sharp angulation anterior of the

medial part of the last whorl, broad callus covering the

umbilical region, wider aperture, very much stronger

denticles on the columella, raised lip along the basal

edge of the last whorl, and sunken base between this

raised lip and the columellar lip.

Family Neritidae Rafinesque, 1815

Genus Nerita Linnaeus, 1758

Type Species: Nerita peloronta Linnaeus, 1758, by

subsequent designation (Montfort, 1810); Recent, south

Florida, West Indies, and Bermuda.

Discussion: Nerita sensu lato ranges from Early

Cretaceous (Hauterivian), and the earliest record is

from the Ono Member of the Budden Canyon

Formation, Trinity County, northern California (Saul

and Squires, 1997). The new species described below

represents the first record of an early Campanian Nerita

from the study area.

Subgenus?

Nerita (subgenus?) orovillensis new species

(Figures 6-8)

Diagnosis: A Nerita with approximately 18 to 19

beaded spiral ribs. Columellar lip with four or five

obscure teeth.

R. L. Squires and L. R. Saul, 2005

Description: Shell medium small (up to 11.6 mm in

height and 16.3 mm in diameter, same specimen),

broader than high, globose. Last whorl rapidly expand-

ing. Protoconch unknown. Teleoconch consisting of 2.5

to 2.75 whorls. Uppermost spire very low. Suture

obscure. Earliest 1.5 teleoconch whorls apparently

smooth, rest of teleoconch covered with approximately

18 narrow spiral ribs bearing small beads; interspaces

between ribs approximately as wide as interspaces.

Beads on ribs becoming smaller and slightly elongate

on base of last whorl, especially in parietal region. Spiral

rib adjacent to suture can be slightly stronger than other

ribs. Aperture large, nearly circular. Outer lip flared,

interior smooth. Columellar lip with five somewhat

obscure teeth, most posterior tooth widest and longest.

Deck area broad, sloping, and sharply demarcated from

base of last whorl. Growth lines prosocline.

Holotype: LACMIP 13324, 11.6mm in height,

16.8 mm in diameter.

Paratype: LACMIP 13325.

Type Locality: LACMIP loc. 24337.

Geologic Age: Early Campanian.

Distribution: Chico Formation, Pentz Road member

(informal), near Pentz, Butte County, northern Califor-

nia.

Etymology: Named for Oroville, California.

Discussion: The new species is based on two speci-

mens. The external surfaces are moderately well pre-

served, but the columellar lip and especially the deck

area are poorly preserved.

The new species is remarkably similar to Nerita

(Theliostyla) crooki Clark (1938: 700, pl. 4, figs. 1, 2)

from the Markley Formation east of San Francisco,

Solano County, northern California. Squires (2003: table

2.1, fig. 2.1) assigned this formation to the middle

Eocene (“Tejon Stage”). The new species differs from N.

(T.) crooki by having fewer and wider teeth on the

columellar lip, fewer ribs on the last whorl with relatively

wider interspaces, and ribs near the middle of the last

whorl not noticeably broader than adjacent ribs.

The new species is also very similar to Nerita

umzambiensis Woods (1906: 311, pl. 37, figs. 14-15;

Bandel and Kiel, 2003: 51-52, pl. 1, figs. 4-5) from

the Santonian/Campanian Umzamba Formation in

southeastern South Africa. The new species differs

from N. umzambiensis by having fewer teeth on the

columella lip, ribs on the base of the last whorl, and

a deck area sharply demarcated from the base of the last

whorl.

The new species somewhat resembles Nerita (The-

liostylaP) kennedyi Squires and Saul (2002: 185-187,

figs. 31-34) from the upper lower to lower middle

Eocene (“Domengine Stage”) Santiago Formation,

northern San Diego County, southern California. The

new species differs from N. (T.?) kennedyi by having

Page 137

beads that are not elongate, wider interspaces between

the ribs, and fewer, stronger, and wider teeth on the

columellar lip.

The only other early Campanian neritid known from

the Pacific slope of North America is Neritina (Dostia)

cuneata (Gabb, 1864: 137, pl. 21, fig. 97) from lower

Campanian strata at Tuscan Springs on Little Salt Creek,

Tehama County, northern Califormmia. Gabb’s species

might also be present in 1) upper Campanian and/or

lower Maastrichtian strata in the Pozo area, San Luis

Obispo County (Vedder, 1977) and 2) Maastrichtian

strata along the western edge of the San Joaquin Valley,

California (Woods and Saul, 1986). The new species is

vastly different from Neritina (Dostia) and does not have

its patelliform shape nor its distinctive collabral sculp-

ture.

Superfamily Cerithioidea Férussac, 1819

Family Indeterminate

Discussion: The new genus described below is most

likely a cerithioid, on the basis of its sigmoidal growth

lines, high spire, sculpture, short siphonal canal (slightly

twisted), smooth columella, and smooth interior of the

outer lip. Some specimens of the new genus have

a narrow spire, like that found in cerithioids, but other

specimens of the new genus have a buccinid-like shell.

The strongly sigmoidal growth lines of the new genus,

however, are unlike that found on buccinid shells. It is

possible that the new genus belongs to a new cerithioid

family.

Genus Bullamirifica new genus

Type Species: Bullamirifica elegans, new species;

Early Campanian, Pentz area, Butte County, northern

California.

Description: Shell medium (up to 83 mm height and

37 mm diameter, same specimen), fusiform to turreted.

Height to diameter ratio 2 to 2.7. Spire high, comprising

Al to 55% of total shell height. Pleural angle 33 to 42°.

Protoconch unknown. Teleoconch whorls six to eight.

Spire whorls with shoulder angulate; last whorl with

periphery angulate. Ramp short to moderately long,

concave to rarely straight-sloped. Suture slightly un-

dulatory, weakly impressed. Collabral sculpture consist-

ing of many narrow ribs, closely to moderately widely

spaced; interspaces smooth. Collabral ribs slightly

opisthocline to opisthocyrt, usually extending from

suture to suture. Collabral ribs present on base or

obsolete; if present, swollen and elongate. Spiral

sculpture consisting of several spiral ribs with variable

width and spacing, especially on last whorl. Spire whorls

with strongest spiral rib on whorl shoulder, several weak

or moderately strong spiral ribs occasionally near

anterior suture, and suture coincident with weak spiral

rib either bearing weak nodes or without nodes. Last

whorl sculpture with three to four widely spaced, strong

spiral ribs on periphery and one or two weaker spiral ribs

Page 138

or several spiral riblets on base. Intersections of collabral

and spiral ribs producing many nodes or strongly

projecting tubercles, either rounded (lmob-like) to

spinose or narrowly elongate. Nodes and knobs most

pronounced on whorl shoulder, especially on last whorl.

Intersections also strong on anterior portion of last whorl

periphery. Aperture short but moderately wide, comma-

shaped; small, narrow arch (canal-like) present where

outer lip meets most posterior part of aperture.

Columellar lip smooth. Siphonal canal short and spout-

like or well developed, moderately short, and can be

twisted to left. Outer lip thin, markedly sinuous, interior

smooth. Growth lines sigmoidal between suture and

shoulder; antispiral sinus coincident with tuberculate

spiral rib.

Geologic Age: Coniacian to early late Campanian.

Etymology: Combination of the Latin bulla, mean-

ing knob, and the Latin mirifus, meaning to cause

wonder.

Discussion: Three species can be herein assigned to

this new genus. Two of these species, Bullamirifica

verruca and Bullamirifica elegans, are based on entirely

new material. The third species was originally tentatively

assigned by Dailey and Popenoe (1966) to Pseudoglau-

conia Douvillé, 1921. Dailey and Popenoe (1966) stated

that this particular species belongs in a new genus, but

they withheld their description until better specimens

were obtained. Although representatives of the new

genus have sigmoidal growth lines (see Wenz, 1940: 764,

fig. 2214) similar to that of Psewdoglauconia, Bullamir-

ifica differs considerably from Pseudoglauconia by

having a bucciniform rather than a tapered/conical

shape, angulate rather than flat-sided whorls, and

tubercles on the sides of the whorls instead of only near

the suture, and the suture between the penultimate and

last whorl is not extremely deep and widely sunken.

Unfortunately, the aperture of Pseudoglauconia is not

known and none of the apertures on the available

specimens of Bullamirifica is complete. In spite of the

absence of knowledge about the aperture of Pseudo-

glauconia, Wenz (1940) believed this gastropod genus to

belong in the Cerithiidae.

The shape of the growth lines in Bullamirifica is

similar to that of Batillaria echinoides clavatulata

(Lamarck, 1804) from the middle Eocene (Lutetian) of

the Paris Basin, France. The siphonal canal of this

Eocene species is longer than normally found in

Batillaria Benson, 1842. Ponder and Warén (1988) and

Houbrick (1988) placed Batillaria in superfamily Cer-

ithioidea, family Batillariidae Thiele, 1929. The similarity

in growth-line shape between Bullamirifica and Batil-

laria suggests to us that the new genus might be

a cerithioid. The high turreted spire of Batillaria,

however, is quite unlike the lower, more paucispiral

spire of Bullamirifica.

Bullamirifica has the growth-line shape, shell shape,

sculpture, and twisted siphonal canal similar to that of

THE NAUTILUS, Vol. 119, No. 4

Pseudorapa Holzapfel, 1888, a monotypic genus from

the Vaals Greensand in the Netherlands and Germany.

The age of these strata was determined to be early

Campanian by Albers (1976). Wenz (1941: 1083,

fig. 3079) illustrated Pseudorapa. Bullamirifica differs

from it by having a less twisted siphonal canal and

more variable sculpture, including the possibilities of

having spiral ribs and a relatively narrow shell. In

addition, Pseudorapa has an outer lip that is very

crenulate and a spiral band near the base of the last

whorl that produces a tooth-like projection on the outer

lip.

Bullamirifica verruca new species

(Figures 9-12)

Diagnosis: Small Bullamirifica with prominent round-

ed knobs, nine on shoulder of last whorl. Suture

coincident with noded spiral rib. Base of last whorl

without elongate collabral ridges. Siphonal canal short

and straight.

Description: Small (up to 34 mm estimated height

and 16.8 mm diameter, same specimen), fusiform,

moderately slender. Height to diameter ratio approxi-

mately 2. Spire high, approximately 44% of shell height.

Pleural angle approximately 35 to 37°. Protoconch and

upper spire unknown. Teleoconch whorls approximately

six (estimated). Spire whorls with shoulder angulate, last

whorl with periphery angulate. Ramp short, concave.

Suture slightly impressed, possibly undulatory. Collabral

sculpture consisting of many ribs, widely spaced;

interspaces smooth. Collabral ribs slightly opisthocline

and extending from suture to suture. Collabral ribs

mainly prevalent on ramp. Spiral sculpture consisting of

several ribs with variable strength and spacing, especially

on last whorl. Intersections of collabral and spiral ribs

producing many strongly projecting rounded tuberculate

knobs or, less commonly, nodes; knobs and nodes most

prominent on shoulder and usually extending posteriorly

across ramp and become narrow ridges. Spire whorls

with strongest spiral rib on whorl shoulder, very faint

spiral riblets present between whorl shoulder and

anterior suture. Suture coincident with moderately weak

spiral rib bearing small nodes. Penultimate whorl with

spiral rib on shoulder bearing nine, moderately closely

spaced knobs. Last whorl with three spiral ribs on

periphery, strength of ribs progressively decreasing

anteriorly: strong rib on shoulder and bearing nine very

prominent tuberculate knobs; middle rib moderately

strong and bearing more numerous nodes (about half

sized of those on shoulder) that tend to become smaller

and even obsolete adaxially; and most anterior periphery

rib weakest and weak nodes tending to be obsolete

adaxially. Base of last whorl usually with one or two

spiral ribs, both bearing nodes (best developed near

outer lip) or bearing no nodes, and with strength of

spiral ribs decreasing anterior in direction; occasionally,

anterior region of base with only very weak spiral riblets.

R. L. Squires and L. R. Saul, 2005 Page 139

Figures 9-21. New cerithioid? gastropods. Specimens coated with ammonium chloride. 9-12. Bullamirifica verruca new genus and

species, LACMIP loc. $133. 9. Paratype LACMIP 13327, apertural view, height 27 mm, diameter 19.2 mm. 10-11. Holotype LACMIP

13326, height 29.7 mm, diameter 17.8 mm. 10. Apertural view. 11. Abapertural view. 12. Paratype LACMIP 13327, basal view,

diameter 18.9 mm. 13-17. Bullamirifica elegans new genus and species, LACMIP loc. 24337. 13-14. Holotype LACMIP 13328,

height 66.9 mm, diameter 34.9 mm. 13. Apertural view. 14. Abapertural view. 15-16. Paratype LACMIP 13329, height 60.1mm,

diameter 25.7 mm. 15. Apertural view. 16. Abapertural view. 17. Halisree LACMIP 13328, basal view, diameter 35.7 mm. 18-21.

Bullamirifica ainiktos (Dailey and Popenoe, 1966) new genus. 18-19. Hypotype LACMIP 13330, LACMIP loc. 10691, height 60 mm,

diameter 24.1 mm. 18. Apertural view. 19. Abapertural view. 20. Plasto-holotype LACMIP 40435, LACMIP loc. 24125, right- lateral

view, height 58.2 mm, diameter 29.1 mm. 21. Hyptotype LACMIP 13331, LACMIP loc. 24124, basal view, diameter 16.7 mm.

Page 140

Aperture round with very small posterior “arch.”

Aperture elliptical, columellar lip smooth, outer lip thin;

siphonal canal short and spout-like.

Holotype: LACMIP 13326, incomplete specimen

with two whorls (upper spire missing), 29.7 mm height,

17.8 mm diameter.

Paratype: LACMIP 13327.

Type Locality: LACMIP loc. 8133.

Coniacian.

Distribution: Redding Formation, Member IV, Oak

Run area, northern California.

Geologic Age:

Etymology: Latin verrucus, wart.

Discussion: This new species is based on three

specimens. It differs from Bullamirifica elegans new

species below by smaller size and presence ae rounded

rather than elongate knobs, fewer knobs on shoulder of

last whorl, no elongate collabral ridges on base of last

whorl, a noded spiral rib coincident with the suture, and

a straight siphonal canal. Bullamirifica verruca differs

from Bullamirifica ainiktos by being smaller with

a shorter spire and having rounded and much more

projecting nodes, many fewer nodes on shoulder of last

whorl, much less tendency for elongate collabral ridges

on ramp, and much weaker spiral lbs between shoulder

and anterior suture.

Bullamirifica verruca is very similar to the cerithioid

Tympanotonus (Tympanotonus) robustus Dockery

(1993: 47, pl. 7, fig. 1) in the shape of the spire, strong

nodes on the spire, and shape of the growth lines next to

the outer lip. Tympanotonus (T.) robustus, which is of

Campanian age and from Mississippi, however, has no

siphonal canal.

Bullamirifica elegans new species

(Figures 13-17)

[?] Pseudoglauconia? aff. P. ainiktos Dailey and Pope-

noe.—Elder and Saul, 1993: pl. 2, fig. 11.

Diagnosis: Large Bullamirifica with prominent

opisthocline collabral ridges on upper spire and base

of whorl. Shoulder of last whorl with 11 nodes. Suture

coincident with unnoded weak spiral rib. Siphonal canal

short but well developed and twisted to left.

Description: Medium large (up to 83 mm estimated

height and 37 mm diameter, same specimen), fusiform,

moderately wide, rarely slender. Height to diameter

ratio approximately 2 .2. Spire high, approximately 40%

of shell height. Pleural angle approximately 33 to 42°,

rarely approximately 30°. Sproroconel and uppermost

spire unknown. Teleoconch whorls approximately eight

(estimated). Whorls with angulate shoulder. Ramp short

and slightly concave. Collabral sculpture consisting of

many moderately strong, narrow ribs; interspaces

smooth. Collabral ribs extending from suture to suture.

THE NAUTILUS, Vol. 119, No. 4

Collabral ribs sigmoidal between posterior suture and

shoulder, opisthocline between shoulder and anterior

suture. Collabral ribs somewhat swollen on base of last

whorl. Spiral sculpture consisting of several ribs with

variable strength and spacing. Intersections of collabral

and spiral ribs producing many swollen, axially elongate

nodes or, in some cases, rounded tuberculate knobs:

nodes and knobs most prominent on shoulder. Spire

whorls with strongest spiral rib on whorl shoulder and

occasionally two moderately strong, noded spiral ribs

between whorl shoulder and anterior suture. Suture

coincident with weak spiral riblet. Penultimate whorl

with spiral rib on shoulder bearing 11 widely spaced

nodes. Last whorl with three or four spiral ribs on

periphery, strongest rib on shoulder and bearing 11

nodes. Spiral “alae on periphery of last whorl usually show

strength progressively decreasing anteriorly, occasionally

rib immediately anterior of shoulder weaker than other

ribs on periphery. Growth lines sigmoidal, with antispiral

sinus coincident with shoulder. Aperture elliptical,

columellar lip smooth, outer lip thin; siphonal canal

short but well developed and slightly twisted to left.

Holotype: LACMIP 13328; nearly complete specimen

of five whorls (upper spire missing), 66.9 mm height,

34.9 mm diameter.

Paratype: LACMIP 13329

Type Locality: LACMIP loc. 24337.

Geologic Age:

Campanian.

Early Campanian to possibly middle

Distribution: Early Campanian: Chico Formation,

Pentz Road member (informal), near Pentz, Butte

County, northern California; Possibly middle Campa-

nian: Pigeon Point Formation, San Mateo County,

northern California.

Etymology: Latin elegans,

choice.

meaning very fine or

Discussion: The new species is based on six speci-

mens from the Pentz area, and preservation is moder-

ately good on all of them. Nearly all the specimens are

moderately wide, but a few are somewhat slender, as the

specimen illustrated in Figures 15-16.

There might be one specimen of the new species from

the Pigeon Point Formation. This museum specimen,

which cannot be located, was figured by Elder and Saul

(1993: pl. 2, fig. 11). It was identified by them as

Pseudoglauconia® sp. aff. P. ainiktos Dailey and Pope-

noe, but it has a profile somewhat more like Bullamir-

ifica elegans and basal nodes like B. elegans. As will be

dhecuanedl later, there are specimens of B. ainiktos from

the Pigeon Point Formation. It seems likely that some of

the Bullamirifica specimens from the Pigeon Point

Formation represent transitional forms iekeeen B.

elegans and B. ainiktos.

Bullamirifica elegans differs from Bullamirifica ver-

ruca in having larger size, prominent collabral ribs

R. L. Squires and L. R. Saul, 2005

extending from suture to suture on the spire whorls,

swollen collabral ribs on the neck, more nodes on

shoulder of the penultimate and last whorls, and

siphonal canal longer and slightly twisted to the left. In

addition, B. elegans occasionally has strong unnoded

spiral ribs between suture and shoulder.

Bullamirifica elegans differs from Bullamirifica ainik-

tos by usually having a wider pleural angle, usually a less

elevated spire, wider and stronger nodes on shoulder,

fewer nodes on shoulder of last whorl, and base with

swollen collabral ribs instead of fine spiral ribs.

Bullamirifica elegans also has variability in morphology,

whereas B. ainiktos does not.

Bullamirifica elegans is very similar to the cerithioid

Tympanotonus (Tympanotonus) binodosus Dockery

(1993: 47, pl. 7, fig. 2) in the shape of the spire, strong

nodes on the spire, and shape of the growth lines next to

the outer lip. Tympanotonus (T.) binodosus of Campa-

nian age and from Mississippi, however, has no siphonal

canal.

Bullamirifica ainiktos (Dailey and Popenoe, 1966) new

combination

(Figures 18—25)

Pseudoglauconia? ainiktos Dailey and Popenoe, 1966:

21-22, pl. 6, figs. 3, 5, 6.

Pseudoglauconia? aff. P. ainiktos Dailey and Popenoe.—

Elder and Saul, 1993: pl. 2, fig. 10.

Cimolithium miyakoense (Nagao, 1934). — Perrilliat-

Montoya, 1968: 20, pl. 4, fig. 2

Vicarya (Shoshiroia) yabei erie, 1960.— Perrilliat-

Montoya, 1968: 21, pl. 6, fig. 1.

Diagnosis: Medium-size Bullamirifica with usually

slender turreted whorls. Ramp on upper spire short,

with narrow collabral ribs crossed by weaker spiral ribs;

ramp on later whorls much longer and smooth and

coneave. Spire whorls concave on posterior half but

angulate medially, with moderately strong tubercles.

Shoulder of last whorl with 14 nodes. Base of last whorl

with many, closely spaced spiral riblets. Siphonal canal

short and possibly straight.

Description: Medium, up to 67.5 mm height estimat-

ed and 25.4 mm diameter, same specimen. Shell usually

slender, turreted, occasionally wide, fusiform. Height to

diameter ratio approximately 2.7. Spire high, approxi-

mately 55% of total shell height. Pleural angle approx-

imately 33 to 37°. Protoconch unknown. Teleoconch

whorls approximately eight (estimated). Spire whorls

with shoulder angulate. Ramp usually short and concave,

occasionally long and straight-sloped. Suture obscured.

Collabral sculpture consisting of many spiral ribs closely

spaced and narrow. Collabral ribs opisthocyrt, most

prominent on ramp. Spiral sculpture consisting of

several ribs, moderately weak and closely spaced.

Intersections of collabral and spiral ribs producing many

nodes, most prominent and somewhat spinose on

shoulder. Spire whorls with strongest spiral rib on whorl

Page 141

shoulder and five moderately strong, non-noded spiral

ribs between shoulder and anterior suture; most anterior

one of these ribs coincident with suture. Penultimate

and last whorls with approximately 14 nodes on

shoulder. Last whorl with three spiral ribs on periphery,

progressively weaker anteriorly, and bearing nodes. Base

of last whorl bearing fine spiral riblets. Siphonal canal

short and twisted to left. Columellar lip smooth. Outer

lip sinuous; interior smooth.

Holotype: LACMIP 40435, incomplete specimen (tip

of spire and siphonal canal missing), 39 mm height,

29 mm diameter.

Type Locality: LACMIP loc. 4125.

Geologic Age:

nian.

Middle Campanian to Late Campa-

Distribution: Middle Campanian: Pigeon Point For-

mation, near Pigeon Point, San Mateo County, northern

California. Middle late Campanian: Punta Baja Forma-

tion, southwest of El Rosario, Baja Califormia, Mexico.

Late Campanian: Jalama Formation, Santa Barbara

County, southern California.

Discussion: Description of the new species is based

on 25 specimens: 18 from the Jalama Formation, five

from the Pigeon Point Formation, and two from the

Punta Baja Formation. The Jalama specimens have poor

to moderately good preservation, although the apices are

broken off aun the siphonal canal missing. The Pigeon

Point specimens also have poor to moderately good

preservation, but the matrix is very hard to remove. The

Punta Baja specimens have poor preservation.

The Pigeon Point Formation specimen illustrated by

Elder and Saul (1993: pl. 2, fig. 10) has a whorl profile

like that found on B. ainltos but it has wider pleural

angle, similar to that found on B. elegans. Unfortunately

ie specimen is missing. As wareationied earlier, it seems

likely that some of the Bullamirifica specimens from the

Pigeon Point Formation represent transitional forms

between B. elegans and B. ainiktos.

Bullamirifica. ainiktos differs from Bullamirifica ver-

ruca by larger size and having less rounded and much

lower meres, more nodes on shoulder of last whorl,

much more tendency for elongate collabral ridges on

ramp, and much stronger spiral ribs between shoulder

and anterior suture.

Bullamirifica ainiktos differs from Bullamirifica ele-

gans by smaller size, having a narrower spire, usually

a more elevated spire, narrower and weaker nodes on

shoulder, more nodes on shoulder of last whorl, and base

with fine ribs instead of swollen collabral ribs.

Perrilliat-Montoya (1968) identified one of the Punta

Baja Formation specimens (Figures 23-25 herein) of B.

ainiktos as Vicarya (Shoshiroa) yabei and identified the

other specimen as Cimolithium miyakoense. With Wenz

(1940: 740-741, fig. 2145) as a basis for comparison, we

consider that Bullamirifica ainiktos differs considerably

from the former gastropod by having a non-conical shell,

Page 142 THE NAUTILUS, Vol. 119, No. 4

Figures 22-34. New cerithioid? and trichotropid gastropods. Specimens coated with ammonium chloride, unless otherwise noted.

22-25. Bullamirifica ainiktos (Dailey and Popenoe, 1966) new genus. 22. Hypotype LACMIP 13332, USGS loc. M-8601, height

35.8 mm, diameter 20 mm. 23-25. Plasto-hypotype IGM 1325, height 80 mm, diameter 36.9 mm. 23. Apertural view. 2d.

Abapertural view. 25. Basal view. 26-29. Minytropis melilota new genus and species. 26. Paratype LACMIP 13334, LACMIP loc.

10787, apertural view, height 7 mm, diameter 2 mm. 27-29. Holotype LACMIP 13333, LACMIP loc. 10786, abapertural view,

height 62 mm, diameter 2.2 mm. 27. Abapertural view. 28. Oblique apertural view of tip; arrow indicates where protoconch ends

and teleoconch starts. 29. Protoconch, height 0.5 mm, diameter 0.8 mm; arrow indicates where protoconch ends and teleoconch

starts; SEM photomicrograph (onecated. 30- 34. Paxitropis dicriota new genus and species. 30-31. Paratype LACMIP 13336,

LACMIP loc. 23643, height 12.7 mm, diameter 6.8 mm. 30. Apertural view. 31. Abapertural view. 32. Holotype LACMIP 13335,

LACMIP loc. 23639, apertural view, height 14.7 mm, diameter 6.6 mm. 33. Paratype LACMIP 13337, LACMIP loc. 23642,

apertural view, height 10.1 mm, diameter 4.4 mm. 34. Paratype LACMIP 13338, LACMIP loc. 24217, abapertural view, height

11 mm, diameter 4.3 mm.

R. L. Squires and L. R. Saul, 2005

more rows of nodes, and absence of a notch on the outer

lip near the suture and not having a thick, extensive

callus that covers part of the noded sculpture near the

suture on the last whorl. Using Kase (1984: 135-136, pl.

21, figs. 1-9; fig. 20) as a basis for comparison, we

consider that Bullamirifica ainiktos differs considerably

from the latter gastropod by having a much lower spire,

more strongly sinuous growth lines, and a well de-

veloped, twisted siphonal canal.

Superfamily Capuloidea Fleming, 1822

Family Capulidae Fleming, 1822

PSubfamily Trichotropinae Gray, 1850

Discussion: We include a subfamily name because

high-spired capulids, like those described below, are so

distinctive from cap-shaped capulids. The subfamily

allocation is tentative, pending much-needed taxonomic

work on capulids.

Genus Minytropis new genus

Type Species: Minytropis melilota new species; Late

Cretaceous, Santonian, northern California.

Description: Shell very small, estimated maximum

height just over 10 mm. Shell thin, narrowly fusiform-

elongate. Protoconch flat-topped and smooth, approxi-

mately two whorls. Teleoconch whorls rounded. Sculp-

ture consisting of several strong spiral ribs with wide

interspaces, both ribs and interspaces crossed by well

delineated prosocline growth lines. Aperture oval, pro-

duced anteriorly to ohare with narrow siphonal canal.

Outer lip thin with no varix or inner denticulations.

Inner lip callused, narrow.

Geologic Age:

Etymology: Combination of the Greek minys, mean-

ing little, small, or short; and the Latin tropis, meaning

keel.

Discussion: The placement of Minytropis among the

trichotropids is suggested by the strongly prosocline and

well marked growth lines, strong spiral sculpture, and

the short, open siphonal canal. Minytropis resembles

Opposirius Iredale, 1931, which is known only from the

Recent of Australia (Wenz, 1940). Minytropis differs

from Opposirius by having a narrower aperture, a longer

siphonal canal, and a rounded rather than a bladelike

inner lip.

Finlay and Marwick (1937) reported that Opposirius is

more similar to Certhioderma Conrad, 1860, than to any

other trichotropid genus. In comparison to Certhioderma,

Minytropis is similar in having a flat-topped, smooth

protoconch and in having a short siphonal canal, but

Minytropis differs by having a much narrower shell,

longer and much narrower aperture, stronger spiral

sculpture, sculpture not recticulate on adult whorls, and

no hint of an umbilicus. Wenz (1940) reported Cerithio-

derma to have a geologic range of Late Cretaceous to

Early and Late Santonian.

Page 143

Oligocene, but Marwick (1965) reported that this genus is

also extant. Dockery (1993) reported Cerithioderma from

Campanian strata in Mississippi. High-spired trichotro-

pids very similar to Cerithioderma are best represented in

New Zealand Tertiary strata (Maxwell, 1992). According

to Maxwell (1992), Trichosirius Finlay, 1926, might be

a junior synonym of Cerithioderma.

Minytropis melilota new species

(Figures 26-29)

Description: Small, up to 10.2 mm estimated height

and 3.5 mm diameter, same specimen. Shell narrowly

fusiform-elongate. Spire high, 60% (estimated) of shell

height. Plena angle approximately 22°. Protoconch flat-

topped, with two whorls, and smooth (0.5 mm height,

0.8 mm diameter). Teleoconch of approximately seven

to eight subangulate whorls, last whorl nearly one-third

total height. Satire deeply impressed, nearly coincident

with very weak rib. Sculpture nearly cancellate on

earliest teleoconch whorl, but four spiral riblets slightly

stronger than numerous collabral threads. Sculpture on

subsequent whorls progressing into five strong, flat-

topped ribs on middle spire, six ribs on lower spire, and

approximately eight to nine ribs on last whorl; most

posterior spiral alo weak to very weak on all these whorls.

Interspaces of nearly equal width on spire whorls but

becoming twice as wide as ribs on last whorl. On last

whorl, ribs on whorl face equally strong but on base, ribs

become much less prominent and more closely spaced.

Growth lines strongly prosocline, marked by fine

collabral threads strongest on early whorls where they

produce slight beading of spiral ribs. Growth lines less

prominent and closer-spaced on later whorls; growth

lines can be somewhat prominent on base of last whorl.

Aperture oval, produced anteriorly to short, narrow

siphonal canal. Outer lip thin, arcuate, crenulated by

ribs. Columellar lip narrow, weakly callused, and barely

forming a pseudo-umbilical chink between base of whorl

and indistinct fasciole.

Dimensions of Holotype: Nearly complete specimen

(siphonal canal mostly missing), 6.2 mm height, 2.2 mm

diameter.

Holotype: LACMIP 13333.

Type Locality: LACMIP loc. 10786.

Paratype: LACMIP 13334.

Geologic Age: Early and Late Santonian.

Distribution: Lower Santonian: Redding Formation,

Member V, Clover Creek, Shasta County, northern

California. Upper Santonian: Chico Formation, Musty

Buck Member, Chico Creek, Butte County, northern

California.

Etymology: Latin melitota, meaning sweet clover (in

reference to the occurrence of the new species near

Clover Creek).

Page 144

THE NAUTILUS, Vol. 119, No. 4

Discussion: Description of this new species is based

on 29 specimens, most of which represented by small

fragments. The better preserved specimens are all from

the Redding Formation, however, none of these speci-

mens is as large as the larger ones from the Chico

Formation localities. Only two specimens, both from

LACMIP loc. 10786, have retained the protoconch. The

protoconch of one of these specimens is illustrated in

Figures 27-29. The other specimen’s protoconch is

mostly embedded in hard matrix. The flat-topped

protoconch of M. meliota supports the position of this

gastropod among the Trichotropidae.

The new species can be distinguished from the similar

looking Opposirius idoneus Iredale (1931: 210, pl. 22,

fig. 7; Wenz, 1940: fig. 2631), an extant species in

southwestern Australia, by having a narrower shell, one

more rib on the spire, two to four more ribs on the base

of the last whorl, a much longer siphonal canal, an

a rounded columella. Opposirius idoneus is the type

species of Opposirius Iredale, 1931.

Genus Paxitropis new genus

Type Species: Paxitropis dicriota; Late Cretaceous,

late Santonian to early Campanian, northern Califor-

nia.

Description: Shell medium small, narrowly pagodi-

form. Spire whorls with prominent keel medially. Suture

bordered posteriorly by relatively weak spiral rib

becoming keel-like on last whorl and, in concert with

aforementioned keel, producing bicarinate last whorl.

Shell base with several weak spiral riblets. Growth lines

prosocline. Columellar lip callused, abaxial margin well-

delineated. Siphonal canal short but distinct. Anompha-

lous.

Geologic Age: Late Santonian to early Campanian.

Etymology: Combination of the Latin pax, meaning

peace (in reference to the peaceful Pacific coast); and

the Latin tropis, meaning keel.

Discussion: Paxitropis is similar to Trichotropis Bro-

derip and Sowerby, 1829, a genus known (Sohl, 1960)

from the Late Cretaceous to Recent. The modern

distribution of Trichotropis is restricted to circum-boreal

waters, in both the northern and southern oceans and, as

Sohl (1960) noted, this modern distribution is in sharp

contrast to its Cretaceous distribution in temperate

shallow-water deposits. Paxitropis differs from Tricho-

tropis by having a smaller size, much narrower shell

(including a narrower aperture), a siphonal canal, much

lower variability in sculpture, and being anomphalous.

Paxitropis resembles the trichotropid Icuncula Ire-

dale, 1924, which is known from the Recent of the Indo-

Pacific (Wenz, 1940). Paxitropis differs from Icuncula by

having no hint of an umbilicus, less prominent keels that

are unwavy, one less keel on the last whorl, and no axial

riblets on the ramp. The aperture of Paxitropis is not

entirely known.

The new genus has a whorl profile very similar to the

turrid Austrocarina Laseron, 1954, known only from the

Recent of New South Wales and Victoria, southeastern

Australia and Tasmania. Paxitropis, however, is not

a turrid because it does not have the turrid growth lines,

which tend to be opisthocline anterior to the notch and

across the whorl periphery curving toward prosocline

very near the siphonal area.

Paxitropis dicriota new species

(Figures 30-34)

Description: Shell small, up to height estimated

14.5 mm and 6.6 mm diameter. Shell pagodiform. Spire

high, approximately 60% of shell height. Pleural angle

approximately 30°. Protoconch unknown. Teleoconch

approximately six strongly angulated whorls, with wide

ramp on middle spire whorls and on last whorl. Sculpture

dominated by prominent spiral rib, medially located on

shoulder and keel-forming on later whorls. Early

teleoconch whorls with two spiral ribs anterior to carina.

On mature whorls, these additional ribs become mere

threads and equivalent in strength to growth lines, thus

producing microscopic cancellate surface, especially on

ramp. Suture impressed, nearly coincident with weakly

noded subsutural spiral rib located immediately posterior

to suture. Subsutural spiral rib relatively weak on spire

whorls but becoming keel-like on last whorl. Last whorl

with two sharp keels, one just above middle whorl height

and second one weaker and occasionally minutely noded,

emergent at anterior suture. Shell base with several weak

spiral ribs. Growth lines prosocline, most prominent on

shell base, near outer lip. Aperture lanceolate and

produced anteriorly to short, but distinct siphonal canal.

Outer lip thin, angulate, crenulated by ribs. Columellar

lip callused, abaxial margin well-delineated. Columellar

lip narrow, barely forming pseudo-umbilical chink

between base of whorl and indistinct fasciole.

Holotype: LACMIP 13335, 14.7 mm height, 6.6 mm

diameter.

Paratypes: LACMIP 13336, 13337, and 13338.

Type Locality: LACMIP loc. 23643.

Geologic Age: Late Santonian to Early Campanian.

Distribution: Upper Santonian: Redding Formation,

Member VI, area east of Redding, Shasta County,

northern California. Lower Campanian: Chico Forma-

tion, Ten Mile Member, Chico Creek, Butte County,

northern California; Ladd Formation, upper Holz Shale

Member, Santa Ana Mountains, Orange County, south-

ern California.

Etymology: Combination of the Greek di, meaning

two, and the Greek criota, meaning ringed.

Discussion: Twenty incomplete specimens were avail-

able, and the six most complete of these are the basis for

the above description. The only specimen from the

R. L. Squires and L. R. Saul, 2005

Redding Formation is geologically the oldest specimen

(LACMIP loc. 24217). The only specimen from the

upper Holz Shale is from LACMIP loc. 21536.

The new species is very similar to the Pliocene to

Recent Trichotropis bicarinata (Sowerby, 1825; Pitt and

Pitt, 1989: pl. 1, figs. 3-4; Egorov and Alexeyev, 1998:

25, figs. 22-23), which is a Pacific upper boreal Arctic

species, occurring in the waters off northern Japan,

Kamchatka, southern Chukchi Sea, and Queen Char-

lotte Islands in British Columbia (Egorov and Alexeyev,

1998). The new species differs from T. bicarinata

by having a much narrower last whorl and a much

narrower aperture. These same differences distinguish

the new species from Trichotropis vokesae Pitt and Pitt,

1989, which is extremely similar to T. bicarinata.

Trichotropis vokesae is of Pliocene age and from

northwestern Ecuador, and the new species differs from

T. vokesae in the same ways that it differs from T.

bicarinata.

The new species can be distinguished from the similar

looking Icuncula torcularis (Tenison-Woods 1879;

Wenz, 1940: fig. 2630), an extant species living off the

coasts of southeastern Australia and Tasmania, by having

less projecting and non-wavy keels, one less keel on the

last whorl, and an absence of axial riblets on the ramp.

Icuncula torcularis is the type species of Icuncula

Iredale, 1924.

The new species is remarkably similar to the extant

Austrocarina recta (Hedley, 1903; 1922: 223, fig. 3;

Powell, 1966: 34, pl. 3, fig. 2; 1969: 364, pl. 276; Wilson,

1994: 187) but differs from A. recta by having a shell

approximately 14 mm in height instead of 6 mm, a more

projecting and usually wider keel on the shoulder of the

adult whorls, several weak spiral ribs on base of last

whorl, several spiral ribs on earliest teleoconch whorls,

straight rather than a curved profile of the base,

columellar lip callused with abaxial margin well-de-

lineated, and no tendency to develop collabral ribs on

earliest teleoconch whorls.

ACKNOWLEDGMENTS

The authors are especially grateful for the careful and

tireless collecting of Eric Géhre of Oroville, California.

Without his help and willingness to donate the material,

the paper would have been greatly reduced in its scope.

James H. McLean, Natural History Museum of Los

Angeles County, Malacology Division, provided valuable

comments about the identification of the new trochid.

Carmen Perrilliat (IGM) graciously sent us excellent

casts of the Punta Baja Formation material. David Haasl

(UCMP) spent considerable effort unsuccessfully trying

to locate the hypotypes of Bullamirifica ainiktos from the

Pigeon Point Formation. Jorge Vazquez, California State

University, Northridge, kindly took SEM photomicro-

graphs of the protoconch of Minytropis melilota. Steffen

Kiel (Smithsonian Institution) and an anonymous re-

viewer critiqued the manuscript.

Page 145

LITERATURE CITED

Adams, A. 1855. Further contributions toward the natural

history of the Trochidae: with the description of a new

genus and of several new species, from the Cumingian

collection. Proceedings of the Zoological Society of

London for 1854: 33-41.

Addicott, W. O. 1973. Oligocene molluscan biostratigraphy and

paleontology of the lower part of the type Temblor

Formation, California. U. S. Geological Survey Profes-

sional Paper 791: 1-48.

Albers, H. J. 1976. Feinstratigraphie, Faziesanalyse und Zyklen

des Untercampans (Vaalser Griinsands = Hervien) von

Aachen und dem niederlindisch-belgischen Limburg.

Geologisches Jahrbuch, A, 34, 3-68.

Almgren, A. A. 1986. Benthic foraminiferal zonation and

correlations of Upper Cretaceous strata of the Great

Valley of California—a modification. In P. L. Abbott (ed.),

Cretaceous Stratigraphy Western North America. Pacific

Section, Society of Economic Paleontologists and Miner-

alogists 46: 137-152.

Anderson, F. M. and G. D. Hanna. 1935. Cretaceous geology

of Lower California. Proceedings of the California

Academy of Sciences, Series 4, 23(1): 1-34.

Arnold, R. 1908. Descriptions of new Cretaceous and Tertiary

fossils from the Santa Cruz Mountains, California.

Proceedings of the United States National Museum 34:

345-390.

Bandel, L. K. and S. Kiel. 2003. Relationships of Cretaceous

Neritimorpha (Gastropoda, Mollusca), with the descrip-

tion of seven new species. Bulletin of the Czech

Geological Survey 78: 49-62.

Bandel, K. and W. Stinnesbeck. 2000. Gastropods of the

Quiriquina Formation (Maastrichtian) in central Chile:

paleobiogeographic relationships and the description of

a few new taxa. Zentralbatt fiir Geologie und Paliontolo-

gie Teil 1(Heft 7/8): 757-788.

Benson, W. H. 1842. Mollusca. In: T. Cantor (ed.) General

Features of Chusan, with Remarks on the Flora and

Fauna of that Island. Annals and Magazine of Natural

History, Series 1, 9: 486-490.

Boehlke, J. E. and P. L. Abbott. 1986. Punta Banda Formation,

a Campanian submarine canyon fill, Baja California,

Mexico. In: P. L. Abbott (ed.) Cretaceous Stratigraphy

Western North America. Pacific Section, Society of

Economic Paleontologists and Mineralogists 46: 91-101.

Broderip, W. J. and G. B. Sowerby, I. 1829. Observations on

new or interesting Mollusca contained, for the most part,

in the Museum of the Zoological Society. Zoological

Journal 4(15): 359-376.

Clark, B. L. 1938. Fauna from the Markley Formation (upper

Eocene) on Pleasant Creek, California. Bulletin of the

Geological Society of America 49: 683-730.

Conrad, T. A. 1860. Descriptions of new species of Cretaceous

and Eocene fossils of Mississippi. Journal of the Academy

of Natural Sciences of Philadelphia, Series 2, 4(3):

275-298.

Dailey, D. H. and W. P. Popenoe. 1966. Mollusca from the

Upper Cretaceous Jalama Formation, Santa Barbara

County, California. University of California Publications

in Geological Sciences 65: 1-27.

Davies, A. M. 1971. Tertiary Faunas, a text-book for oilfield

palaeontologists and students of geology, Volume 1, The

Composition of Tertiary Faunas, revised by F. E. Eames.

George Allen and Unwin, London, 571 pp.

Page 146

THE NAUTILUS, Vol. 119, No. 4

Dockery, D. TIII. 1993. The streptoneuran gastropods

exclusive of the Stenoglossa, of the Coffee Sand (Campa-

nian) of northeastern Mississippi. Mississippi Department

of Environmental Quality, Office of Geology, Bulletin 129:

1-191.

Douvillé, H. 1921. Mélanges paleontologiques: genre Eova-

sum, Glaunconiidae, Pleuroceratidae’ Pirenidae, genre

Itruvia. Journal de Conchyliologie 66: 1-18.

Egorov, R. and D. Alexeyev. 1998. Treasure of Russian Shells.

Volume 2 . Trichotropidae Moscow, 36 pp.

Elder, W. P. P, amd L. R. Saul. 1993. Paleogeographic implica-

tions a molluscan assemblages in the Upper Cretaceous

(Campanian) Pigeon Point Formation, California. In: G.

Dunne and K. McDougall (eds.) Mesozoic Paleogeogra-

phy of the Western United States-II. Pacific Section,

Society of Economic Paleontologists and Mineralogists 71:

Dp. Ui 186.

Elder, W. P. and L. R. Saul. 1996. Taxonomy and _bio-

Lane of Coniacian through Maastrichtian Anchura

(Gastropoda: Aporrhaidae) of the North American Pacific

slope. Journal of Paleontology 70: 381-397.

Férussac, D. 1819. Histoire naturelle générale e pariticuliére

des mollusques terrestres et fluviatiles. Volume 1, Paris,

128 pp.

Finlay, H. J. 1926. A further commentary on New Zealand

mollusean systematics. Transactions of the New Zealand

Institute 57: 320-485.

Finlay, H. J. and J. Marwick. 1937. The Wangaloan and

associated molluscan faunas of Kaitangata-Green Island

subdivision. New Zealand Department of Scientific and

Industrial Research, Geological Survey Branch, Palaeon-

tological Bulletin 15: 1-140.

Fleming, J. 1822. Philosophy of Zoology. Volume 2. Edin-

burgh, 618 pp.

Gabb, W. M. 1864. Description of the Cretaceous fossils.

Geological Survey of California, Palaeontology 1: 57-243.

Goudkoff, P. P. 1945. Stratigraphic aaleiong of Upper

Cretaceous in the Great Valley, California. American

Association of Petroleum Geologists Bulletin 29:

956-1007.

Gradstein, F. M., J. G. Ogg and A. G. Smith. 2004. A geologic

time scale 2004. Cambridge University Press, Cambridge,

589 pp.

Grant, U. S., IV and H. R. Gale. 1931. Catalogue of the marine

Pliocene and Pleistocene Mollusca of California and

adjacent regions. Memoirs of the San Diego Society of

Natural History 1: 1-1036.

Gray, M. E. 1850. Figures of molluscous animals, selected

from various authors. Volume 2, London, pls. 79-199.

Groves, L. T. 2004. New species of Late Cretaceous

Cypraeidae (Gastropoda) from California and British

Columbia and new records from the Pacific slope. The

Nautilus 118: 43-51.

Hedley, C. 1903. Scientific results of the trawling expedition of

H.M.CS. “Thetis” off the coast of New South Wales in

February and March, 1896. Mollusca. Part II. Scaphopoda

and Gastropoda. Memoirs of the Australian Museum 4:

327-402.

Hedley, C. 1922. A revision of the Australian Turridae.

Records of the Australian Museum 13(6): 213-259.

Hickman, C. S. and J. H. McLean. 1990. Systematic revision

and suprageneric classification of trochacean gastropods.

Natural History Museum of Los Angeles County. Science

Series 35: 1-169.

Holzapfel, E. 1888. Die Mollusken der Aachener Kreide.

Paleontographica 34: 29-180.

Houbrick, R. S. 1988. Cerithioidean phylogeny. In: W. F.

Ponder, D. J. Eernisse and J. H. Waterhouse (eds),

Prosobranch Phylogeny. Malacological Review, Supple-

ment 4: 88-128.

Iredale, T. 1924. Results from Roy Bell’s molluscan collections.

Proceedings of the Linnaean Society of New South Wales

49: 179-278.

Iredale, T. 1931. Australian molluscan notes. I. Records of the

Australian Museum 18: 201-235.

Kamada, Y. 1960. On the associated occurrence of Vicarya and

Vicaryella in the Japanese Tertiary, with the first de-

scription of a Paleogene species of Vicarya from Japan.

Science Reports of Tohoku University, Series 2, Special

Volume 4: 281-295.

Kase, T. 1984. Early Cretaceous marine and brackish-water

Gastropoda from Japan. National Science Museum,

Tokyo, 263 pp.

Keen, A. M. 1960. Superfamily Trochacea Rafinesque, 1815.

In: R. C. Moore (ed.) Treatise on Invertebrate Paleon-

tology. Pt. I. Mollusca 1. Geological Society of America

and University of Kansas Press, Lawrence, pp. 1246-

1275.

Kiel, S. and K. Bandel. 2001. Trochidae (Archaeogastropoda)

from the Campanian of Torallola in northem Spain. Acta

Geologica Polonica 51(2): 137-154.

Kilmer, F. H. 1963. Cretaceous and Cenozoic stratigraphy and

paleontology, El Rosario area. University of California,

Berkeley, unpublished Ph. D. dissertation, 149 pp.

Kuroda, A. T., T. Habe and K. Oyama. 1971. The Sea Shells of

Sagami Bay. Maruzen Co., Ltd., Tokyo, 1281 pp.

Lamarck, J. B. 1804. Mémoires sur les fossiles des environs de

Paris. Annales de Muséum National d’Histoire Paris.

Tome 5, variously paged. Reprinted 1978, Paleontological

Research Institution, Ithaca, New York.

Laseron, C. F. 1954. Revision of the New South Wales

TurridaeAustralian Zoological Handbook. Royal Zoologi-

cal Society of New South ’ Wales, Sydney, 56 pp.

Lesson, R. P. 1835. Illustrations de zoologie ou recueil de

figures d’animaux peintes d’apres. Paras, 17 pls.

Linnaeus, C. 1758. Systema naturae per regna tria naturae.

Regnum animale. Editio decima reformata. Volume 1.

Laurentii Salvii, Stockholm, 824 pp.

McLean, J. H. 1978. Marine shells of southern California,

revised edition. Natural History Museum of Los Angeles

County, Science Series 24: 1-104.

Marwick, J. 1965. Upper Cenozoic Mollusca of Wairoa district,

Hawke’s Bay. New Zealand Geological Survey Paleonto-

logical Bulletin 39: 1-83.

Maxwell, P. A. 1992. Eocene Mollusca from the vicinity of

McCulloch’s Bridge, Waiho River, South Canterbury,

New Zealand: paleoecology and systematics. New Zealand

Geological Survey Paleontological Bulletin 65: 1-280.

Montfort, P. D. 1810. Conchyliologie systématique et classifi-

cation méthodique des coquilles. Volume 2. F. Schoell,

Paris, 176 pp.

Nagao, T. 1934. Cretaceous Mollusca from the Miyako

District, Honshi, Japan. Journal of the Faculty of Science.

The Hokkaido Imperial University, Series 4, 2: 177-277.

Perrilliat-Montoya, M. C. 1968. Fauna del Cretacio y del

Terciario del norte de Baja California. Univ ersidad

Nacional Auténoma de México, Instituto de Geologa,

Paleontologa Mexicana 25: 1-26.

R. L. Squires and L. R. Saul, 2005

Page 147

Philippi, R. A. 1887. Die Tertiiren und Quartiiren versteiner-

ungen Chiles. F. A. Brockhaus, Leipzig, 266 pp.

Pitt, W. D. and L. J. Pitt. 1989. A new species of Trichotropis

(Gastropoda: Mesogastropoda) from the Esmeralda beds,

Onzole Formation, northwestern Ecuador. Tulane Studies

in Geology and Paleontology 22: 131-136.

Ponder, W. F. 1998. Superfamily ‘Capuloidea. In: P. L. Beesley,

G. J. B. Ross and A. Wells (eds.), Mollusca: the Southern

Synthesis. Fauna of Australia. Volume 5. CSIRO Publish-

ing, Melbourne, Part B, pp. 774-775.

Ponder, W. F. and A. Warén. 1988. Appendix. Classification of

the Caenogastropoda and Heterostropha—a list of the

family-group names and higher taxa. In: W. F. Ponder, D. J.

Eernisse and J. H. Waterhouse (eds.) Prosobranch

Phylogeny. Malacological Review, Supplement 4: 288-326.

Powell, A. W. B. 1966. Tine molluscan families Speightiidae and

Turridae: An evaluation of the valid taxa, both Recent and

fossil, with lists of characteristic species. Bulletin of the

Auckland Institute and Museum 5: 1-184.

Powell, A. W. B. 1969. The family Turridae in the Indo-Pacific.

Part 2. The subfamily Turriculinae. Indo-Pacific Mollusca

9(10): 207-416.

Quintero, I. and J. Revilla. 1966. Algunas especies nuevas y

otra poco conocidas. Notas y Comunicaciones del

Instituto Geologico y Minero de Espana 82: 27-86.

Rafinesque, C. S. 1815. Analyse de la Nature ou Tableau de

Univers et des Corps Organisées. Barracvecchia, Palermo,

224 pp

Saul, L. R. 1983. Turritella zonation across the Cretaceous-

Tertiary boundary, California. University of California

Publications Geological Sciences 125: 1-165.

Saul, L. R. and W. P. Popenoe. 1992. Pacific slope Cretaceous

bivalves of the genus Calva. Natural History Museum of

Los Angeles County, Contributions in Science 433: 1-68.

Saul, L. R. and R. L. Squires. 1997. New species of neritid

gastropods from Cretaceous and Lower Cenozoic strata of

the Pacific slope of North America. The Veliger 40:

131-147.

Saul, L. R. and R. L. Squires. 2003. New Cretaceous

cerithiform gastropods from the Pacific slope of North

America. Journal of Paleontology 77: 442-453.

Smith, J. P. 1900. The development and phylogeny of

Placenticeras. Proceedings of the California Academy of

Sciences, Series 3, 1(7): 180-240.

Sohl, N. F. 1960. Archeogastropoda, Mesogastropoda and

stratigraphy of the Ripley, Owl Creek, and Prairie Bluff

formations. U. S. Geological Survey Professional Paper

331-A: 1-151.

Sowerby, G. B., I. 1825. A catalogue of the shells contained in

the collection of the late Earl of Tankerville. London, 92

Squires, R. L. 2003. Turnovers in marine gastropod faunas

during the Eocene-Oligocene transition, west coast of the

United States. In: D. R. Prothero, L. C. Ivany and E. A.

Nesbitt (eds.) From Greenhouse to Icehouse: the Marine

Eocene-Oligocene Transition. New York: Columbia Uni-

versity Press, pp. 14—35.

Squires, R. L. and L. R. Saul. 1997. Late Cretaceous

occurrences on the Pacific slope of North America of

the melanopsid gastropod genus Boggsia Olssen, 1929.

The Veliger 40: 193-202.

Squires, R. L. and L. R. Saul. 2001. New Late Cretaceous

gastropods from the Pacific slope of North America.

Journal of Paleontology 75: 46-65.

Squires, R. L. and L. R. Saul. 2002. New information on Late

Cretaceous, Paleocene, and Eocene neritid gastropods

from the North American Pacific slope. The Veliger 45:

177-192.

Squires, R. L. and L. R. Saul. 2003a. Additions to Late

Cretaceous shallow-marine gastropods from California.

The Veliger 46: 145-161.

Squires, R. L. and L. R. Saul. 2003b. New Late Cretaceous

(Campanian and Maastrichtian) marine gastropods from

California. Journal of Paleontology 77: 50-63.

Squires, R. L. and L. R. Saul. 2004, The pseudomelaniid

gastropod Paosia from the marine Cretaceous of the

Pacific slope of North America and a review of the age and

paleobiogeography of the genus. Journal of Paleontology

78: 484-500.

Tenison-Woods, J. E. 1879. On some new species of

Tasmanian marine shells. Proceedings of the Royal

Society of Tasmania for 1878: 32-40.

Thiele, J. 1929-1935. Handbuch der systematischen Weich-

tierkunde. Gustav Fischer, Jena, pp. 1-1154.

Vedder, J. G. 1977. Preliminary list of Late Cretaceous

mollusks from the Pozo district, San Luis Obispo County,

California. In: D. G. Howell, J. G. Vedder and K.

McDougall (eds.), Cretaceous Geology of the California

Coast Ranges, West of the San Andreas Fault. Pacific

Section Society of Economic Paleontologists and Miner-

alogists, Pacific Coast Paleogeography Field Guide 2: pp.

107-109.

Wenz, W. 1938-1944. Gastropoda. Teil 1: Allgemeiner

Teil und Prosobranchia. In: O. H. Schindewolf (ed.),

Handbuch der Paliozoologie, Band 6. Gebriider Born-

traeger, Berlin, pp. 1-1639. [Reprinted 1960-1961].

Wilson, B. 1994. Australian marine shells. Prosobranch

gastropods, Part 2 (Neogastropoda). Odyssey Publishing,

Kallaroo, Western Australia, 370 pp

Woods, A. J. C. and L. R. Saul. 1986. New Neritidae from

southwestern North America. Journal of Paleontology 60:

636-655.

Woods, H. 1906. The Cretaceous fauna of Pondoland. Annals

of the South African Museum 4: 275-350.

APPENDIX 1

LOCALITIES CITED

Localities are LACMIP, unless otherwise noted. All

quadrangle maps listed below are U. S. Geological

Survey maps.

10786. Near crest of south slope of divide between Basin

Hollow and Clover creeks, at approximately southeast

corner of the northwest 1/4 of section 33, T. 32 N, R. 2

W. Lower Santonian. Redding Formation, Member V

(lower part). Coll.: W. P. Popenoe and D. W. Scharf,

August 8, 1931.

10787. Near crest of north slope of divide between Basin

Hollow and Clover creeks, near northeast corner of

northwest 1/4 of section 33 and not more than 122 m

south of section line, T. 32 N, R. 2 W, Millville

Quadrangle (15 minute, 1953), Shasta County, northern

California. Lower Santonian. Redding Formation, Mem-

Page 148

ber V (lower part). Coll: W. P. Popenoe and D. W.

Scharf, August 8, 1931.

12582. Turritellas and oysters along beach just south of

La Bocana Roja Formation outcrop, approximately 2 km

north of south tip on west side of Punta Baja, Mexico.

Middle upper Campanian. Punta Baja Formation. Coll.:

L. R. Saul, October 12, 1990.

21536. Corona Quadrangle, Santa Ana Mountains,

Orange County, southern California. Lower Campanian.

Ladd Formation, upper Holz Shale Member. Coll.: T.

Bear, 1940.

93639. East bank of Chico Creek in concretions in

massive, greenish-gray sandstone, 373m south and

293 m west of northeast corner of section 23, T. 23 N,

R. 2 E, Paradise Quadrangle (15 minute, 1953), Butte

County, northern California. Lower Campanian. Chico

Formation, Ten Mile Member. Coll.: L. R. and R. B.

Saul, August 20, 1952.

23642. On W bank of Chico Creek, west of big westward

projection of east lava cap and approximately N 27°W of

BM 1770, 122 m south and 632 m west of northeast

comer of section 26, T. 23 N, R. 2 E, Paradise

Quadrangle (15 minute, 1953), Butte County, northern

California. Lower Campanian. Chico Formation, Ten

Mile Member. Coll.: L. R. and R. B. Saul, August 20,

1952.

23643. Concretionary sandstone on west side of Big

Chico Creek, 670 m south and 762 m west of northeast

corner of section 26, T. 23 N, R. 2 E, Paradise

Quadrangle (7.5 minute, 1953), Butte County, northem

California. Lower Campanian. Chico Formation, Ten

Mile Member. Coll.: L. R. Saul and R. B. Saul, August,

1952.

24124. Hard, medium-grained, gray, arkosic sandstone,

158 m north of Jalama Creek, 3.3 km west and 0.6 km

THE NAUTILUS, Vol. 119, No. 4

north of southeast corner of the topographic sheet,

Lompoc Hills Quadrangle (7.5 minute, 1947). Upper

Campanian. Jalama Formation. Coll.: D. Dailey and J. R.

Dorrance, October, 1929.

24125. Hard, fine to medium-grained arkosic sandstone,

2.8 km west and 0.53 km north of southeast corner of

the topographic sheet, Lompoc Hills Quadrangle

(7.5 minute, 1947). Upper Campanian. Jalama Forma-

tion. Coll.: D. Dailey and J. R. Dorrance, October 29,

1929.

24217. Hard sandstone slabs in bed of Clover

Creek, 213m north and 366m west of southeast

comer of section 22, T. 32 N, R. 2 W, Millville

Quadrangle (15 minute, 1953), Shasta County, northern

California. Upper Santonian. Redding Formation,

Member VI. Coll: W. P. Popenoe and D. Dailey,

August 27, 1959.

24337. Along east side of Highway 70, 792 m north and

305 m west of southeast corner of section 36, T. 21 N, R.

3 E, Cherokee Quadrangle (7.5 mintue, 1949), Butte

County, northern California. Lower Campanian. Chico

Formation, Pentz Road member (informal). Coll.: E.

Gohre.

UCMP B-3388. In south-trending arroyo which reaches

coast about 2.8 km north of the tip of Punta Baja.

Marine invertebrates from massive, chert conglomerate

at confluence of this arroyo and northwest-trending

tributary about 122 m upstream from confluence. This

locality is about 488 m northwest of the Punta Baja road.

Middle upper Campanian. Punta Baja Formation. Coll.:

F. H. Kilmer, date unknown.

USGS M-8601. In lens in sandstone on beach 0.5 km

southeast of Bolsa Point, Pigeon Point Quadrangle, San

Mateo County, northern California. Middle Campanian.

Pigeon Point Formation. Coll.: W. P. Elder, 1989?

THE NAUTILUS 119(4):149-152, 2005

Page 149

Daffymitra lindae, a new genus and species of Volutomitridae

(Neogastropoda) from the Bellingshausen Abyssal Plain

Yuri I. Kantor

Severtzov Institute

M. G. Harasewych

Department of Invertebrate Zoology

National Museum of Natural History

Smithsonian Institution

Washington DC 20013-7012 USA Moscow 117071

RUSSIA

Russian Academy of Sciences

Leninski Prospect, 33

ABSTRACT

Daffymitra lindae, new genus, new species, is described from

the Bellingshausen Abyssal Plain off Ellsworthland, Antarctica.

Known only from its shell, this new taxon is included in the

family Volutomitridae, but differentiated from all known living

genera and species on the basis of its inflated shell shape, with

an attenuated anterior and distinctive siphonal canal, as well as

by the shape and disposition of its three recessed, obliquely

oriented columellar plaits. This new taxon represents the first

record of Volutomitridae from abyssal depths. The concholo-

gical similarity of Daffymitra to the Upper Cretaceous genus

Volutomorpha raises the possibility that Daffymitra may be

a surviving descendent of a lineage presumed extinct since the

end of the Cretaceous, and suggests that a reassessment of the

relationships between the various Cretaceous genera assigned

to Volutoderminae and the earliest Volutomitridae.

INTRODUCTION

The Volutomitridae is a small family of rachiglossan

neogastropods characterized by: a fusiform shell (reach-

ing 50 mm) with 2-5 columellar plaits, and a paucispiral,

mammilate protoconch; a radula with distinctive, wish-

bone-shaped rachidian teeth and small, needle-like

lateral teeth that are present in most species; a mid-

oesophagus that is long, convoluted, with a muscular

posterior end; a gland of Leiblein that is only partially

separated from the mid-oesophagus; and a single

accessory salivary gland (Ponder, 1972, 1998; Kantor

and Harasewych, 1992; Bouchet and Kantor, 2004). The

family ranges into the North Pacific, North Atlantic and

the Caribbean, but centers of diversity are concentrated

in the southern hemisphere, particularly in the waters

surrounding Antarctica, Australia, New Zealand, and

especially New Caledonia (Bouchet and Kantor, 2004).

Six genera and 50 species are currently recognized in the

Recent fauna (Bouchet and Kantor, 2004), with a single

genus (Paradmete) containing six species represented in

the Antarctic and Magellanic fauna (Numanami, 1996;

Bouchet and Kantor, 2004). Volutomitrids are known

from the sublittoral zone to depths of 1980 m (based on

dead specimens), but the majority of species are

confined to outer continental shelf and upper continen-

tal slope depths (Bouchet and Kantor, 2004: Appendix;

see Figure 7 herein).

In the course of our studies of Antarctic and sub-

Antarctic neogastropods sampled by the United States

Antarctic Program (USAP) and housed in the collections

of the National Museum of Natural History (USNM), we

encountered a single specimen of a volutomitrid that was

collected on the Bellinghausen Abyssal Plain. This

specimen represents a new genus and species within

the family Volutomitridae, as well as the first record of

the family from abyssal depths. This new genus and new

species is described herein, and comparisons made with

other volutomitrid taxa.

SYSTEMATICS

Class Gastropoda Cuvier, 1797

Order Neogastropoda Wenz, 1938

Family Volutomitridae Gray, 1854

Genus Daffymitra new genus

Type Species: Daffymitra lindae new species, by

original designation.

Description: Shell medium-sized for the family (to

28 mm), with low, conical spire, broadly inflated body

whorl. Protoconch conical, large, 2.6 mm in diameter, of

2 5/8 smooth whorls. Teleoconch thin, of about 3

smoothly rounded whorls, with well demarcated, broad

siphonal canal. Axial sculpture of distinct, very narrow,

broadly spaced ribs that extend from suture to siphonal

canal. Spiral sculpture of narrow, cords, alternating in

prominence. Aperture high (74% of shell length) broadly

ovate. Outer lip thin, smooth. Columella with three

columellar folds (central strongest) and siphonal fold.

Shell white. Periostracum thin, olive brown. Operculum,

anatomy, radula, unknown.

Etymology: This genus is named after Daffy, a tor-

toiseshell longhair cat that belongs to the senior author's

wife Linda.

Page 150

THE NAUTILUS, Vol. 119, No. 4

Figures 1-6. Daffymitra lindae new species, holotype. 1. Apertural view of the shell. 2. Oblique view, shell is rotated slightly to

expose the columellar plaits. 3. Lateral view of the shell. 4. Dorsal view of the shell. 5. Apical and 6. lateral views of the protoconch.

Transition to teleoconch is marked by arrow.

Daffymitra lindae new species

(Figures 1-7)

Description: Shell (Figures 1-4) of moderate size (to

27.9 mm), thin, fragile, with matte surface, inflated,

fusiform, tapering anteriorly, with conical spire. Proto-

conch (Figures 5-6) large, mammilate, 1970 um in

height, diameter increasing from 676 um to 2570 um in

2 5/8 convex whorls. Protoconch-teleoconch transition

distinct (Figures 5, 6, arrow), marked by onset of weak

closely spaced prosocline ribs. Teleoconch of 3 1/8.

strongly convex, ovate whorls with rounded shoulder.

Suture impressed. Axial sculpture of thin, sharply

demarcated, weakly prosocline raised ribs, 42 on last

whorl, 34 on penultimate whorl. Ribs closely spaced on

first teleoconch whorl, becoming more widely spaced on

later whorls, but again closely ‘spaced along final 1/8"

whorl. Spiral sculpture of very low, narrow cords,

subequal in width, alternating in prominence, covering

entire shell surface, about 50 on final whorl, 10 on

penultimate whorl. Aperture large (0.74 shell length),

broadly oval, smooth, deflected fron shell axis by ‘TB’.

Outer lip very thin, weakly reflected, edge forming final

axial rib, with shallow anal sinus at suture. Golumella

weakly sinuate, convex posteriorly, distinctly concave

medially, and again anterior to 3 obliquely oriented,

recessed folds (Figure 2). Central fold most pronounced,

anteriormost fold even more obliquely oriented than

central and posterior fold. Parietal callus, broad, very

thin. Siphonal canal broad, long, well delimited from

M. G. Harasewych and Y. I. Kantor, 2005

Number of Species

(0) 5 10 15 20 25

Depth (m)

Volutomitridae, all species

Paradmete only

Figure 7. Bathymetric distribution of the Recent species of

Volutomitridae, of the Antarctic genus Paradmete (data from

Bouchet and Kantor, 2004: Appendix), and of Daffymitra

lindae. A: continental shelf; B: upper continental slope; C:

lower continental slope; D: continental rise; E: abyssal plain, F:

hadal depths.

aperture. Shell color white. Periostracum thin, olive

brown, covering entire shell. Operculum, radula and

anatomy unknown.

Type Locality: Bellingshausen Abyssal Plain, 61°27’ S,

94°58’—95°22' W, in 4419-4804 m [R/V ELTANIN

cruise 23, sta. 1621, 10 Apr. 1966].

Type Material: Holotype, USNM 1080443, shell

length 27.9 mm, final whorl length 23.4 mm, aperture

length 21.5 mm, shell width, 13.5 mm.

Distribution: Known from the type locality only.

Etymology: This species is named for the senior

author’s wife, Linda Lee Harasewych.

Remarks: Despite the absence of anatomical and

radular data, this new species can be unambiguously

assigned to the family Volutomitridae on the basis of its

distinctive shell shape, sculpture, presence of the

diagnostic paucispiral mammilate protoconch, and weak

columellar folds.

Page 151

Seven genera are currently recognized within the

family Volutomitridae (Cernohorsky, 1970; Bouchet and

Kantor, 2004). The large size, long, broad aperture, and

thin shell of this new species, as well as the presence of

three obliquely oriented and deeply recessed columellar

folds and a siphonal fold preclude its assignment to

either the fossil genus Proximitra Finlay, 1927, or the

Recent genera Conomitra Conrad, 1865; Microvoluta

Angas, 1877. Peculator Iredale, 1924; or Magdalemitra

Kilburn, 1974.

Conchologically, the new species is closer to the

genera Volutomitra H. and A. Adams, 1853 and

Paradmete Strebel, 1908, which are considered to be

closely related (Powell, 1951: 165; Cernohorsky, 1970:

91). The 13 known species of Volutomitra are widely

distributed in the World Ocean, ranging from South

Africa, Southern Australia, New Zealand and to the

Bering Sea in the Pacific, and from Colombia to the

northern part of the Atlantic Ocean. Daffymitra lindae

differs from all known species of Volutomitra in having

a proportionally shorter spire and inflated rather than

fusiform shell, coarse spiral sculpture and sharp, narrow,

broadly spaced axial ribs, as well as columellar folds that

are weak, recessed within the aperture and obliquely

oriented rather than being strong, prominent, and nearly

perpendicular to the waltnanellenr ¢ axis. The only species of

Volutomitra with pronounced axial sculpture, V. erebus

Bayer, 1971, from Colombia, has axial ribs that are

thicker, more rounded, orthocline, and more densely

spaced.

The genus Paradmete, contains six species, all

confined to Antarctic and sub-Antarctic waters. Daffy-

mitra lindae may easily be distinguished from Paradmete

fragillima (Watson, 1882), the type species, as well as

from P. briedensis Numanami, 1996, and P. arnaudi

Numanami, 1996, by its larger size, shorter spire,

inflated rather than narrowly fusiform shell, and

distinctive narrow, prosocline axial ribs, as well as by

its well demarcated siphonal canal. The Magellanic

Paradmete crymochara (Rochebrune and iMelille, 1885)

approaches Daffymitra lindae in size, but differs in its

elongate, fusiform shape, absence of a distinct siphonal

canal, and presence of four columellar folds. The

distinctive Paradmete percarinata Powell, 1951, can be

recognized by its prominent peripheral carina, sharply

dhamitered shell and pronounced columellar folds that

are nearly perpendicular to the columellar axis. Most

similar to Daffymitra lindae is Paradmete curta (Strebel,

1908), which reaches a similar size, has a low spire, and

has axial ribs, which, however, are opisthocline rather

than prosocline. Daffymitra differs in lacking a strong

shoulder and in having an inflated shell shape with an

attenuated anterior and distinctive siphonal canal.

The shell of Daffymitra lindae bears a surprising

resemblance to some members of the Mesozoic genus

Volutomorpha, particularly V. mutabilis Wade, 1926 (see

Wade, 1926: pl. 37, fig. 10, pls. 40, figs. 6, 9; Sohl, 1964:

pl. 39, figs. 1, 2, 6). Volutomorpha was restricted to the

Upper Cretaceous faunas of the Gulf and Atlantic

Page 152

coastal plains (for a review, see Sohl, 1964: 252-254),

and was “the giant of Cretaceous gastropods” (Wade,

1926: 20) with shell lengths extrapolated to exceed

45 cm. Pilsbry and Olsson (1954: 19) included Voluto-

morpha in the Cretaceous subfamily Volutodermatinae,

which they placed in the family Volutidae together with

Volutomitrinae. More recently, Dzhalilov (1977: 93)

proposed a new subfamily Volutomorphinae, also within

Volutidae, while Bouchet et al. (2005: 255) considered

Volutomorphinae a synonym of Volutodermatinae,

which they transferred from Volutidae to the extinct

family Pholidotomidae.

While Daffymitra is easily distinguished from Voluto-

morpha by its far smaller size, thinner shell, absence of

thick axial ribs, and lack of a pronounced shoulder, this

conchological similarity raises the intriguing possibility

that Daffymitra is a “living fossil,” a surviving de-

scendent from a group presumed to have become extinct

at the end of the Cretaceous. Further research is clearly

required to reevaluate the relationships between the

various Cretaceous genera assigned to Volutoderminae

and the earliest Volutomitridae.

DISCUSSION

The family Volutomitridae has a broad geographic range,

but has previously been known only from continental

shelf and continental slope faunas, while the genus

Paradmete has been reported only from shelf and upper

slope depths (Figure 7). The greatest diversities for both

the family and the genus occur at upper continental

slope depths.

Although Daffymitra lindae is represented by a single

empty shell, the fragility of the shell, the presence of

periostracum, and the fact that it was collected below the

aragonite compensation depth indicate that the speci-

men could not have been dead for long, and that the

species inhabits the area in which this specimen was

collected. Thus, this taxon represents the first record of

Volutomitridae from abyssal depths. In a survey of

Antarctic and Magellanic Buccinoidea, Harasewych and

Kantor (2004) found that the abyssal buccinoidean fauna

of the region has no genera in common with the

sublittoral or bathyal faunas, but that credible sister taxa

and likely origins for at least some of the abyssal genera

occur on the adjacent continental slope. Based on shell

morphologies, the genera Volutomitra, Paradmete, and

Daffymitra appear to represent a lineage within

Volutomitridae distinct from the predominantly austral

genera Proximitra, Conomitra, Microvoluta, Peculator,

and Magdalemitra. The genus Paradmete, a member of

the upper slope fauna of Antarctica, is likely the sister

taxon of the abyssal genus Daffymitra.

THE NAUTILUS, Vol. 119, No. 4

ACKNOWLEDGMENTS

This research was supported by a grant from the NSF —

USAP United States Antarctic Program [Contract

Number OPP-9509761]. We are grateful to Bruce

Marshall for bringing to our attention the similarity of

Daffymitra and Volutomorpha.

LITERATURE CITED

Bouchet, P., J. Fryda, B. Hausdorf, W. F. Ponder, A. Valdés and

A. Warén. 2005. Part 2. Working Classification of the

Gastropoda [in] Bouchet, P. and J. P. Rocroi. Classification

and Nomenclator of Gastropod Families. Malacologia 47:

1-397.

Bouchet, P. and Y. I. Kantor. 2004. New Caledonia: the major

centre of biodiversity for volutomitrid mollusks (Mollusca:

Neogastropoda: Volutomitridae). Systematics and Bio-

diversity 1(4): 467-502.

Cernohorsky, W. O. 1970. Systematics of the families Mitridae

and Volutomitridae (Mollusca: Gastropoda). Bulletin of

the Auckland Institute and Museum 8: 1-190.

Dzhalilov, M. R. 1977. Cretaceous gastropods of Southeastern

Central Asia. Donish, Dushanbe, 1-202 pp.

Harasewych, M. G. and Y. I. Kantor. 2004. The Deep-Sea

Buccinoidea (Gastropoda: Neogastropoda) of the Scotia

Sea and Adjacent Abyssal Plains and Trenches. The

Nautilus 118: 142.

Kantor, Y. I. and M. G. Harasewych. 1992. Morphology of the

digestive system of Volutomitra alaskana Dall, 1902

(Gastropoda, Pectinibranchia, Volutomitridae), with notes

on the possible mechanism of feeding. Ruthenica 2:

45-53.

Numanami, H. 1996. Taxonomic study on Antarctic Gastro-

pods collected by Japanese Antarctic Research Expedi-

tions. Memoirs of National Institute of Polar Research

Series E (Biology and Medical Science) 39: 1-244.

Pilsbry, H. A. and A. A. Olsson. 1954. Systems of the Volutidae.

Bulletin of American Paleontology 35(152): 275-306, 4 pls.

Ponder, W. F. 1972. The morphology of some mitriform

gastropods with special reference to their alimentary and

reproductive systems (Neogastropoda). Malacologia 11:

295-342.

Ponder, W. F. 1998. Family Volutomitridae. pp. 842-843. In:

P. L. Beesley, G. J. Ross and A. Wells (eds) Mollusca: The

Southern Synthesis. Fauna of Australia. Vol. 5. CSIRO

Publishing: Melbourne, Part B viii, 565-1234 pp.

Powell, A. W. B. 1951. Antarctic and SubAntarctic Mollusca:

Pelecypoda and Gastropoda. Discovery Reports 26:

47-196, pls. 5-10.

Sohl, N. F. 1964. Neogastropoda, Opisthobranchia and Basom-

matophora from the Ripley, Owl Creek, and Prairie Bluff

Formations. United States Geological Survey Professional

Paper 331-B: i-iv, 153-344, pls. 19-52, tables 1-2.

Wade, B. 1926. The fauna of the Ripley Formation on Coon

Creek, Tennessee. United States Geological Survey Pro-

fessional Paper 137: 1-272, 72 pls., 2 figs.

THE NAUTILUS 119(4):153-156, 2005

Page 153

Exilia alanbeui, a new species from the Neogene of central Chile:

the first record of Exilia (Gastropoda: Ptychatractidae) from

South America

Sven N. Nielsen

Freie Universitat Berlin

Institut fiir geologische Wissenschaften

Fachrichtung Palaontologie

Malteserstrasse 74-100

Haus D, 12249 Berlin

GERMANY

[email protected]

ABSTRACT

A new species of the ptychatractid genus Exilia, E. alanbeui

new species, is described from late Neogene sediments of the

Navidad Formation, central Chile. This new species represents

the first record, fossil or Recent, of the genus Exilia from the

South American continent. Given that Recent species of Exilia

are restricted to bathyal depths, the presence of representa-

tives of the genus in fine siltstones of the Navidad and Ranquil

formations lend support to a previous interpretation of bathyal

depths for these deposits.

INTRODUCTION

The genus Exilia Conrad, 1860, has been placed in the

family Turbinellidae by Maxwell (1988) and in the

subfamily Ptychatractinae within the Turbinellidae by

Kantor et al. (2001). The subfamily Ptychatractinae was

removed from the Turbinellidae and raised to family

level by Riedel (2000), as was suggested previously by

Kantor and Bouchet (1997). Riedel (2000) placed the

Turbinellidae together with other columellar plaits-

bearing groups in his suborder Volutina while the

Ptychatractidae were placed in the suborder Muricina

based mainly on radula type and absence of columellar

plaits. The most recent classification is that presented by

Bouchet and Rocroi (2005), placing Ptychatractidae

within Pseudolivoidea. The genus Exilia has been

revised by Bentson (1940) and more recently by Kantor

et al. (2001). The latter authors synonymized nine

nominal genera, i.e., Mitraefusus Bellardi, 1873, Mesor-

hytis Meek, 1876, Surculina Dall, 1908, Phenacoptygma

Dall, 1918, Palaeorhaphis Stewart, 1927, Zexilia Finlay,

1926, Graphidula Stephenson, 1941, Benthovoluta

Kuroda and Habe, 1950, and Chathamidia Dell, 1956,

containing fossil and Recent species with Exilia and

recognized a total of nine living species. Fossil species

were not formally treated by them but a number of

species were figured for comparison. Previously, the

Recent species of the synonymized genus Benthovoluta

had been reviewed by Cernohorsky (1973) and Har-

asewych (1987). Species of Exilia are known from the

Late Cretaceous (Coniacian) onward and fossil species

are known from all continents except Africa and South

America, “probably a reflection of insufficient studies of

Cretaceous and Tertiary mollusks in these regions”

(Kantor et al., 2001, p. 92). Some Cretaceous through

Miocene species come from shallow-water deposits

while deep-water occurrences are known since the late

Eocene (Kantor et al., 2001). Recent species of Exilia are

restricted to bathyal depths at tropical latitudes and in

the New Zealand region (Kantor et al., 2001).

GEOLOGY OF THE FOSSIL-BEARING LOCALITIES

The Navidad Formation (Darwin, 1846) and its equiva-

lents have recently been re-interpreted as late Miocene

bathyal basin deposits containing displaced intertidal to

outer shelf sediments (Finger et al., 2003; Nielsen et al.,

2003). Several fossiliferous sites have been sampled for

micro- and macrofossils. Of these, two localities from the

Navidad Formation (Figure 1B) and two from the

Ranquil Formation (Figure 1C) yielded specimens of

Exilia described here.

Locality PPP (Figure 1B) is a grey deep-water

siltstone that today forms the intertidal platform at

Punta Perro. Based on planktonic foraminifera this

locality has been dated as late Miocene (Tortonian) by

Tsuchi et al. (1990) and Ibaraki (1992), while new dating

by Finger et al. (2003) suggests an even younger age.

The fauna is of an unusual composition for the Navidad

Formation, containing otherwise unknown species of

Struthiochenopus (Zinsmeister and Griffin, 1995) and

Xenophora (Nielsen and DeVries, 2002). The specimens

coming from Pupuya (Figure 1B) were collected by

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THE NAUTILUS, Vol. 119, No. 4

Isla Coronel

Sta. Maria

Pta. Lavapié

Pta. Perro

Figure 1.

1 @Curanilahue

/ Los Alamos

15S 20 25km

Fossil localities for Exilia alanbeui new species. in central Chile. A. Location of working areas. B. Localities of the

Navidad Formation, Platform Punta Perro (PPP), Pupuya. C. Localities of the Ranquil Formation, Punta El Fraile (FRM), northern

end of Caleta Ranquil (RQT), southwestern end of Caleta Ranquil.

V. Covacevich and D. Frassinetti a little south of the

village Pupuya and come from a grey siltstone for which

no ages are available. However, ihe faunal content (e.g.,

Nigleemn et al., 2004) justifies correlation with PPP. The

sediment at the type locality Punta El Fraile (FRM,

Figure 1C) consists of grey siltstone and underlies

a yellowish sandstone typical for the Ranquil Formation.

The limit between these two lithological units has been

observed to be a sharp, concordant one including some

pebbles at the boundary. The sediments at Caleta

Ranquil (ROT and MIB, Figure 1C) consist of a grey

siltstone similar to that present at Punta El Fraile. The

geological relation with other lithologies present at this

locality, like reddish sandstone with beds of glauconitic

sandstone and coarse light-grey sandstone, are not yet

clear. However, both localities of the Ranquil Formation

are similar to those of the Navidad Formation in

lithology, and faunal content and were also dated as late

Miocene by Finger et al. (2003).

MATERIALS AND METHODS

Specimens described in this study are deposited in the

collections of the Departamento de Paleontologia de

Invertebrados Museo Nacional de Historia Natural,

Santiago de Chile (SGO.PI) and Senckenberg Museum,

Frankfurt, Germany (SMF). Photographs were taken

using a Leicaflex SL2 or Olympus Camedia c730

camera. Images taken with the former camera were

scanned from Ilford FP4 125 black and white 35 mm

negatives using an Acer ScanWit 2720S film scanner. All

images were processed with Adobe Photoshop 7.0.

SYSTEMATIC PALEONTOLOGY

Family Ptychatractidae Stimpson, 1865

Genus Exilia Conrad, 1860

Type Species: — Exilia pergracilis Conrad, 1860, by

monotypy; Midway Group, lower Eocene, Alabama, USA.

Exilia alanbeui new species

(Figures 2-14)

Description: Shell narrow-fusiform, with spire occu-

pying only little more than half of total height.

Protoconch unknown. Teleoconch of about seven

straight to slightly convex whorls, lacking pronounced

shoulder. Axial sculpture consists of rounded, slightly

S. N. Nielsen, 2005

Page 155

Figures 2-14. Exilia alanbeui new species. 2-4. Holotype FRM 019 (height 28.35 mm). 5-6. Paratype FRM 012-3 (height

30.2 mm). 7-8. Paratype FRM 012-2 (height 15.46 mm). 9-10. Paratype FRM 012-1 (height 9.3 mm). 11. Paratype FRM 012-4

(height 18.03 mm). 12. Paratype FRM 012-5 (height 17.62 mm). 13. Paratype FRM 012-6 (height 15.2 mm). 14. PPP 017 (PPP,

height 23.9 mm).

backwards bent, collabral ribs. Penultimate whorl bears

about 16 ribs, which become obsolete on base of whorl.

Spiral sculpture well and equally defined, consisting of

narrow, low, rounded, subequal cords, crossing axial

elements unchanged, separated by interspaces of about

equal width. Fine secondary spiral threads present in

some interspaces. On penultimate whorl 10-14 spiral

cords present, 8—9 further cords present on base of last

whorl and about 30 weaker, closely spaced cords on

siphonal canal. Aperture narrow-elongate. Outer lip

thin. Siphonal canal narrow, long, straight. Inner lip

smooth. Columella smooth and almost straight, with

weak swelling at entrance of siphonal canal.

Etymology: This species is named in honour of Alan

G. Beu (Institute of Geological and Nuclear Sciences,

New Zealand), who always helps when needed.

Type Material: Holotype SGO.PI.6371 (FRM, height

28.35 mm), paratypes SGO.PI.6372 (FRM, one speci-

men, height 10.5 mm), SGO.PI.6373 (FRM, 21 speci-

mens), SMF 327749 (FRM, 19 specimens).

Other Material Examined: SGO.PI.6374 (PPP,

height 23.9 mm), SGO.PI.6375 (PPP, one specimen),

SGO.PI.5303 (Pupuya, four specimens), SGO.PI.6376

(RQT, one specimen, height 9.25 mm), SGO.PI.6377

(MIB, one specimen, height 29.91 mm).

Type Locality: The grey siltstone at Punta El Fraile

(FRM), Arauco, southern central Chile.

Occurrence: PPP, Pupuya, FRM, MIB, RQT; Navi-

dad and Ranquil formations, Late Miocene or Early

Pliocene, central Chile.

Discussion: _—_Exilia alanbeui resembles the type

species, E. pergracilis (see Bentson, 1940, pl. 2,

fig. 25; Kantor et al., 2001, figs. 1 A-D), and

E. lincolnensis Weaver, 1916, but differs from those

North American Eocene species in having a less

convex whorl profile, almost straight instead of sinusoi-

dal axial sculpture and a more angulated whorl base.

It differs from the New Zealand Miocene species

E. nodulifera (Marwick, 1931), E. leachi (Marwick,

1931) and E. wellmanni Maxwell, 1988 (see Maxwell

1988) in having stronger axial sculpture. Most species

of Exilia have more convex whorls and especially

the Recent species have weaker axial sculpture

(see Harasewych, 1987 and Kantor et al., 2001).

The geographically closest species, E. cortezi, has

wider whorls and different axial sculpture and_ is

therefore not considered to be closely related to

E. alanbeui. Exilia alanbeui is known only from

localities consisting of deep water siltstones which

have been dated as probably Messinian in age based

on foraminifera (Finger et al., 2003). Therefore,

this species is younger than the above mentioned

Miocene species from New Zealand. Regarding

the similarity of the two species, E. alanbeui may prove

to be closely related to the New Zealand species

E. wellmanni.

Page 156

THE NAUTILUS, Vol. 119, No. 4

CONCLUSIONS

The occurrence of Exilia alanbeui in central Chile is the

first record of the genus from the South American

continent (except the Recent E. cortezi (Dall, 1908) from

the Galapagos Islands) and therefore closes a consider-

able biogeographic gap. Although shallow water species

of Exilia are known from Miocene deposits of Europe,

the occurrence of Exilia in siltstones of the Navidad and

Ranquil formations is here interpreted to support

microfossil data placing these deposits in a bathyal

environment (Finger et al., 2003).

Exilia alanbeui is another Chilean Miocene gastropod

species that shows many similarities to New Zealand

taxa. Trans-Pacific biogeographic connections between

Chile and New Zealand during the Miocene have

already been shown for some vetigastropods (Nielsen

et al., 2004) and xenophorids (Nielsen and DeVries,

2002).

ACKNOWLEDGMENTS

I thank Klaus Bandel (Universitat Hamburg, Germany),

who helped in numerous ways and provided the

infrastructure to continue this work while I was in

Hamburg. Paulina Vasquez (TU Berlin, Germany) found

the figured specimen from PPP and was a cheerful help

during field work. Daniel Frassinetti (SGO.PI, Chile)

provided access to collections under his care. Tom

DeVries (Burton, USA) improved the language of an

earlier draft. Constructive reviews by Philippe Bouchet

(Muséum National d’Histoire Naturelle Paris, France)

and an anonymous reviewer are gratefully acknowl-

edged. The material has been collected during field work

financed by the Deutsche Forschungsgemeinschaft

grant Ba 675/25 during the years 2000 to 2002.

LITERATURE CITED

Bentson, H. 1940. A systematic study of the fossil gastropod

Exilia. University of California Publications 25: 199-238.

Bouchet, P. and J. P. Rocroi. 2005. Classification and nomen-

clator of gastropod families. Malacologia 47, in press.

Cernohorsky, W. O. 1973. The taxonomy of Benthovoluta

hilgendorfi (von Martens) and allied turbinellid genera

(Mollusca: Volutacea). Records of the Auckland Institute

and Museum 10: 123-131.

Darwin, C. 1846. Geological observations on South America.

Smith, Elder & Co., London, 279 pp:

Finger, K., A. Encinas, S. Nielsen and D. Peterson. 2003.

Microfaunal indications of Late Miocene deep-water

basins off the central coast of Chile. 10° Congreso

Geologico Chileno, Concepcién, Chile. Abstract Volume

CD-ROM. 8 pp.

Harasewych, M. G. 1987. A revision of the genus Benthovoluta

with notes on the evolution of the subfamily Ptychatracti-

nae (Prosobranchia: Turbinellidae). The Nautilus 101:

166-181.

Ibaraki, M. 1992. Planktonic foraminifera from the Navidad

Formation, Chile: their geologic age and paleoceano-

graphic implications, pp. 91-95. In K. Ishizaki and

T. Saito (eds.), Centenary of Japanese Micropaleontology.

Terra Scientific Publishing Company, Tokyo.

Kantor, Y. I. and P. Bouchet. 1997. The Ptychatractinae: an

endemic deep-sea clade of the Turbinellidae? American

Malacological Union, 63% Annual Meeting (Santa Bar-

bara, California), Abstracts: 37-38.

Kantor, Y. I., P. Bouchet and A. Oleinik. 2001. A revision of the

Recent species of Evxilia, formerly Benthovoluta (Gastrop-

oda: Turbinellidae). Ruthenica 11: 81-136.

Maxwell, P. A. 1988. Late Miocene deep-water Mollusca from

the Stillwater Mudstone at Greymouth, Westland, New

Zealand: paleoecology and systematics. NZ Geological

Survey Paleontological Bulletin 55: 120 pp.

Nielsen, S. N. and T. J. DeVries. 2002. Tertiary Xenophoridae

(Gastropoda) of western South America. The Nautilus

116: 71-78.

Nielsen, S. N., T. J. DeVries, A. Encinas, K. L. Finger and D.

Peterson. 2003. Towards an understanding of the age of

the Navidad Formation. 10° Congreso Geoldgico Chileno,

Concepcion, Chile. Abstract Volume CD-ROM. 7 pp.

Nielsen, S. N., D. Frassinetti and K. Bandel. 2004. Miocene

Vetigastropoda and Neritimorpha (Mollusca, Gastropoda)

of Central Chile. Journal of South American Earth

Sciences 17: 73-88.

Riedel, F. 2000. Ursprung und Evolution der “héheren”

Caenogastropoda. Berliner Geowissenschaftliche Abhand-

lungen E32. 240 pp.

Tsuchi, R., T. Shuto, T. Takayama, I. Koizumi, A. Fujiyoshi, R.

Nomura, M. Ibaraki, H. Duque-C, R. Tirado-S, M.

Aldana-A, E. Villavicencio-R and R. Martinez-P. 1990.

Trans-Pacific correlation of Neogene geologic events.

Reports of Andean Studies, Shizuhoka University, Special

vol. 3: 1-7.

Zinsmeister, W. J. and M. Griffin. 1995. Late Cretaceous and

Tertiary aporrhaid gastropods from the southem rim of

the Pacific Ocean. Journal of Paleontology 69: 692-702.

THE NAUTILUS 119(4):157-163, 2005

Page 157

Spawn of Amphissa sp. and Cosmioconcha sp. (Caenogastropoda:

Columbellidae) from the Colombian Caribbean

Patricia Miloslavich

Ana Karinna Carbonini

Departamento de Estudios Ambientales

Universidad Simén Bolivar

P.O. Box 89000

Caracas 1080

VENEZUELA

[email protected]

Bogota D.C.

COLOMBIA

Juan Manuel Diaz

Instituto Alexander von Humboldt

Néstor E. Ardila

Instituto de Investigaciones Marinas

INVEMAR k

Santa Marta, A.A. 1016

COLOMBIA

ABSTRACT

We describe the egg capsules and embryos of Amphissa sp. and

Cosmioconcha sp. collected in the Caribbean Sea off Colombia

between 20-160 m depth. Amphissa sp. had one layer of 24

egg capsules attached to the shell. Cosmioconcha sp. was

completely covered by several layers of egg capsules; only the

outer layer capsules contained embryos, the rest had the

escape aperture open. Egg capsules of both species were

translucent, dome-shaped, with an oval escape aperture at the

center of the dome top. The surface was smooth, with no ridges

nor sutures, and attached to the shell by an oval basal

membrane that was surrounded by a thin, irregular flange. Egg

capsules of Amphissa sp. measured 2 mm in diameter and

contained 6 embryos/capsule at the gastrula stage; egg capsules

of Cosmioconcha sp. measured 1 mm in diameter and

contained 25 embryos/capsule at all stages of development

(eggs, trochophores, and veliger larvae). Gastrulae of Amphissa

sp. measured 750-900 fm in length and development in this

species is probably direct. Uncleaved eggs of Cosmioconcha sp.

measured 150 jum in diameter, the veliger had an operculum,

a transparent shell measuring 188-219 jm in length, and

a small velum; hatching occurs as planktonic larvae. No nurse

eggs were observed in the two species but late cannibalism

among sibling embryos may occur in Cosmioconcha sp.

INTRODUCTION

The family Columbellidae is very diverse in number of

species in the Southern Caribbean. In this region, a total

of 13 genera (Columbella Lamarck, 1799; Rhombinella

Radwin, 1968; Amphissa H. and A. Adams, 1853;

Zafrona Iredale, 1916; Anachis H. and A. Adams,

1853; Nassarina Dall, 1889; Mitrella Risso, 1826;

Cosmioconcha Dall, 1913; Aesopus Gould, 1860; Deci-

pifus Olsson and McGinty, 1958; Mazatlania Dall, 1900;

Strombina Morch, 1852; Nitidella Swainson, 1840, and

Pyrene Réding, 1798) and more than 30 species have

been recorded (cf. Radwin, 1877 a, b; 1978; Costa,

2005); however, columbellid classification, as well as the

identity and interpretation of the characters that support

this classification, is not well resolved (DeMaintenon,

1999). The shells of these species are usually small and

fusiform, within a size range of 5 to 24 mm and the outer

lip usually bears denticles. The egg capsules and

reproduction of several species have been described

worldwide: Thorson (1940) summarized the different

forms of egg capsules found in species from the Iranian

Gulf, Knadeen (1950, 1995) described the egg capsules

and development of species of tropical West Africa and

the Azores, Amio (1957, 1963) described the eggs, larvae

and embryology of Japanese species, DAsaro (1970)

described the egg capsules of Panamanian Pacific

species, Marcus ad Marcus (1962) described the

reproductive biology of several species from Brazil and

Fortunato et al. ( 1998) characterized the reproduction of

Bifurcium bicanaliferum (Sowerby, 1832) from the

Pacific coast of Panama. In the Caribbean region,

Bandel (1974) described the egg capsules of 10 species

of Columbellidae from Santa Marta, Colombia; Pench-

aszadeh et al. (1983) characterized the reproduction of

Mazatlania consentini Philippi, 1836 (as M. aciculata)

(Lamarck, 1822) and Cipriani and Penchaszadeh (1993)

that of Strombina francesae J. Gibson-Smith, 1974, and

Strombina pumilio (Reeve, 1859), all three from

Venezuela. Despite these studies, egg capsule morphol-

ogy and developmental mode is still unknown for most of

the recognized species.

Bandel (1974) distinguished 6 morphological groups

of columbellid egg capsules according to the material he

observed in Colaba and the ere These are: (1)

shallow domes on a broad, irregularly rounded disk of

adhesion only extending on a narrow rim beyond the

capsule walls as in Mitrella ocellata (Gmelin, 1791)

reported as Mitrella argus (d’Orbigny, 1842), (2) cone

shaped capsule, the calls rise from the round adhesion

disk and end in a projecting edge forming a collar

around the escape aperture, which is concave as in

Costoanachis sparsa (Reeve, 1859), (3) cupola shaped

with a suture dividing the capsule in 2 halves, oval base

and the basal membrane extends in an irregular rim

beyond the capsule walls, the escape aperture is

asymmetrically located as in Columbella mercatoria

(Linnaeus, 1758), (4) flask shaped and oval or round at

the base, radial symmetry, the round adhesion disk is

smooth and transparent, the capsule is sculptured with

ridges as in Nitidella nitida (Lamarck, 1822), (5) brick

Page 158

76°0'00"W 74°0'00"W 72°0'00"W

Punta Galinasg- = “85

ro)

)

12°0'00"N

10°0'00"N

~ 10°0'00'N

E-159,-5~/

Puerto® ~~

Escondido

COLOMBIA 025 9

50 100=

Km iS

72°0'00"W

76°0'00"W 74°0'00"W

Figure 1. Map of the collecting localities in the

Colombian Caribbean.

shaped egg capsules, escape aperture is absent as in

Anachis sp. and (6) oval shaped standing on a peduncle

as in Columbella tryngas Pils. A common spawning

feature among these species was that they attached the

egg capsules to hard substrates such as rocks, stems and

algae (Sargassum), aquarium walls and hydroid stalks.

Another spawning strategy found in the family involves

attaching the egg capsules to the shells of conspecific

living adults, such is the case of Mazatlania consentini,

Strombina pumilio, S. francesae and Bifurcium bicana-

liferum (Penchaszadeh et al., 1983; Cipriani and

Penchaszadeh, 1993; Fortunato et al., 1998). There is

no report in the literature for the Columbellid family of

females using the shells of other living species as

a substrate for egg laying.

In this paper, we aesenbe the egg capsules and

embryos of Amphissa sp. (Amphissa “EL, and Adams,

1853) and Cosmioconcha sp. (Cosmioconcha Dall, 1913)

two apparently yet undescribed species from the Colom-

bian Caribbean continental shelf. We also report on the

number of egg capsules spawned by individual females

and some reproductive parameters such as number and

size of embryos contained in the egg capsules.

MATERIALS AND METHODS

One specimen of Amphissa sp. and three specimens of

Cosmioconcha sp. were collected during 2001 by the

Colombian ship B/I Ancon during an expedition to the

Colombian Caribbean shelf (Figure 1). The specimen of

Amphissa sp. was collected at Puerto Escondido, station

E-159 at 158 m in muddy bottom (9°17' N, 76°26’ W).

The three specimens of Cosmioconcha sp. were

collected at Punta Gallinas, station E-85 at 22 m depth

in muddy-sand bottom (12°27' N, 71°41’ W). Samples

were obtained by trawling for 10 minutes with a trawling

net type V measuring 12.7 m in total length and with

a mesh size of 10 mm anmonen Trawl Co, Inc, USA).

THE NAUTILUS, Vol. 119, No. 4

Samples were washed and separated in a 2 mm mesh

seine, observed alive and preserved in ethanol 70%. The

material of both species is deposited at the Museo de

Historia Natural Marina de Colombia (MHNMC),

INVEMAR, catalog numbers INV MOL3746 for Am-

phissa sp. and INV MOL5396 for Cosmioconcha sp.

The following reproductive aspects were studied: (1)

number and size of egg capsules attached to each specimen,

(2) number and size of embryos within each capsule and

(3) observation of the different stages of development.

RESULTS

The specimen of Amphissa sp. (Figure 2) measured

12.5 mm and5 mm in shell length and width respectively;

the protoconch of this species measured 480 [1m in length

and had two whorls. A seemingly related species is

A. acuminata (Smith, 1915) from the southern West

Atlantic. Specimens of Cosmioconcha sp. (Figure 3)

measured between 1] and 12 mm in shell length and

between 4.5 and 5 mm in shell width; the protoconch

measured 570 tim in length and had 3-3'2 whorls. The

taxonomic status of this species has to be confirmed.

A total of 24 egg capsules were attached to the shell of

Amphissa sp. (Figure 4); these were arranged in one

layer and covered most of the shell. The three specimens

of Cosmioconcha sp. were completely covered by egg

capsules in several layers that added 3 mm to the size of

the shell (from 5 to § mm) (Figure 5). The egg capsules

of the internal layers had no embryos and had their

escape apertures open; the closed capsules containing

embryos were limited to the external layer (Table 1).

The egg capsules of Amphissa sp. (Figure 6) were

translucent, dome-shaped, with an oval escape aperture

located on the center of the dome top. The capsule

surface was smooth with no ridges nor sutures. They

were attached to the shell by an oval basal membrane

that was surrounded by a thin, irregular flange. Egg

capsules measured around 2mm in diameter and

contained around 6 gastrulae embryos in each capsule.

The egg capsules of Cosmioconcha sp. (Figure 7) were

very similar to those of Amphissa sp. but smaller,

measuring around 1 mm in diameter and containing

around 25 embryos in each capsule at the three

developmental stages examined: eggs, trochophore, and

veliger larvae (Table 2). No nurse eggs were observed.

The open egg capsules were filled with debris composed

of organic matter and muddy sediment. Between the egg

capsules, within the empty spaces, several specimens of

the genus Turbonilla were found.

All egg capsules of Amphissa sp. contained embryos in

the gastrulae stage in a number of six per capsule. The

gastrulae were yellow, elongated, measuring 750-

900 tm in length and 600-660 pm in width (Table 3).

Each egg capsule of Cosmioconcha sp. contained

embryos in the same stage of development; however,

three different stages were identified in the spawn:

uncleaved eggs, trochophore and veliger larvae (Table 3).

P. Miloslavich et al., 2005

Page 159

Figures 2-5. Shells and attached egg capsules. 2. Adult shell of Amphissa sp. and detail of protoconch. Scale bar = 2 mm, detail =

200 um. 3. Adult shell of Cosmioconcha sp. and detail of protoconch. Scale bar = 2 mm, detail = 100 tm. 4. Adult shell of Amphissa

sp. covered by attached egg capsules. Scale bar = 2 mm.

bar = 2 mm. All photos taken from preserved material.

Table 1. Cosmioconcha sp. Number of layers and summary of

the attached Be capsules for each of the three specimens

collected (ND = not determined).

(ee)

Specimen 1

Number of layers 4 6 4

Total number of attached capsules

Total number of capsules in external layer ND 217 157

Number of open empty capsules 447 623 385

Number of closed empty capsules 3 0 1

Number of closed capsules containing 86 152 108

embryos

Number of open capsules containing 1 0 0

embryos

5. Adult shell of Cosmioconcha sp. covered by attached egg capsules. Scale

Empty embryonic shells were observed in a few capsules

with embryos at the veliger stage. The uncleaved eggs

measured 150 um in dhamneie? and the tr ochophore

measured 165-212 um in length. The veliger had an

operculum and was char avieniae’ by a transparent,

fragile shell measuring 188-219 um in length and 156-

194 um in width. Te: velum was small, measuring 144—

219 um across from one lobe to the other and 31-94 um

in height; the cilia of this velum were very small,

measuring 6-13 Lm.

DISCUSSION

Amphissa sp. has not been previously reported in the

Colombian Caribbean; the sediment at the locality

Pace 160

Figures 6-7. 6. Egg

capsule of Amphissa sp. Scale bar =

500 pm. 7. Egg capsule of Cosmioconcha sp. Scale bar =

500 pm. All photos taken from preserved material.

where the specimen was collected is muddy, suggesting

a broader habitat distribution for the species. The shell

of this species resembles somewhat that of A. acuminata

(Smith, 1915), a southwestern Atlantic species (Costa,

2005), but it is not as slender and the whorls are

more rounded; the teleoconch consists of six rounded

whorls and the sculpture of uniform spiral lirae and

fine axial ribs, forming small rounded nodules at the

THE NAUTILUS, Vol. 119, No. 4

intersections; aperture narrow, suboval; outer lip a lit-

tle thickened exteriorly, with about five slender, short

lirae within. For a detailed description of A. acuminata,

see Simone and Leme (2001) and Costa (2005). The

living species apparently most closely related to Cosmio-

concha sp. is Costoanachis helenae Costa, 1983, whose

distribution range is apparently restricted to the

tropical sector of the Brazilian coast, from Amapa to

Rio de Janeiro, in depths ranging from 8 to 100 m (Rios,

1994; Costa, 2005). We compared the egg capsules of

Cosmioconcha sp. to those of C. helenae collected off

Boipeba, an island on coast of the Brazilian state of

Bahia (deposited at the Museu Nacional do Rio Janeiro,

MNRJ 1029). The specimen measured 11.9 mm in

length and 5.2 mm in width. Several spawning events

had occurred on this shell since there were two layers of

egg capsules, the external with about 30 and the internal

onlin about 15 capsules, a number considerably low-

er than the number of capsules found in our species

of Cosmioconcha (see Table 2). The capsules were

very similar in shape to those of Cosmioconcha sp. but

their size was larger; they measured 1.4 mm in length

and 1.3mm in width, and the aperture measured

0.6 mm in length and 0.5 mm in width (n= 10 capsules

measured).

Cosmioconcha sp. was previously reported from the

Colombian Caribbean as Anachis cf. fraudans Jung,

1969, a very closely related form from the Miocene-

Pliocene of Trinidad (Diaz and Puyana, 1994). It also

resembles C. helenae (Costa, 1983) in shell size and

form, as well as in the number and form of embryonic

whorls. However, the latter has a more slender spire, the

upper half of its last whorl is sculptured with 11-18 axial

ribs and its shell color is pale brown with white spiral

bands (Costa, 2005). The last whorl of Cosmioconcha sp.

lacks axial ribs and the shell is pale yellow under the dark

brown periostracum. Costoanachis helenae has so far

been only recorded from the central Brazilian coast,

from Amapa to Rio de Janeiro (Rios, 1994; Costa, 2005).

The reproductive strategy of spawning or attaching the

egg capsules to the adult shell of conspecifics is

comparable to the spawning of other Caribbean species

found in Venezuela, (Penchaszadeh et al., 1983; Cipriani

and Penchaszadeh, 1993). However, it was not reported

Table 2. Summary of the egg capsule characteristics of Amphissa sp. and Cosmioconcha sp. Values represent mean = SD, numbers

in parenthesis indicate range (n = number of egg capsules measured).

Species

Shell length (mm)

Capsule

length (mm)

Capsule

width (mm)

Capsule

height (mm)

Aperture

length (mm)

’ Aperture

width (mm)

Embryos

per capsule

Amphissa sp. 2.2+0.1 2.0+0.01 0.8+0.2 0.90+0.04 0.74£0.07 5.9+1.2 (4-8)

(12.5) (2.0-2.4) (1.9-2.1) (0.6-1.0) (0.78-0.93) (0.60—0.90) n=l]

n=18 n=18 n=18 n=11 n=11 (Gastrulae)

Cosmioconcha sp. 1.15+0.15 1.01+0.12 0.36+0.06 0.35+0.05 0.27+0.03 25.0+2.9 (19-31)

(11-12) (0.75-1.5) (0.75-1.47) (0.24—0.50) (0.27-0.54) (0.02-0.02) n=23

n=63 n=63 n=62 n=62 n=62 (All stages)

P. Miloslavich et al., 2005

Page 161

Table 3. Size of embryos of Amphissa sp. and Cosmioconcha sp. during intracapsular development. Values represent mean + SD

and are reported in um, numbers in parenthesis indicate range (n = number of embryos measured).

Species Egg Gastrula Trochophore Veliger Protoconch (in adult)

Amphissa sp. = 840+ fees 900) = = 500-600

Cosmioconcha sp. 150+0 — 189+19 (165-213) 200+14 (188-219) 570

n=16 n=4 n=7

for any of the 10 columbellid species studied by Bandel

(1974) in the Santa Marta region, which attached the egg

capsules to other hard substrates such as rocks, Sargassum

plants, stalks of hydroids and, when in captivity, to the

glass walls of the aquarium. There are no records in the

literature of columbellids spawning on the shells of other

species, a strategy that has been observed in the sandy

beach gastropod Olivancillaria deshayesiana Ducros,

1857, from Argentina, which spawns the egg capsules on

the adult shells of Buccinanops monilifer Kiener, 1834,

and Buccinanops duartei Klappenbach, 1961, either

directly on the adult shell or on top of the egg capsules

of these species, which are also attached to the shell of

conspecific individuals (Borzone, 1985). The fact that

several layers of egg capsules are found, as well as different

stages of embryonic development and empty egg capsules,

on Cosmioconcha sp. suggests that several different events

of oviposition have occurred using this specimen as

substrate. Penchaszadeh et al. (1983) reported that

females of Mazatlania consentini (as M. aciculata) attach

their egg capsules to the shells of other individuals of the

population with shells larger than 9 mm, apparently

exclusively males, as shown by histological sections of

the gonads of bearing individuals (Penchaszadeh, 1981).

Fortunato et al. (1998) stated that the egg masses of

Bifurcium bicanaliferum usually have several layers as

well, whereas those of the Venezuelan Caribbean

Strombina pumilio, S. francesae, and M. consentini have

only one layer of egg capsules attached to the shell as we

observed in Amphissa sp. Regarding this remarkable

difference in the number of egg capsules that a single

animal is bearing, we hypothesize that the egg capsules of

species that bear few of them on the shell arranged on

a single layer are easily detachable since they are usually

individually attached, as is the case in M. aciculata. In this

species, up to 20 egg capsules can be observed at the same

time on one single adult shell; these have embryos at

different stages of development, which means that not all

egg capsules are spawned simultaneously. Once the

veliger larvae hatch, the empty capsule usually falls off

the shell, sometimes leaving only the basal membrane

attached to the shell, and new capsules are spawned

(Penchaszadeh, 1981). On the other hand, egg capsules of

species with several layers are more resistant since they are

attached not only to the adult shell by the basal membrane,

but also form a very compact structure of egg capsules

attached between them by lateral membranes (B.

bicanaliferum, Cosmioconcha sp.). As pointed out by

Pechenik (1986), gastropod egg capsules are structurally

and chemically complex, the different layers are composed

in general among and other components by protein and

carbohydrates (Bayne, 1968; Miloslavich, 1996; Rawlings,

1999). Hunt (1966) also reported that since the capsule

walls are extremely resistant to chemical treatment, the

protein and carbohydrate components should be strongly

linked in a glycoprotein complex and thus stabilized by

sclerotization. In this way, the bond between two egg

capsules is much stronger than the bond between an egg

capsule and a shell.

The description of the egg capsule of Amphissa sp. is

similar to the description provided by Bandel (1974) for

Zafrona pulchella (cited as Anachis pulchella) in its

dome shape. However, the dome of Amphissa sp. lacks

the division into lower and upper parts separated from

each other by one or a few concentric rings, it does not

have a suture that continues the axis of the escape

aperture and the membrane is not striped nor wrinkled.

On the contrary, the surface is very smooth and only very

thin imperceptible lines are visible. A comparable

feature among both species is that the number of

embryos per capsule was very similar and that no nurse

eggs nor cannibalism seem to occur.

The egg capsules of Cosmioconcha sp. also fit in the

first category of egg capsule morphology described by

Bandel (1974), a simple dome attached by a rounded

base. It is very similar to the capsule of Amphissa sp. but

half its size in length, width and height. This capsule

shape and general morphology are also comparable to

the two species of the Strombina group from the

Caribbean described by Cipriani and Penchaszadeh

(1993), which have about the same size than the capsules

of Amphissa sp., and to B. canaliferum from the Pacific

coast of Panama described by Fortunato et al. (1998),

which has about the same size than the capsule of

Cosmioconcha sp.

No nurse eggs seem to be present in the two species

studied. In comparison to other columbellid gastropods,

the reproductive strategy of Amphissa sp. is similar to

that of S. francesae and S. pumilio: large egg capsules

(more than 2 mm in length), large eggs (more than

500 [1m) and few embryos (about 4 to 8) that hatch as

large crawling juveniles (larger than 0.9 mm). We did

not observe the hatching stage of Amphissa sp., but the

large size of the gastrula (about 840 pm) indicates that it

Page 162

THE NAUTILUS, Vol. 119, No. 4

comes from a large egg which would most probably lead

to direct development as in the two species of Strombina

studied by Cipriani and Penchaszadeh (1993). On the

other hand, the reproductive strategy of Cosmioconcha

sp. is similar to that of Bifurcium bicanaliferum, Anachis

sparsa, and Cosmioconcha sertulariarum (dOrbigny,

1839): small egg capsules (about 1 mm in length), small

eggs (about 140-200 um) and many embryos (more than

20) that hatch as small veliger larvae (about 300 um)

(Fortunato et al., 1998: Marcus and Marcus, 1962). Late

cannibalism among sibling embryos inferred from the

presence of empty shells inside a few egg capsules of

Cosmioconcha sp. has also been reported in B.

bicanaliferum by Fortunato et al. (1998). Although late

cannibalism might take place in the egg capsules of

Cosmioconcha sp., it is very unlikely that this limited

extraembryonic food source will allow the larva to grow

from 200 tm to 600 um, which is the protoconch size

found in the adult shell. It is most likely that

Cosmioconcha sp. will hatch as a veliger larva spending

some time in the plankton as is also the case of B.

bicanaliferum, which grows from 1.5 whorls to 2.5

whorls in the planktonic period (Fortunato et al., 1998).

In the case of Amphissa sp., we can conclude from the

size of its protoconch that this species must hatch as

a crawling juvenile since the gastrula stage is already as

large as the protoconch, a situation similar to what

happens with the two Strombina species from the

Venezuelan Caribbean studied by Cipriani and Pench-

aszadeh (1993).

We hope that in a near future, and thanks to the

collection efforts carried out by INVEMAR in the

southern Caribbean, more specimens will be collected.

This will not only allow for the proper description of

these species through use of traditional characters such

as those from adult shell, anatomy, and radular

morphology, but may also help increase the number of

observations on egg capsules and the development of

other embryonic stages.

ACKNOWLEDGMENTS

This work was partially supported by a Decanato de

Investigacion y Desarrollo, Universidad Siméon Bolivar

grant to the Grupo de Ciencias Marinas, by a FONACIT

(S1-2001-000764) grant to the first author. Material

collecting was possible through the “Macrofauna I and

II” projects funded by COLCIENCIAS and INVEMAR.

We are especially indebted to Paulo Marcio Costa,

Departamento de Invertebrados, Malacologia, Museu

Nacional, Rio de Janeiro, and Marta DeMaintenon,

University of Hawaii at Hilo for helping with the

problematic taxonomic status of our material. Paulo

Marcio Costa also provided us with one reproductive

specimen of Costoananchis helenae from Brazil for

comparative purposes. We also wish to thank Pablo

Penchaszadeh, Facultad de Ciencias Exactas y Natur-

ales, Universidad de Buenos Aires, for our fruitful

discussions in the subject and José L. Garcia Rond6n,

Artis Publishers, for his invaluable help with the

photographs.

LITERATURE CITED

Amio, M. 1957. Studies on the eggs and larvae of marine

gastropods I. Journal of the Shimonoseki College of

Fisheries 7: 107-127.

Amio, M. 1963. A comparative embryology of marine

gastropods, with ecological considerations. Journal of the

Shimonoseki College of Fisheries 12: 231-357.

Bandel, K. 1974. Spawning and development of some

Columbellidae from the Caribbean Sea of Colombia.

The Veliger 16: 271-282.

Bayne, C. J. 1968. Histochemical studies of the egg capsules of

eight gastropod molluscs. Proceedings of the Malacologi-

cal Society of London 38: 199-212.

Borzone, C. A. 1995. Ovicapsulas de Prosobranquios (Mol-

lusca: Gastropoda) de una playa arenosa expuesta del sur

del Brasil. Iheringia, Série Zoologia 79: 47-58.

Cipriani, R. and P. Penchaszadeh. 1993. How does Strombina

reproduce? Evidence from two Venezuela species (Proso-

branchia: Columbellidae). The Veliger 36: 178-184.

Costa, F. H. A. 1983. Anachis helenae, nova espécie de

Columbellidae do litoral brasileiro (Mollusca: Gastro-

poda). Anais da Sociedade Nordestina de Zoologia 1:

95-99.

Costa, P. M. S. 2005. Estudo taxonémico dos representantes da

familia Columbellidae Swainson, 1840 (Mollusca, Caeno-

gastropoda) da costa brasileira. Doctoral Dissertation

(CD-ROM), Universidade Federal do Rio de Janeiro,

Museu Nacional, Rio de Janeiro, 291 pp.

DeMaintenon, M. 1999. Phylogenetic analysis of the Colum-

bellidae (Mollusca: Neogastropoda) and the evolution of

herbivory from carnivory. Invertebrate Biology 118:

258-288.

Diaz, J. M. and M. Puyana. 1994. Moluscos del Caribe

Colombiano. Un Catélogo Ilustrado. Colciencias-Funda-

cién Natura-INVEMAR, 291 pp.

Fortunato, H., P. E. Penchaszadeh and P. Miloslavich. 1998.

Observations on the reproduction of Bifurcium bicanali-

ferum (Sowerby, 1832) (Gastropoda: Columbellidae:

Strombina group) from the Pacific Coast of Panama.

The Veliger 41: 208-211.

Hunt, S. 1966. Carbohydrate and amino-acid composition of

the egg capsule of the whelk Buccinum undatum L.

Nature 210: 436-437.

Knudsen, J. 1950. Egg capsules and development of some

marine prosobranchs from Tropical West Africa. Atlantide

Report 1: 85-130.

Knudsen, J. 1995. Observations on reproductive strategy and

zoogeography of some marine prosobranch gastropods

(Mollusca) from the Azores. Agoreana, Supplement:

135-158.

Marcus, E. and E. Marcus. 1962. Studies on Columbellidae.

Boletim da Faculdade de Filosofia, Ciéncias e Letras, Sao

Paulo, 24: 335-384.

Miloslavich, P. 1996. Biochemical composition of prosobranch

egg capsules. Journal of Molluscan Studies 62: 133-135.

Pechenik, J. A. 1986. The encapsulation of eggs and embryos

by molluses: an overview. American Malacological Bulle-

tin 4: 165-172.

P. Miloslavich et al., 2005

Page 163

Penchaszadeh, P. E. 1981. Estudios sobre modalidades

reproductivas de gasterépodos prosobranquios del Caribe

Sur. Trabajo de Ascenso a Profesor Titular de la

Universidad Simon Bolivar, Caracas, 101 pp. (Unpub-

lished work.)

Penchaszadeh, P. E., G. De Mahieu, V. Farache and M. E.

Lera. . Ecology of the sandy beach gastropod Mazatlania

aciculata in Quizandal (Carabobo, Venezuela). In: McLa-

chlan, A. and T. Erasmus (eds.). Sandy beaches as

ecosystems. Proceedings of the First International Sym-

posium on Sandy Beaches, South Africa, Junk Publishers,

pp. 655-660.

Radwin, G. E. 1977a. The family Columbellidae in the Western

Atlantic. The Veliger 19: 403-417.

Radwin, G. E. 1977b. The family Columbellidae in the

Westem Atlantic. Part Ila. The Pyreninae. The Veliger

20: 119-133.

Erratum

Radwin, G. E. 1978. The family Columbellidae in the Western

Atlantic. Part IIb. The Pyreninae (continued). The Veliger

20: 328-344.

Rawlings, T. A. 1999. Adaptations to physical stresses in the

intertidal zone: The egg capsules of neogastropod

molluscs. American Zoologist 39: 230-243.

Rios, E. C. 1994. Seashells of Brazil. 2"? edition. Universidade

do Rio Grande, Rio Grande, 368 pp. 113 pls.

Simone, L. R. L. and J. L. M. Leme. 2001. Anatomia

comparativa e sistematica de Amphissa acuminata e

Amphissa cancellata (Gastropoda, Caenogastropoda, Co-

lumbellidae) da costa sudeste do Brasil. Cadernos do

Centro Universitario Sio Camilo, Sao Paulo, 7(2):

105-114.

Thorson, G. 1940. Studies on the egg masses and larval

development of gastropods from the Iranian Gulf. Danish

Scientific Investigations in Iran. Copenhagen, 2: 159-235.

Due to an editorial lapse in the latest article by Cristian Ituarte (2005), please substitute the last sentence in the

Etymology section (page 97, right-hand column, lines 24, 25) as follows:

Replace ~..., which underwent a major cultural change during the past 10,000 years.”

For: “..., which has been a major pathway for cultural exchange during the past 10,000 years.”

LITERATURE CITED

Ituarte, C. 2005. The Sphaeriidae (Bivalvia) from northwesterm Argentina including three new species of Pisidium. The Nautilus 119:

93-104.

THE NAUTILUS 119(4):164-168, 2005

Page 164

Pterorytis pacanana new species (Gastropoda: Muricidae):

circumstantial evidence for late Pliocene El Nino events in

southern Peru

Thomas J. DeVries"

Burke Museum of Natural History and Culture

University of Washington

Seattle, WA 98195 USA

ABSTRACT

Pterorytis pacanana new species was discovered in upper

Pliocene bioclastic sandstone near Chala, southern Peru.

Although it was found farther south than other Pterorytis

species from the eastern equatorial Pacific Ocean, P. pacanana

most resembles P. roxaneae Petuch, 1994, from the Pliocene

Pinecrest beds of Florida. Its presence among endemic

late Pliocene cool-water mollusks from high-energy shoreface

paleoenvironments is thermally anomalous in the same sense

as the rare occurrence of other species from northern Peru

and Ecuador in upper Pliocene and Pleistocene strata

from southern Peru. By analogy with modern thermally

anomalous mollusks that appear episodically off the coast of

southern Peru and Chile, the Pliocene and Pleistocene

examples are circumstantial evidence for the occasional south-

ward incursion of warm equatorial waters during former El

Nino events.

INTRODUCTION

Pliocene deposits in southern Peru are noted for a variety

of muricid gastropods, most belonging to genera still

represented by extant species (e.g., Acanthina Fischer

von Waldheim, 1807; Chorus Gray, 1847; Concholepas

Lamarck, 1801; Crassilabrum Jousseaume, 1880; Mur-

egina Vermeij, 1998; Stramonita Schumacher, 1817;

Xanthochorus Fischer, 1884), but some from genera

entirely or locally extinct (e.g., Herminespina DeVries

and Vermeij, 1997; Trophon, Montfort, 1810) (DeVries,

1995, 1997, 2000, 2003, in press a, in press b; DeVries

and Vermeij, 1997). Specimens of most Pliocene

muricids are not exceedingly difficult to find.

A well-preserved muricid from Pliocene bioclastic

sandstone south of Chala is, literally, the rare exception.

None have been found other than a single shell plucked

from a roadcut of the Panamerican Highway that

overlooks Playa Huacllaco. The lamellar fimbriate

varices of the Huacllaco specimen invite comparison

with ocenebrines from Ecuador, the southeastern

"Mailing address: P.O. Box 13061, Burton, WA 98013 USA

United States, and the northwestern Pacific Ocean. Its

lone appearance in a collection of endemic cool-water

muricids is another example of a thermally anomalous

molluscan species encountered in upper Pliocene and

Pleistocene beds of southern Peru (Muizon and

DeVries, 1985; DeVries, 1986; Ortlieb et al., 1990). A

reasonable hypothesis is that these equatorial species

were introduced to higher austral latitudes by warm-

water incursions during the Pliocene and Pleistocene

comparable to modern incursions that carry Panamic

molluscan larvae poleward during El Nifio events

(DeVries, 1988; Arntz and Tarazona, 1990; Paredes et

al., 1998).

GEOLOGY

Bioclastic sandstone and gravel and balanid coquina

overlie igneous basement in roadcuts along a series of

sweeping curves in the Panamerican Highway where it

descends towards the beach at Playa Huacllaco

(Figure 1). The sediments represent high-energy fore-

shore and intertidal paleoenvironments that once

flanked steep cliffs. The section, previously published

Figure 1. Type locality of Pterorytis pacanana new species

(DV 1628).

T. J. DeVries, 2005

Page 165

Figures 2-8. Pterorytis species. 2, 3, 5. Pterorytis pacanana new species. Upper Pliocene. Holotype, UWBM 97772, length =

30.9 mm, width = 19.6 mm. 2. Apertural view. 3. Abapertural view. 5. Oblique view of spire. 4, 6-8. Pterorytis roxaneae Petuch,

1994. Upper Pliocene. Holotype, Florida Museum of Natural History, Gainesville, UF 66254, length = 33.3 mm, width = 21.5 mm.

4. Lateral view. 6. Apertural view. 7. Abapertural view. 8. Oblique view of spire.

by DeVries (2003), consists of four stratigraphic units.

The ocenebrine specimen was found at the base of Unit

IIL, just below strata where rounded clasts of blackened

andesite first appear in great numbers and below beds

where specimens of Concholepas and Acanthina acquire

their modern form (DeVries, 2000, 2003).

The age of the Huacllaco beds is bracketed by basal

beds with specimens of Concholepas nodosa Hupé,

1854, Acanthina triangularis DeVries, 2003, and Her-

minespina mirabilis (Moricke, 1896), which collectively

indicate an early late Pliocene age (DeVries and

Frassinetti, 2003), and the uppermost and oldest of

several marine terraces, whose elevation and largely

extant taxa suggest a latest Pliocene age (Muizon and

DeVries, 1985).

MATERIALS AND METHODS

The specimen from Peru described in this study was

found by the author. Dimensions affected by breakage

are enclosed by parentheses. The holotype is deposited

at the University of Washington’s Burke Museum of

Natural History and Culture in Seattle, Washington

(UWBM).

SYSTEMATICS

Family Muricidae Rafinesque, 1815

Subfamily Ocenebrinae Cossmann, 1903

Genus Pterorytis Conrad, 1862

Subgenus Pterorytis sensu stricto

Type Species: — Murex umbrifer Conrad, 1832, by

monotypy.

Pterorytis pacanana new species

(Figures 2, 3, 5)

Diagnosis: Shell small; texture waxy, shell thin. Five

lamellar, fimbriate varices; intervarical nodes absent.

Three spiral cords; shoulder spiral cord strong. Sutural

platform horizontal. Labral tooth absent.

Description: Shell 30.9 mm long (first teleoconch

whorls missing), quadrate in profile, very thin, with

waxy texture. Spire estimated to be about 40 percent of

shell length. Siphonal canal about 20-25 percent of shell

length. Protoconch and earliest teleoconch whorls

missing. Upper sides of whorls planar, vertical; base of

body whorl sharply constricted. Shoulder orthogonally

Page 166

angulate, sutures deeply impressed; sutural platform

horizontal to slightly concave. Three last whorls with five

lamellar varices extending from fasciolar ridge to suture,

each varix joining across suture with varix of previous

whorl. Lamellae broader basally, narrower adapically,

extended adapically at shoulder but not spinose;

fimbriate on adapertural face; erect or recurved

adaperturally except apertural lamellae, which is weakly

recurved abaperturally. Intervarical nodes absent. Spiral

sculpture of prominent but ill-defined rounded primary

cord at shoulder angulation and two additional broad low

spiral cords anteriorly, almost obsolete. Additional broad

low secondary cords barely visible adjacent to adaper-

tural face of varices. Lamellar fimbria slightly extended

at intersection with primary spiral cords. Aperture with

inverted tear-drop shape. Parietal rib, anal sulcus absent;

parietal area unexcavated. Columella smooth, inner lip

very weakly concave, adherent anteriorly. Outer lip

without dentition on inner edge. Labral tooth absent.

Siphonal canal open, slightly recurved to right. Siphonal

fasciole strongly arched, without rostrae. Pseudo-umbi-

licus narrow, extending adapically beyond siphonal

canal.

Holotype: UWBM 97772, DV 1628-5, lower Upper

Pliocene, length (30.9) mm, width 19.6 mm.

Type Locality: Roadcut along the Panamerican High-

way, 10 km south of Chala, on a winding descent from

a 200 m elevation terrace towards Playa Huacllaco,

locality DV 1628, north side of first sweeping outside

curve from base of outcrop, south side of road, near

outcrop of igneous basement rock (Figure 1), 15°52’ S,

74°10’ W (Chala 1:100,000 quadrangle).

Occurrence: Upper Pliocene, southern Peru.

Etymology: ‘Paca, Quechua for ‘high, and ‘nan,

Quechua for ‘path, referring to the horizontal to

concave sutural platform bordered above and below by

vertical walls of the whorls.

Remarks: Pterorytis or Ocinebrellus Jousseaume,

1880, is the most appropriate genus in which to place

the Huacllaco specimen, which has five varices on at

least the last three whorls, consistent with the three to

nine varices observed on fossil Atlantic species of

Pterorytis (Emerson, 1959; Vermeij, 2001), the four to

five varices on the modern eastern Pacific Pterorytis

hamatus (Hinds, 1844) (Emerson, 1985), and three to

twelve varices on specimens of Ocinebrellus (Amano and

Vermeij, 1998a). The ocenebrine genera Ceratostoma

Herrmannsen, 1846, Pteropurpura Jousseaume, 1880,

and Microrhytis Emerson, 1959, in contrast, have three

prominent lamellar varices on all or at least the last two

whorls (Vermeij and Vokes, 1997; Amano and Vermeij,

1998a, 1998b). Specimens of Ceratostoma, Pteropur-

pura, Microrhytis, and some Ocenebra Gray, 1847, also

have intervarical nodes, which are absent on the

Huacllaco specimen and specimens of Pterorytis and

Ocinebrellus.

THE NAUTILUS, Vol. 119, No. 4

The absence of a closed siphonal canal does not argue

against assignment of the single Huacllaco specimen to

Pterorytis or Ocinebrellus, since juvenile and some adult

specimens of the two genera may have an open siphonal

canal. The Peruvian specimen, itself partly abraded,

might be a juvenile specimen or have a broken siphonal

canal.

Distinguishing Pterorytis from Ocinebrellus for place-

ment of the Huacllaco specimen is problematic.

Ocinebrellus has four primary spiral cords on the swollen

portion of the body whorl, posterior to the labral tooth, if

present (Vermeij and Vokes, 1997; Amano and Vermeij,

1998a). Pterorytis and the Huacllaco specimen have

three primary spiral cords, with an additional weak

primary spiral between the shoulder and suture. On

smoother specimens of both genera the number of

primary spiral cords can be difficult to enumerate, and

on strongly sculptured specimens the distinction be-

tween primary and secondary spiral cords is unclear if

the ontogeny is unknown.

Other features are equally unsatisfactory for distin-

guishing the two genera. The reflection of lamellar

varices is not reliable, since the lamellae may be

adaperturally reflected, or not, in Ocinebrellus (Amano

and Vermeij, 1998a), abaperturally reflected or erect in

Pterorytis (Vermeij and Vokes, 1997), and erect or

reflected in either direction on the Huacllaco specimen.

The angularity of the shoulder also fails as a distinguish-

ing character. Most specimens of Ocinebrellus have an

angulate shoulder and a horizontal sutural platform (as is

the case for the Huacllaco specimen), but some have

rounded shoulders. Most specimens of Pterorytis have

broad, planar, steeply sloping sutural platforms, but

some have narrower, less inclined sutural platforms

more like those of typical Ocinebrellus and the

Huacllaco specimen.

The presence or absence of a labral tooth is not

diagnostic. Some ocenebrines acquire a labral tooth only

in adulthood; the Huacllaco specimen might be a juve-

nile. Species of Pterorytis that normally have a tooth (P.

umbrifer Conrad, 1832; P. fluviana Dall, 1890) have

specimens lacking a tooth. Specimens of P. roxaneae

Petuch, 1994, never have a labral tooth (Vermeij and

Vokes, 1997). Species of Ocinebrellus (sensu Amano and

Vermeij, 1998a) that normally lack a labral tooth have

specimens with a tooth. (In the more restrictive

classification of Houart and Sirenko (2003), species of

Ocinebrellus do not have a labral tooth.)

Taking into account this taxonomic ambiguity, the

Huacllaco specimen is assigned to Pterorytis. Species of

Pterorytis (P. umbrifer, P. fluviana, P. roxaneae) typically

have smooth-shelled variants such as the specimen from

Huacllaco. Most specimens of Pterorytis, like the

Huacllaco specimen, feature a prominent primary spiral

cord at the shoulder, but nonetheless usually lack spines

or angulations typically seen in specimens of Ocineb-

rellus.

Pliocene Ecuadorian specimens assigned to Ocineb-

rellus by Vokes (1988) are smooth, have a prominent

T. J. DeVries, 2005

shoulder spiral bordering an inclined sutural platform,

and have varices that are only weakly angulate; they may

be referred to Pierorytis ecuadoria (Olsson, 1964). They

have a more fusiform profile and greater number of

varices than the Huacllaco specimen.

The specimen of Pterorytis pacanana most closely

resembles specimens of the Pliocene Pterorytis (Pteror-

ytis) roxaneae Petuch, 1994, (Figures 4, 6-8) from

the Pliocene Pinecrest beds of Florida (Vermeij and

Vokes, 1997). Specimens of both species are thin with

a waxy texture, have reduced spiral sculpture, and lack

a labral tooth. Pinecrest specimens differ from the

Huacllaco specimen in having four varices, not five,

varical lamellae that are broader adapically, not abapi-

cally, and a weak peripheral spiral cord bordering

a sloping sutural platform, rather than the strongly

defined horizontal sutural platform of P. pacanana.

Contrary to Vermeij and Vokes (1997), the holotype of P.

roxaneae is neither excessively worm nor lacking the

labral varix.

DISCUSSION

Pterorytis pacanana is the third or fourth species of

Pterorytis recognized in the eastern Pacific Ocean, after

P. ecuadoria (Pliocene, Ecuador), P. hamatus (Recent,

northern Peru [Alamo and Valdivieso, 1997], a species

with a protoconch unlike that of Ocinebrellus or any

other ocenebrine [R. Houart, personal communication,

2005]), and an unnamed Recent ocenebrine from

northern Peru (Radwin and D’Attilio, 1976). These taxa

define an eastern tropical Pacific complement to a clade

of Pterorytis species from the southeastern United States

whose oldest members date to the late Miocene.

Ocinebrellus, which may be endemic to the northwest-

em Pacific (Amano and Vermeij, 1998a; Houart and

Sirenko, 2003), extends back to the Early or Middle

Miocene. Ocinebrellus seems morphologically more

similar to Pterorytis than other ocenebrine clades, but

the trail of fossil species that might lead from Japanese

Ocinebrellus to Panama and the Caribbean and beyond

to Peruvian and Floridian Pterorytis has yet to be

discovered (Amano and Vermeij, 1998a).

The specimen of Pterorytis pacanana from 16°S is

remarkable for its singular occurrence and equatorial

affinity. Associated taxa (Table 1) are entirely endemic or

cool-water species that became prevalent after a pro-

vincial mid-Pliocene extinction that coincided with

a global cooling event (Dowsett et al., 1996; DeVries,

2001). The rare appearance of Pterorytis in southern

Peru resembles that of a mangrove bivalve, Anadara cf.

A. grandis (Broderip and Sowerby, 1829), whose speci-

mens are found in small numbers in uppermost Pliocene

beds at 15°30’ S with cool-water species (Muizon and

DeVries, 1985; DeVries, 1986). In a more recent

example, specimens of Chione broggi (Pilsbry and

Olsson, 1943) and Cerithium stercusmuscarum Valenci-

ennes, 1833, both living today only as far south as the

Page 167

Table 1. Molluscan species associated with Pterorytis paca-

nana, new species, which was found at the base of Unit III of

the Upper Pliocene beds above Playa Huacllaco, Peru.

(* = Extinct).

Unit III+IV (younger)

Acanthina unicornis (Bruguiere, 1789)

Cancellaria buccinoides Sowerby, 1832

Choromytilus chorus (Molina, 1782)

Chorus grandis (Philippi, 1887) / C. giganteus (Lesson,

1846)

Concholepas camerata DeVries, 2000

Concholepas concholepas (Bruguiére, 1789)

Crassilabrum crassilabrum (Sowerby, 1834)

Crepidula dilatata (Lamarck, 1822)

Eurhomalea lenticularis (Sowerby, 1835)

Glycymeris ovata (Broderip, 1843)

Mesodesma donacium (Lamarck, 1818)

Mulinia edulis (King, 1831)

Oliva peruviana Lamarck, 1811

Piscoacritia new species

Prisogaster niger (Wood, 1828)

Sinum cymba (Menke, 1828)

Xanthochorus cassidiformis (Blainville, 1832)

Unit I+II (older)

* Acanthina triangularis DeVries, 2003

Acmaeids

* Chlamys cf. C. vidali (Philippi, 1887)

Choromytilus chorus (Molina, 1782)

zo Chorus grandis (Philippi, 1887)

Concholepas camerata DeVries, 2000

Es Concholepas nodosa (Mo6ricke, 1896)

Fissurella spp.

Herminespina mirabilis (Méricke, 1896)

Lithophaga sp.

x Piscoacritia collapsa DeVries and Hess, 2004

Stramonita new species

Tegula (Chlorostoma) new species

Xanthochorus buxeus (Broderip, 1833)

Xanthochorus new species

Sechura coastline of northern Peru (5°S) (Alamo and

Valdivieso, 1997), are occasionally found in upper

Pleistocene terrace deposits near San Juan de Marcona

(15°20'S), Sacaco (15°30’S), and Ilo (17°40'S) (DeVries,

1986, 1988: Ortlieb et al., 1990). Ortlieb et al. (1990)

proposed that the late Pleistocene thermally anomalous

species were introduced southward from equatorial

latitudes during El Nifio events. Several such immigra-

tions of equatorial mollusks have been documented

during modern El Nifios events (Artz and Tarazona,

1990; Paredes et al., 1998). The rare Pliocene equatorial

species in southern Peru were probably introduced in

the same manner.

ACKNOWLEDGMENTS

I would like to thank Brandur Karlsson of Reykjavik,

Iceland, for his assistance in the field and Greg Herbert

(University of South Florida) for helpful discussions on

Page 168

ocenebrine taxonomy. G. J. Vermeij and R. Houart

provided helpful critiques in their reviews of the

manuscript.

LITERATURE CITED

Alamo, V. and V. Valdivieso. 1997. Lista sistematica de

moluscos marinos del Pert. Instituto del Mar del Pert,

Callao, 183 pp.

Amano, K. and G. J. Vermeij. 1998a. Taxonomy and evolution

of the genus Ocinebrellus (Gastropoda: Muricidae) in

Japan. Paleontological Research 2: 199-212.

Amano, K. and G. J. Vermeij. 1998b. Origin and biogeographic

history of Ceratostoma (Gastropoda: Muricidae). Venus

57; 209-223.

Amitz, W. A. and J. Tarazona. 1990. Effects of El Nifio 1982-

1983 on benthos, fish, and fisheries off the South

American Pacific coast. In: P. W. Glynn (ed.) Global

ecological consequences of the 1982-1983 El Nifio-

Southern Oscillation. Elsevier: Amsterdam, pp. 323-360.

DeVries, T. J. 1986. The geology and paleontology of tablazos

in northwest Peru. Doctoral dissertation, The Ohio State

University: Columbus, 964 pp.

DeVries, T. J. 1988. A review of geological evidence for ancient

El Nifio activity in Peru. Journal of Geophysical Research

(Oceans) 92(C13): 14,471-14,479.

DeVries, T. J. 1995. Concholepas Lamarck, 1801 (Neogastro-

poda: Muricoidea): A Neogene genus native to South

America. The Veliger 38: 284-297.

DeVries, T. J. 1997. A review of the genus Chorus Gray, 1847

(Gastropoda: Muricidae) from western South America.

Tulane Studies in Geology and Paleontology 30: 125-147.

DeVries, T. J. 2000. Two new Neogene species and the

evolution of labral teeth in Concholepas Lamarck, 1801

(Neogastropoda: Muricoidea). The Veliger 43: 43-50.

DeVries, T. J. 2001. Contrasting patterns of Pliocene and

Pleistocene extinctions of marine mollusks in western

North and South America. Geological Society of America,

Abstracts with Programs 33(3): A-35.

DeVries, T. J. 2003. Acanthina Fischer von Waldheim, 1807

(Gastropoda: Muricidae), an ocenebrine genus endemic to

South America. The Veliger 46; 332-350.

DeVries, T. J. a. (In press). The Late Cenozoic history of

Xanthochorus Fischer, 1884 (Gastropoda: Muricidae) in

western South America. The Veliger.

DeVries, T. J. b. (In press). Late Cenozoic Muricidae from

Peru: Seven new species and a biogeographic summary.

The Veliger.

THE NAUTILUS, Vol. 119, No. 4

DeVries, T. J. and C. D. Frassinetti. 2003. Range extensions

and biogeographic implications of Chilean Neogene

mollusks found in Peru. Boletin del Museo de Historia

Natural, Chile 52: 141-157.

DeVries, T. J. and G. J. Vermeij. 1997. Herminespina: New

genus of Neogene muricid gastropod from Peru and

Chile. Journal of Paleontology 71: 610-615.

Dowsett, H., J. Barron and R. Poore. 1996. Middle Pliocene

sea surface temperatures: a global reconstruction. Marine

Micropaleontology 27: 13-25.

Emerson, W. K. 1959. The gastropod genus Pterorytis.

American Museum Novitates 1974: 1-8.

Emerson, W. K. 1985. Murex hamatus Hinds, 1844, a living

West American species assigned to the Neogene paciphile

genus, Pterorytis Conrad (Gastropoda: Muricidae). The

Nautilus 99: 14-17.

Houart, R. and B. I. Sirenko. 2003. Review of the Recent

species of Ocenebra Gray, 1847 and Ocinebrellus

Jousseaume, 1880 in the northwest Pacific. Ruthenica

13: 53-74.

Muizon, C. de and T. J. DeVries. 1985. Geology and

paleontology of the Pisco Formation in the area of Sacaco,

Peru. Geologische Rundschau 74(3): 547-563.

Ortlieb, L., T. DeVries and A. Diaz. 1990. Ocurrencia de

Chione broggi (Pilsbry and Olsson, 1943) (Pelecypoda) en

depositos litorales Cuaternarios del sur del Pert: Im-

plicaciones paleoceanograficas. Boletin de la Sociedad

Geolégica del Pert 81: 127-134.

Paredes, C., J. Tarazona, E. Canahuire, L. Romero, O. Cormejo

and F. Cardozo. 1998. Presencia de moluscos tropicales

de la provincia panameiia en la costa central del Pert y su

relacién con los eventos “El Nifio”. Revista Peruana De

Biologia (Universidad Nacional Mayor De San Marcos)

5(2): 123-128.

Petuch, E. J. 1994. Atlas of Florida Fossil Shells. Chicago

Spectrum Press, Evanston, 394 pp.

Radwin, G. E. and A. D’Attilio. 1976. Murex Shells of the

World. Stanford University Press, Stanford, 284 pp.

Vermeij, G. J. 2001. Innovation and evolution at the edge:

origins and fates of gastropods with a labral tooth.

Biological Journal of the Linnean Society 72: 461-508.

Vermeij, G. J. and E. H. Vokes. 1997. Cenozoic Muricidae of

the western Atlantic region. Part XII - the subfamily

Ocenebrinae (in part). Tulane Studies in Geology and

Paleontology 29: 69-118.

Vokes, E. H. 1988. Muricidae (Mollusca: Gastropoda) of the

Esmeraldas beds, northwestern Ecuador. Tulane Studies

in Geology and Paleontology 21: 1-50.

THE NAUTILUS 119(4):169-173, 2005

Page 169

A new species of Falsimargarita (Gastropoda: Vetigastropoda:

Trochidae) from the South Atlantic Ocean

Eliézer de Carvalho Rios

Museu Oceanografico

Funda¢ao Universidade de Rio Grande

Cx. Postal 379

96200-970 Rio Grande

BRAZIL

Sao Paulo

Cx. Postal 42494

BRAZIL

[email protected]

04299-970 Sao Paulo

Luiz Ricardo L. Simone!

Museu de Zoologia da Universidade de

ABSTRACT

A new trochid species, Falsimargarita stephaniae, is described

from about 1200 m depth off the Malvinas (Falkland) Islands,

South Atlantic Ocean. The new species is distinguished from

the most similar congeneric ones by its exceedingly large spiral

cords located only along the periphery of its shouldered whorls.

A re-hydrated specimen allowed for the description of some

details of the anatomy of the new species, including head-foot,

buccal mass, and radula.

INTRODUCTION

Representatives of the trochid genus Falsimargarita

Powell, 1951, can be distinguished by shell characters

such as external iridescence, well-defined spiral whorls,

strong spiral sculpture, opened umbilicus, and thin shell

wall. The genus encompasses five species occurring in

the cold or freezing deep waters off Antarctica and the

Magellanic region of South America. The taxon was

more recently revised by Dell (1990), who outlined the

diagnostic characters of the genus and described two

species.

The analysis of a specimens collected by a boat deep-

fishing for king crab and tuna revealed the presence of

the new species. This paper is part of a larger project of

revision of western Atlantic molluscan species, which at

the moment is focused on the study of deep-sea trochids.

MATERIALS AND METHODS

A single specimen with dry soft parts was available for

study. Dry soft parts were carefully removed and re-

hydrated in physiological solution and 3-4 drops of KOH

20% for 3 h, then transferred to 70% ETOH. Only the

Author for correspondence

head-foot was adequately extracted. The dissection was

performed with the specimen immersed under the

fixative, in a stereomicroscope. All drawings were done

under camera lucida. The radula was removed and

cleaned in a boiling solution of KOH for 1 h, then

cleaned by sonication in water. The examination was

done under a Zeiss electron microscope at the

Laboratério de Microscopia Eletrénica of Museu de

Zoologia da Universidade de Sao Paulo.

Institutional abbreviations used in this article are:

MORG, Museu Oceanografico da Fundagao Universi-

dade de Rio Grande, Rio Grande, Brazil: MZSP, Museu

de Zoologia da Universidade de Sao Paulo, Sao Paulo,

Brazil.

SYSTEMATICS

Genus Falsimargarita Powell, 1951

Type species: Margarites gemma Smith, 1915; by

original designation, Antarctica.

Falsimargarita stephaniae new species

(Figures 1-12)

Diagnosis: Shell with broad spire, 5 prominent large

and tall spiral cords restrict to periphery; umbilicus

protected by strong plate.

Description: SHELL of medium size (16.6 mm), tro-

choid to turbiform, whitish, iridescent-gray to pale-

reddish; wall relatively thin, light. Protoconch of one

smooth, glossy whorl. Separation protoconch-teleoconch

poorly defined. Spire with 3.5 teleoconch whorls; each

whorl highly convex, relatively high and shouldered;

superior half weakly descendent, sculptured by 6-7 low

and narrow spiral and numerous axial lines, both equally

predominating; inferior half abruptly descendent, sculp-

Page 170

THE NAUTILUS, Vol. 119, No. 4

Figures 1-5.

tured by five strong and prominent spiral cords, two

adapical and one abapical cords clearly larger, latter cord

coinciding with suture; smooth area bearing only growth

lines present between cords of abapical half of whorl.

Body whorl about twice spire width, sculptured with five

strong spiral cords that continue unchanged from spire;

five spiral cords restricted to periphery area of body

whorl; base sculptured with 18 spiral lines successiv ely

and gradually broader and more spaced toward umbi-

licus. Body whorl well separated from umbilicus by

larger spiral cord, almost a low carina. Umbilicus open,

Falsimargarita stephaniae new species. 1-4. Shell of holotype, apertural, abapertural, apical, and umbilical views

(larger diameter 16.6 mm). 5. Operculum, outer view.

deep, surface with simple growth lines; a strong plate-

like expansion of the inner lip separates this latter

from umbilicus. Aperture rounded, ample. Inner lip

slightly deflected on abapical half, somewhat thick;

adapical half marked only by thin glazed area on

body whorl; no callus present. Outer lip rounded,

thin, with small projections corresponding to spiral

sculpture.

Heap-roor: Total length about % length of last shell

whorl. Head protruded, occupying about % of total

head-foot volume. Snout with about % of foot size,

E. C. Rios and L. R. L. Simone, 2005

Page 171

Figures 6-11. Falsimargarita stephaniae new species. 6. Buccal mass, ventral view, ventral wall opened longitudinally along

median line and deflected to expose jaws. Scale bar = 1 mm. 7-8. Left jaw plate, outer and inner views. Scale bar = 0.5 mm. 9-11.

Radula. 9. General view; 10. Detail of central and lateral teeth. Scale bars = 100 um. 11. Detail of lateral teeth, arrow indicating

fifth lateral tooth. Scale bar = 50 um. Abbreviations: jw, jaw; mo, mouth; od, odontophore; ra, radula; rs, radular sac.

cylindrical, broad; distal surface flattened, fully covered

with small papillae; each papilla cylindrical, tip rounded;

mouth central. Tentacles long (about twice snout

length), narrow, tip rounded. Ommatophore with about

Ys of tentacle length and approximately with same width;

located just posterior to tentacles; eyes dark, on

ommatophore tips. Foot occupying about 7 of head-

foot volume. Mesopodium constituting most of foot,

outline somewhat triangular; sole flat, simple; anterior

furrow of pedal glands bordered by thick margins,

restricted anteriorly, not protruding beyond lateral

edges. Epipodium divided into two apparently symmet-

rical lateral flaps, covering entire dorsal surface of

mesopodium, from snout base to opercular pad; bearing

eight pairs of long epipodial tentacles projecting about

twice longer than epipodial width, each tentacle pro-

truding on ventral but not on dorsal epipodial surface;

each epipodial tentacle bearing papillae at ventral

surface of basal region; papillae increasing in number

and size toward middle tentacles; some epipodial

tentacular papillae bifid at tip. Opercular pad with

edges as continuation of epipodium. Columellar muscle

thick, encompassing a half whorl.

OpercuLUM: Circular, horny, multispiral; nucleus

central; occupying entire shell aperture.

DicEsTIvE System: Buceal mass somewhat larger than

snout internal space. Buccal cavity having a pair of very

large, dark-brown jaw plates, outline somewhat elliptical,

Page 172

THE NAUTILUS, Vol. 119, No. 4

Figure 12. Falsimargarita stephaniae new species. Head-

foot, ventral to slightly lateral right view, epipodium deflected

upward. Scale bar = 1 mm. Abbreviations: ep, epipodium; et,

epipodial tentacle; fs, foot (mesopodium) sole; he, head; mo,

mouth; om, ommatophore; pg, anterior furrow of pedal glands;

sn, snout; te, cephalic tentacle.

both jaws occupying most of dorsal and lateral surfaces

of buccal cavity; posterior and lateral regions of jaws low,

medial and anterior regions taller, with projected edges;

series of small cusps present along anterior and medial

edges, each cusp pointed and well separated from each

other. Odontophore about half projected into buccal

cavity. Radular ribbon about three times odontophore

length; about half of radular ribbon projected beyond

posterior end of odontophore.

RaDuLa: Rachidian tooth encompassing about '/4 of

radular ribbon width, triangular, narrowing somewhat

abruptly, strongly curved over its own base; cutting edge

sharply pointed, margins with very slender, elongated

cusps. First to fourth lateral teeth long, slender,

narrowing gradually, curved inward; distal half bearing

edges with slender, elongated cusps; tip sharply pointed.

Rachidian and four more central lateral teeth thin,

flexible. Fiftieth lateral tooth thick, hook-like, curved

inward; base broad, thick; distal region arched, re-

sembling a thick scythe. Marginal teeth slender, tall,

about 20 pairs per row, slightly broader toward medial

region; base ruler-like, weakly curved inward; distal half

sharpening gradually, with several slender, elongated

cusps along edges about 25 pairs per tooth; tip sharply

pointed.

Holotype: MORG 49650 (shell and operculum),

MZSP 46559, diameter: 16.6 mm; height: 14.6 mm;

includes re-hydrated soft parts and radula.

Type Locality: Argentina, off Islas Malvinas (45°S

58°W), 1200 m depth (fishing boat col., x/2004, Helen

Racz leg.).

Distribution: Known only from type locality.

Etymology: The Latinized specific epithet honors the

collector's mother, Ms. Teodora Stefania.

DISCUSSION

Although it has been recognized that definitions of

generic boundaries in the Trochidae merits further

revision, we are reasonably certain of its generic

allocation of the new species, because the species

possesses the conchological attributes reported in the

Introduction for Falsimargarita.

There are two other genera also occurring in the

South Atlantic Ocean that also exhibit iridescent shells,

a character associated with the presence of thin outer

shell layers. One of these genera is Margarella Thiele,

1893 (see Zelaya, 2004); the new species cannot be

allocated into this genus because of its larger size,

presence of thin shell wall, and absence of a parietal

callus. The other genus is Gaza Watson, 1879; Falsi-

margarita stephaniae can not be included in Gaza given

its taller shell, more rounded spiral whorls, and absence

of a flap covering the umbilicus.

Additional comparisons and discussion about Falsi-

margarita is provided by Dell (1990: 93). At first glance,

the new species could also be assigned to the Indo-

Pacific genus Otukaia Ikebe, 1942; however, the new

species has lower profile, more elaborate sculpture, and

a more widely open umbilicus.

Falsimargarita stephaniae differs from the remaining

congeneric species F. iris (Smith, 1915), F. gemma

(Smith, 1915), F. thielei (Hedley, 1916), F. georgiana

Dell, 1990 and F. benticola Dell, 1990) by having

shouldered whorls, and by the strength of the spiral

folds. The other species have a rounded whorl profile

and a uniformity of spiral sculpture. Only F. thielei

possesses differentiable spiral cords resembling those of

F. stephaniae; however, F. stephaniae additionally differs

from F. thielei by having a larger number of those

outstandingly large spiral cords at the shell periphery

and a taller spire.

The bathymetry is also a distinctive among Falsi-

margarita species. Falsimargarita gemma, F. iris, and

F. thielei occur in depths to 400 m. Falsimargarita

benthicola and F. georgiana are found in deeper waters

around 3000 m, while F. stephaniae occurs at interme-

diary depths, around 1200 m.

Until the discovery of the new species, the only

Falsimargarita known to occur in latitudes north of 50°S

was F. iris, which reaches 35°S (Rosenberg, 2004).

Falsimargarita stephaniae is the second species recorded

for these latitudes.

ACKNOWLEDGMENTS

We thank Helen Racz for the donation of the studied

specimen; José H. Leal, The Bailey-Matthews Shell

Museum, Sanibel, Florida, for literature; Lara Guimar-

aes, MZSP, for help with SEM; Diego Zelaya and one

anonymous referee for comments on the manuscript.

This study is partially developed with financial help from

the State of Sao Paulo through grants from Fapesp

E. C. Rios and L. R. L. Simone, 2005

(Fundacaéo de Amparo a Pesquisa do Estado de Sao

Paulo), processes (4/00309-2, 04/02333-8.

LITERATURE CITED

Dell, R. K. 1990. Antarctic Mollusca, with special reference to

the fauna of the Ross Sea. Royal Society of New Zealand

Bulletin 27: i-iv, + 1-311.

Hedley, C. 1916. Australia—Antarctic expedition 1911-1914.

C — Zoology and Botany. Government Printer. Adelaide

4(1): 1-80 + pls. 1-9.

Notice

Page 173

Powell, A. W. B. 1951. Antarctic and Subantarctic Mollusca: Pelecy-

poda and Gastropoda. Discovery Reports 26: 47-196 + pls. 5-10.

Rosenberg, G. 2004. Malacolog version 3.3.3, Western Atlantic

Gastropod Database. http://data.acnatsci.org/wasp/

index.php (accessed on 07/11/2005).

Smith, E. A. 1915. Mollusca. Part I. Gastropoda Prosobranchia,

Scaphopoda, and Pelecypoda. British Antarctic (“Terra

Nova”) Expedition, 1910. Natural History Report. Zoology

2: 61-112, pls. 1-2.

Zelaya, D. G. 2004. The genus Margarella Thiele, 1893

(Gastropoda: Trochidae) in the southwestern Atlantic

Ocean. The Nautilus 118: 112-120.

)

BIVALVIA 2006 - INTERNATIONAL CONGRESS ON BIVALVIA, BELLATERRA (BARCELONA), SPAIN, 22—27 JULY 2006

The congress, to be held at the Universitat Autonoma de Barcelona, calls together neontologists and palaeontologists with research

interests in bivalve mollusks. Plenary talks include population genetics, evolution of ontogeny, evolutionary paleontology,

biomineralization, and freshwater conservation biology, but contributions need not be restricted to these topics. In addition, there

will be a planning session for a new bivalve treatise.

Two one-day excursions—one each on recent and fossil bivalves—will be organized.

Interested parties are asked to register and submit abstracts via the congress webpage http://bivalvia2006.uab.es.

Further inquiries may be directed to Niko Malchus ([email protected]). Please include in the subject line “Bivalvia 2006”.

THE NAUTILUS 119(4):174, 2005 Page 174

Notice

2006 MEETINGS OF THE AMERICAN MALACOLOGICAL SOCIETY AND THE WESTERN SOCIETY OF MALACOL-

OGISTS, SEATTLE, WASHINGTON, 29 JULY—3 AUGUST 2006

The 72"? annual meeting of the American Malacological Society and the 39th annual meeting of the Western Society of Malacologists

will be held jointly in Seattle from 29 July—3 August 2006 under the coordination of AMS and WSM co-president Dr. Roland C.

Anderson. The meeting’s main venue will be the University of Washington. Reasonably priced housing will be available at the

University dormitories and the University Inn Motel. The opening night reception will be held at the Burke Museum, located on

campus, and the closing banquet will be at UW’s University Club, also on campus. Thursday, 3 August, will be devoted to field trips.

The meeting will include three symposia: one on cephalopod behavior organized by Jennifer Mather of the University of Lethbridge,

one on chitons organized by Douglas Eernisse of the California State University at Fullerton, and one on opisthobranchs organized

by Sandra Millen of the University of British Columbia, Canada.

There will be a sale of malacological reprints to benefit the student fund of WSM and the traditional spirited auction of books and

molluscan memorabilia (no shells) that will benefit the student funds of both organizations. Several notable items of cephalopod art

have already been donated as well as a copy of R. T. Abbott's 94 edition of American Seashells. Bring some of your reprints, books,

and molluscan art to benefit this very worthy cause! Reprints and auction items can be sent to Roland Anderson at the address below.

For further information please contact AMS and WSM president:

Dr. Roland C. Anderson

1483 Alaskan Way

Seattle, WA 98101 USA

[email protected]

Neve ler I tL Us

Volume 119

2005

AUTHOR INDEX

ANRIDTIGAS INI, Ingo es ande tacos aoe Combi au oe tbe nan ata taa tInG 157 INIISTESIEN ERS Be sree tssstsisys csi orae olay Waco aereenais Gee ERPs 153

[BVATRUUHIOIK. ISS obo ea SEO O ORS RD EIR OUT DR STE CaO REC Iee 43 INTORIBISAU Vier ome pean ys masse sian srs toacrcseaer Nayosines Mee nc fguraseneE 83

(CARIBOINIINIZ GAG Ifo se rapeedo secs bicot mone Reecdame ea nanee arenes 157 [ROR EU Biel Oe aeacee.ctre cau Sac am Meets tam ean ain aera ern eae tetas 169

(CONRIDDINIIEIRU MSs ntroe sees mone nob an eet aee a eae Oeeer Ener arenes 105 IPAGHONMN Ol: Pasa sige ou soomasoa veut cmon com oun sees tocomedes eee)

(COARIRAINIANS Nite 2 eA e oeamcenueboseen oud asta cde te taster tere ne 83 ROS Ke Ne apeereseres stator toes te cfavnctoiarrt Siepaniecd ysis oases nsieesea ste orator wePDE il

IDYoNWinito), Iho! | BaecocgucodesaabHboosdeanvans db anoeoaceeannanee: 164 IRVOIGAINEE! Dieters anh o @ dum etotbo aera cinta Do POA RC ie eer ene mm EEN cate 109

(FAURE OEE Vile taney eitan retraction aero reraenae enlace 15 IRVOVREUS I Blosira wine hs cra ame Re SD enn ret er Sa ER re 116

ID iA02;, If, LS 55 Sob oes anoe des Cena sO Sas Hoenn Senne amte onae 157 S\itdite JiR, sngscedebooyseacsdaoted anoet guadeeanpescoeondaas as 133

HTETENRUANS Fe VVav(@ EL sg N/053 Gy cpepens ares. Jere pae hatetors cisctsvevencisestess Aare nien Sooradicavadeters 149 SVANZAUIST Rts) boar as one nn a einieah Grice ia toe inerita pele enlac aaa 11

IRIARTNTSLID) 12s ID sse6 nomseans oan nen daaandasene soe nes onSEa ee onsaae 1 STEN @ NUE Se el eI eee ey a ae 169

lenuauninn: (Ch bey aos eb amawelbeso Ne how ON Een Cac ano ree nee ae 93 SIDAZOINGIOG \[sbobesaboodeopponctossunaoons sopaovEdsaneoaaunsenoe NT

TRAINARO IR 75 deraeeetoen sae dite ars Hive Se Ott Baek ete ena Reon 149 SMG Oy Cho ea aeaecmarcedadeadees sonereeomannn abcgoracmeionts 109

IXGIILIDIN, JSP erode naa seb aO Sona con mame cee eae eect es 15, 90 SOWITIDS IR Is. a oom amen naa eae ae ned oaan eauaeme none acodaas 133

IML ARIOMNINIE WEr owoaeoenanceends ue doded ses Bede nOnE meet 109 STRONG, E. Een Se Oe MAO OREO T buate conto UL Oe none Goma Tene 119

IN UTA STEAWAG Ely cre (etary ices eieie vsi-stoe a take exes sess gl cteenraletsaaciehe a 157 VST ISHWAT Arter omc acne sr ees Tee tect rian 43, 91

IMUINGRONG Rvs conse daaeaadensoo ce ccs ta cada Somomma cea ea ean 11 NVI TINIE OFF aaah casi nlondcrd se Rano eeH ean cor mor saae saa aenae memmn aor 50

NEW TAXA PROPOSED IN VOLUME 119 (2005)

GASTROPODA

Akera julieae Valdés and Barwick, 2005, new species (Akeridae) ... 2... 11 te 44

Bullamirifica Squires and Saul, 2005, new genus (Indeterminate family).................. 0.00.00 00002020 e eee 137

Bullamirifica elegans Squires and Saul, 2005, new species (Indeterminate family)..........................0.00.. 140

Bullamirifica verruca Squires and Saul, 2005, new species (Indeterminate family)........................0.00005. 13

Calliotropis pulvinaris Vilvens, 2005, new species (Trochidae).................................5............. 50

Coralliophila trigoi Mariottini, Smriglio and Roldan, 2005, new species (Muricidae)...........................005. 110

Daffymitra Harasewych, 2005, mew gemms (VollutomminmGae) ...20c0c0neccsoccsvacsoansaagesecesagseasoonneans 149

Daffymitra lindae Harasewych, 2005, new species (Volutomitridae)................. 00. ee ee 150

BuiliaalanbeumNielseny.2005smewsspecies) (Btychatractidae) yaar ee ae ae eee eae one arnt ae aoe e as 154

Falsimargarita stephaniae Rios and Simone, 2005, new species (Trochidae) ................. 00.00.20. e se eee eee 169

Minions Scuines aul Samil, 2005, mer eemus (Cayoullickte). .o.0.0c0cecesecsaesoccen es onasonaassasososonece 143

Minytropis melilota Squires and Saul, 2005, new species (Capulidae)......................020.0202020-22 sees 143

Nerita orovillensis Squires and Saul, 2005, new species (Neritidae).......................................... 136

Paryphantopsis abstrusa Slapcinsky, 2005, new species (Charopidae).................. 5.0052 e eee eee eee 29

Paryphantopsis koragae Slapeinsky, 2005, new species (Charopidae) .................. 5.0.0 e ee eee eee eee, 31

Paryphantopsis lets Skpcinelsy, 2005, many gaecies (Clramaaie) ....00000c0c0ccsnensccccsccsveeaecosocccsen t 33

Paryphantopsis matawanensis Slapeinsky, 2005 a(Charopidae) icc, es cards ous ws Ae Senay Gatco: ane Moyoacie owen ete 3

Paryphantopsis ubwamensis Slapcinsky, 2005, new species (Charopidae) ....... 2... 2... eee 37

Raryphantopsis yaw Slapcmskys 2000s mews species (Charopidae)y ass 45 454042420200 40 08 oe os oes oe see se eee se 39

Poxiimopis Squines amcl Sail, A005, mony eens (Chyoulticke), ooo. 0s020ceceses0ccsaaesccncececsasonssononseane 144

Paxitropis dicriota Squires and Saul, 2005, new species (Capulidae).............. 0.50.00 eevee eee eee eee 144

Picroryiis jacoana IDWintes, 20S, many gaecies (MiwHCICE). . 50205050008 0000405050009080505505655055500000" 165

Tegulla jeamae Soiines amel Samll, 20S, many Gaecies (Wola) . o.oo 202000020080e0080590000005000050c00000R" 13

Tiroplnon jromodhia! Pastore, A005, mesy jaeCeS (MiumieICAS). «oo 00000000es0cesocoscsoenaoaegesacncnaecogecas 76

BIVALVIA

rlemmiong Ikoe amel Ilarcanelll, 2005, mewy gemus (Whatomiclk) 5 oo 50505 c0es0s0ecoacs cos sosoe oss bodeseeoosuesDe 1

Pisichionm, chide Naerte, A005, mew Gaxecies (Solnaenik) . 5 o>0c00000700000¢900008950000000000500004504500505 100

Pisichionm ccloua Maerte, 2005, mew Goecies (Solmxenticke). .4o.0ccc0vescsesecececossacanseuscscsssgnasonsbane 97

Pisicionmn omazoaca Wane, 2005, mew Goecies (Splanenticke) . . . os 0002020000 a0s0ane yee aesesoageos0 Des o50aGS 94

Philippe Bouchet

Arthur E. Bogan

Robert H. Cowie

Kevin S. Cummings

Marta J. deMaintenon

Emilio F. Garcia

Daniel L. Graf

M. G. Harasewych

Roland Houart

REVIEWERS FOR VOLUME 119

Steffen Kiel

Sadao Kosuge

Bruce A. Marshall

Paula M. Mikkelsen

Russ Minton

Guido Pastorino

Maria del Carmen Perrilliat

Richard E. Petit

Timothy Rawlings

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