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NAUTILUS

Volume 121, Number 1

March 28, 2007

ISSN 0028-1344

CONTENTS

Richard L. Squires Paleocene pareorine turritellid gastropods from the Pacific slope of

LouElla Saul NOVEM AMIETICA: joi tec aicns 1 atbrdiad te. AGBcand: deal aid. Pole Mo al ahh a eh dew, Blues Se a) dane l

Brian F. Coles Vertigo malleata, a new extreme calcifuge land snail (Gastropoda:

Jeff Nekola Vertiginidae) form the Atlantic and Gulf coastal plains of the USA... 2... 17

Meghna Roy Population dynamics of the fingernail clam Sphaerium occidentale

D. Dudley Williams (Lewis, 1856) (Bivalvia: Sphaeriidae) in an intermittent pond ........... 29

Research Note

Ilya V. Buynevich Paleoenvironmental significance of the eastern mudsnail, [lyanassa obsoleta

(Say, 1822), from a microtidal coastal sequence of southern New England 37

BGTACAY oo, faethe inn coe hes hee a aA ga SNe Res, aha near Gog apa Qabed Me, iad Poa ae a aoe Mp ae 40

THE NAUTILUS 121(1):1-16, 2007

Page ]

Paleocene pareorine turritellid gastropods from the Pacific slope

of North Americz

Richard L. Squires

Department of Geological Sciences

California State University

Northridge, CA 91330-8266 USA

[email protected]

Angeles County

LouElla R. Saul

Invertebrate Paleontology Section

Natural History Museum of Los

900 Exposition Boulevard

Los Angeles, CA 90007 USA

[email protected]

ABSTRACT

This paper presents the first detailed study of Mesalia martin-

ezensis (Gabb, 1869) and Mesalia clarki (Dickerson, 1914a), the

only two known pareorine (spout-bearing) turritellid gastro-

pods from the Pacific slope of North America. Both species are

redescribed, in light of new morphologic information that also

confirms their assignment to genus Mesalia Gray, 1847, which

we believe to be congeneric with Sigmesalia Finlay and Mar-

wick, 1937. New stratigraphic information allows for refine-

ment of the chronologic range of each species. Mesalia mar-

tinezensis is of early late Paleocene (late Danian) to early late

Paleocene (early Thanetian) age and ranges from northern

California to northern Baja California. Mesalia clarki is of late

middle to early late Paleocene age (late Selandian to early

Thanetian) age and is known only fom California; in southern

California it is restricted to a coralline-algal facies. Both species

have considerable variability in their spiral sculpture.

Mesalia originated during either the Late Cretaceous (Maas-

trichtian) in northern Africa or the early Paleocene (Danian) in

northern Africa and western Iran. It became widespread during

the warm (greenhouse) conditions of the Paleocene and

Eocene but became geographically restricted during subse-

quent cooler global conditions. Mesalia is an extant genus with

possibly six species, and whose total geographic range is in

coastal waters in southern Portugal, southern Spain, Mediter-

ranean Sea (primarily the western part), Canary Islands, and

the west coast of northern Africa.

INTRODUCTION

The object of this study was to rectify the identification

uncertainities concerning the only two known pare orie

spout-bearing) turritellid gastropod species from shal-

low-marine rocks on the Pacific slope of North America.

They are Mesalia martinezensis (Gabb, 1869) and Mesa-

lia clarki (Dickerson, 1914a). va modern workers

Corresponding author: Richard Squires

email: [email protected]

(e.g., Zinsmeister, 1953) have generally placed both spe-

cies in Mesalia Gray, 1847, buat. some workers (e. g., Saul,

1983a: fig. 2; Squires, 2003: table 2.4) have been ie ssitant

to unequivocally use the genus name because of inad-

equate morphologic information about these species.

The shape of the aperture of the latter one was unknown

until now, and apertural information is critical in distin-

cuishing parerorine gastropods from similar looking tur-

ritellids (see “Systematic Paleontology” for morphological

comparisons). There has also been the possibility that

Mesalia macreadyi Waring, 1914, which has commonly

been put into synonymy with M. martinezensis, could be

a distinct species.

We conclude that there are only two species, both

belonging to Mesalia, which we believe to be congeneric

with Sigme salia Finlay and Marwick, 1937. In addition to

providing new morphologic information about the study

area Mesalia, we retine their geographical (Figure 1) and

chronologic ranges (Figure 2), Both M. martinezensis

and M. clarki have considerable variability in their spiral

sculpture.

Mesalia has long been reported (e.g., Cossmann, 1912)

as originating during the Late Cretaceous in the region

between northern Africa and western Iran. Our review of

the literature shows that the genus most likely originated

during either the Late Cretaceous (Maastrichtian) or the

early Paleoce me (Danian) in this Old World region.

Modem systematic studies of Mesalia are sorely lack-

ing as are detailed ecologic studies of the few extant

species. The classification scheme used here follows that

of Bouchet et al. (2005: 249), and tase ar terms

are taken from Cox (1960). Pacific slope of North

America Turritella zones are taken from eel (1983b).

Institutional abbreviations used in the text are: ANSP:

Academy of Natural Sciences of Philadelphia; LACM:

Natural History Museum of Los Angeles d ounty, Mala-

cology Section; LACMIP: Natural History Museum of

Los Angeles County, Invertebrate I

UCMP: University of California, Berkeley, Museum of

Paleontology.

Paleontology Section;

Page 2

THE NAUTILUS, Vol. 121, No. 1

Latitudinal Distribution

of Species

]-Lower Lake

2-Martinez

3-Mt. Diablo area

4-Big Rock Creek

5-Simi Hills

6-Santa Monica Mountains

7-Mesa San Carlos

California a

Mesalia clarki

Mesalia martinezensis

Figure 1. Locales and latitudinal distribution of the study

area gastropods.

STRATIGRAPHY AND DEPOSITIONAL

ENVIRONMENTS

The geologic ages of the formations and most of the

depositional environments of the formations containing

the two studied species are mentioned in Squires (1997).

Mesalia martinezensis is widespread in the study area

(Figure 1) and always found in siltstone or sandstone

be ds that formed either as storm accumulations of mol-

lusk-rich assemblages in shelfal-marine depths or as lo-

calized displaced shallow-marine mollusks deeper

Age System Series European Mesalia Mesalia Turritella

(Ma)| S¥Stemy/penies Stages | martinezensis| clarki Zones

Lower

sean] |

T. peninsularis

Danian T. peninsularis

qualeyi

T. infragranulata

T.1. pachecoensis

Middle} Upper

Lower Paleogene

Paleocene

Lower

Upper Cret.]| Maastr.

Figure 2. Chronostratigraphic position of the study area taxa

Ages of stage boundaries from Gradstein et al. (2004). Turri-

tella zones from Saul (1983b

depths. These mollusks commonly include shallow-

marine mollusks such as naticid and buccinid gastropods,

as well as glycymerid and crassatellid bivalves. All are

indicative of normal marine salinities.

The type locality of Mesalia martinezensis has been

generally assigned (e.g., Keen and Bentson, 1944) to the

“Martinez Formation” in the vicinity of the city of Mar-

tinez, Contra Costa County, northern California (Figure

1). The history of how early workers referred to “this

originally poorly defined “formation” has been summa-

aed by Mallory (1959). In this present study, we follow

the work of Weaver (1953), who refined the str atigraphy

of the Paleocene and Eocene formations in the v icinity of

the area where the “Martinez group” was first estab-

lished. He established new stratigraphic names, and the

rocks that pertain to the type locality of M. martinezensis

belong in his Paleocene Vine Hill Sandstone.

Mesalia clarki is only known from two locales: 1) its

type locality (UCMP loc. 1540, see “Appendix”) on the

north flank of Mount Diablo, Contra Costa County,

northern California, and 2) from the Santa Monica

Mountains, Los Angeles County, southern California

(Figure 1). Its type locality is near the site of Stew artville,

and approximately 24 km east-southeast of ihe ry oF

Martinez, and, according to Dickerson (1914a: 74), this

locality is “300 to 400 ft. above the base of the ates

in hard, gray-green glauconitic sandstone.” Numerous

mollusks have been found at this locality (Dickerson,

1914a: 75). They consist of turritellid and buccinid gas-

tropods, crassatellid bivalves, and other shallow-marine

species, all indicative of normal marine salinities. Turri-

tella infragranulata pacheocensis Stanton, 1896, which is

found at this locality, is indicative of the upper middle

Paleocene (upper Selandian) (Figure 2). On the geologic

map of Brabb et al. (1971), the locality plots within the

glauconitic sandstone lower member of the “Martinez”

Formation. Megafossils are generally scarce in the “Mar-

tinez” Formation in the vicinity of this type locality (E.

), thus, it seems plausible that

the fossils probably occur in storm-derived, isolated

Brabb, personal commun.)

lenses.

Mesalia clarki in the Santa Susana Formation in the

Palisades Highlands area of the Santa Monica Moun-

tains, southern California, is always found near outcrops

of coralline-algal limestone. Hoots (1931: 91-92, 133-

134, pL L9B) reported that these limestones are resistant,

can be cliff forming, weather white, are nodular and ir-

regularly bedded, up to 35 m thick, up to 1200 m in

lateral extent, and commonly terminate in an abrupt wall.

Additional ge ologic and/or pi aleontologic details concern-

ing these limestones are mentioned in Strathearn et al.

(1988), Colburn (1996), Squires and Saul (1998), Squires

and Kennedy (1998), and Squires and Saul (2001).

At LACMIP loc, 10508, in the Palisades Highlands

area, abundant specimens of M. clarki ave found in a thin

coralline-algal-rich micaceous muddy siltstone bed about

| m_ stratigraphically below a blocky, coralline-algal-

limestone interval approximately 24 m thick. Also fouicd

R. L. Squires and L. R. Saul, 2007

Page 3

in this bed is the large neritid gastropod Corsania (Jan-

uncia) rhoga Saul and Squires, 1997, as well as the bi-

valves Plicatula lapidicina Squires and Saul, 1998, and

Plicatula trailerensis Squires and Saul, 1998. Occurring

in nearby beds in close association with the coralline-

algal deposits are the gastropods Terebralia susana

Squires and Kennedy, 1998, and hie abs ereenellum

Hanna and Hertlein. 1939. All of these aforementioned

mollusks indicate very nearshore, tropical to subtropical

conditions (Squires and Saul, 1997; Squires and

Kennedy, 1998). The latter workers concluded that the

coralline-algal limestones, like those at locality 10508,

were deposited in a protected bay (no deeper than 40 to

70 m) with warm-algal-limestone buildups associated

with shoals on the bay floor. These limestone buildups

are very similar in lithology and sedimentologic/tectonic

setting to limestones in the Paleocene Sepultura and Ba-

hia Ballenas formations in northern Baja California (Ab-

bott et al., 1995), as well as similar to limestones in the

upper Paleocene to lower middle Eocene Sierra Blanca

Limestone in Santa Barbara County, southern California.

These limestones were deposited when tectonic plate-

edge strain in the fore-arc basin caused local basement

highs to form within the otherwise deeper marine envi-

ronment (Whidden et al., 1995; Abbott et al., 1995). It is

likely that the Santa Susana Formation coralline-algal

limestones formed under similar conditions.

Although Mesalia clarki and Mesalia martinezensis

hoe occur in the Santa Susana Formation in the Santa

Monica Mountains, southern Califormia, they never oc-

cur together in the same beds. Mesalia martinezensis is

not associated with the coralline-algal facies there or any-

where else in the study area.

PALEOBIOGEOGRAPHIC IMPLICATIONS

Kiel and Bandel (2004: 120, fig. 71) reported two speci-

mens of Mesalia cf. multilineata (J. Miiller, 1851) from

Cenomanian strata at the Kasssenberg quarry in Ger-

many. If these specimens actually belong to Mesalia, they

would be the geologically oldest. The conical-turriculate

shell with convex whorls bearing strong spiral ribs does

resemble Mesalia, but there are no specimens with an

intact aperture nor with a protoconch. Turritella multi-

lineata J. Miiller (1851: 29, pl. 4, figs. 4, 6) was originally

reported from the lower Campanian Aachen strata of

Germany, thus, it is considerably younger than the Kass-

senberg quarry material. Turrite lla mattiline cata Was also

figured by G. Miiller (1898: 97, pl. 13, figs. 4, 5), who

reported it from middle Santonian to lower Campanian

strata at Braunschweig/Ilsede, Germany. There is also a

mention of T. multilineata in Kollmann and Odin (2001:

441), and they also consider this Campanian species to

belong to genus Turritella. The pleural angle of J.

Miiller’s figure is much narrower than that of Kiel and

Bandel (2004), and in our opinion, Kiel and Bandel’s M.

cf. M. multilineata (J. Miiller) does not look like J.

Miiller’s species. Kiel and Bandel (2004) tentatively syn-

onymized ]. Miiller’s specimens and G. Miiller’s speci-

mens with their Kassenberg quarry specimens. We be-

lieve, however, that Kiel Gnd Bandel’s Cenomanian

specimens have nothing to do with T. multilineata and

represent, at best, a very questionable Mesalia. More

specimens of this possible Mesalia from the Cenomanian

of Germany are needed in order to determine its generic

assignment.

Cossmann (1912: 126) reported the chronologic range

of Mesalia to be Late Cretaceous (Turonian) to Recent,

as did Wenz (1939), who apparently simply reiterated

Cossmann’s findings. We were unable, however, to cor-

roborate any of Cossmann’s Cretaceous occurrences. He

reported Mesalia gazellensis Whitfield (191; 424, pl. 9,

fig. 10) as being from the Turonian of Syria, but the

aperture of his, species is unknown. In addition, the

sculpture is obsolete, which is unlike Mesalia.

Cossmann (1912: 126) listed five Mesalia species of

Late Cretaceous (Senonian) age, and these are discussed

in the following sentences. Arcotia indica Stoliczka

(1868: 215, 469, pl. 16, figs. 12, 12a; pl. 19, fig 6) from

southern India is not a Mesalia. This species is also dis-

cussed later under “Systematic —. Specimens

of Turritella ventricosa Forbes (1846: 123, pl. 13, fig. 3;

Stoliczka, 1868: 227, pl. 17, fig. 15) ‘ne southern Indi ia

are missing the aperture. Turritella martinezensis Gabb

(1869: 159. pl. 28, fig. 51) from California is not of Cre-

taceous age. Mesalia nettoana White (1887: 164-165, p

18, figs. 3, 4) from the Maria Farinha beds in Brazil is

oe age. Mesalia hebe White (1887: 165, pl. 1S, fig.

), also from Brazil, looks like a juvenile specimen of M.

ne eer

Cossmann (1912: 126) also listed two Late Cretaceous

(Maastrichtian) species. The first one is Mesalia jovisam-

monis (Quaas, 1902: 256, pl. 26, figs. 1S—20), which

Quaas reported, in a very generalized way, to be associ-

ated with the Exogyra overwegi biozone at Ammonite

Hill in the Great Sand Sea in western Egypt. This bio-

zone can also be recognized in the Maastrichtian (but not

latest Maastrichtian) part of the Ammonite Hill Member

of the Campanian to Paleocene Dakhla Formation in

western Egypt (Barthel and Herrmann-Degen, 1981).

Tantawy et al. (2001) assigned this member an early to

early late Maastrichtian age, based on planktic forami-

nifera, calcareous siannofossils , and macrofossils. They

also determined, however, that the entire formation

ranges in age from early Maastrichtian to early Danian.

Immediately above the widespread K/T disconformity in

the region, a sedimentologically complex sequence

marks the lower Danian Bir Abu Minquar horizon, which

contains a mixture of Maastrichtian (reworked) and Da-

nian fossils, including both microfossil and macrofossil

species (e.g., including some ammonites.). Unfortu-

nately, Quaas did not “provide any information as to

where actly in the stratigraphic section he collected the

specimens of M. jovisammonis. His specimens were lost,

so it is not possible to match their rock matrix to actué ul

outcrops. Recollecting of this gastropod is necessary to

decipher its exact geologic age.

Page 4 THE NAUTILUS, Vol. 121, No. 1

Figures 3-15. Type species of Mesalia and Sigmesalia, plus comparative pictures of Mesalia solida (Deshayes, 1861). Specimens

coated with ammonium chloride, 3-9. Mesalia mesal (Adanson, 1757), Baie de Hann, Senegal, West Africa (Recent). 3-7.

Hypotype LACM 173163. 3. Apertural view, height 45 mm, diameter 15 mm. 4. Tip of specimen shown in previous figure, height

14 mm, diameter 5.5 mm. 5. Protoconch and earliest spire whorls, apertural/ right-lateral view, height 1 mm, diameter 0.7 mm. 6.

Base, diameter 15.6 mm. 7. Abapertural view, height 45 mm, diameter 15 mm. 8-9. Hypotype LACM 173164. 8. Oblique apertural

view, height 51 mm, diameter 16.3 mm. 9. Close-up of abapertural view of last whorl, diameter 17.2 mm. 10-13. Mesalia koeneni

(Le Renard, 1994), LACMIP hypotype 13397, Grignon, Paris Basin, France (middle Eocene, Lutetian Stage), height 45.8 mm,

diameter 20.3 mm. 10. Apertural view. 11. Close-up of aperture. 12. Right-lateral view (outer lip broken), 13. Base. 14-15. Mesalia

solida (Deshayes, 1561), hypotype LACMIP 13398, Le Guépelle, Paris Basin, France (late Eocene). 14. Apertural view, height 21.6

min, diameter S.1 mm. 15. Protoconch and earliest spire whorls, apertural to slightly right-lateral view, height 1 mm, diameter 0.7

Tim.

R. L. Squires and L. R. Saul, 2007

Page 5

Abbass (1963: 39-40, pl. 2, figs. 20-22) illustrated M.

jovisammonis from eastem Egypt and referred to it as

Mesalia (Woodsalia) jovisammonis of Maastrichtian age.

He did not provide, however, any discussion as to how

this age was determined.

Mesalia cf. M. multisulcata (Lamarck 1804), ea

by Serra (1937: 313-315, pl. 16, figs. 12, 12a, 13) from

near Tripoli, Libya, looks like it might be conspecific

with Mesalia jovisammonis. Serra provided no detailed

stratigraphic or geologic age information.

The second species that Cossmann (1912) listed from

the Maastrichtian is Mesalia fasciata (Lamarck, 1804)

from Iran. Cossmann based this occurrence on work by

Douvillé (1904: 329-330, pl. 47, figs. 23-27), who re-

ported M. fasciata from the * ‘Couches: a Cérithes” beds in

the Luristan region in west-central Iran. Douvillé (1904)

believed that these Iranian specimens of M. fasciata,

whose type locality is in middle Eocene (Lutetian Stage)

strata at Grignon in Paris Basin, France (Eames, 1952:

34), are of Maastrichtian age, but the “Couches a

Cérithes” beds contain the bivalve “Cardita” beaumonti

dArchiac and Haime, 1854, which is diagnostic of earli-

est Danian age in Iran and Pakistan (Douvillé, 1928:

Eames, 1952; Davies, 1975). Mesalia fasciata is long-

ranged geologically (early Paleocene to late Eocene) and

widespread geographically (western Europe to Pakistan)

(Eames, 1952).

Another Mesalia that needs investigation as to its pre-

cise stratigraphic position and geologic age is Mesalia

foucheri Pervinquiére (1912: 44, pl. 3, figs. 6-15), from

the Ghadames (Garat Temblili) region in Tunisia, north-

ern Africa. Pervinquiére (1912: 336) reported the species

as being of Maastricthian age, but no critical geologic

details are provided. He did differentiate between Maas-

trichtian and Danian fossils; thus, like in nearby Egypt

and Libya, the stratigraphic section containing M.

foucheri and other macrofossils in Tunisia, also spans the

K/T boundary.

Two species of so-called Woodsalia Olsson, 1929, from

Upper Cretaceous (Campanian?) strata in northwestern

Peru (Olsson, 1944) might eventually be placed in Me-

salia, once their apertures become known. They oo

Woodsalia paitana Olsson (1944: 69-70, pl. 11, fig. 5)

and W ‘oodsalia paitana robusta Olsson (1944: 70, pl. 11,

figs. 3, 9).

The so-called Mesalia (Mesalia) mauryae Allison

(1955: 414415, pl. 41, fig. 3; Perrilliat, 1989: 149, fig.

51h) from the upper Aptian upper member of the Alisi-

tos Formation, Punta China region, Baja California,

Mexico, is, according to Squires and Saul (2006), Turri-

tella seriatimgranulata Roemer, 1849.

In addition to the above-mentioned Old World Danian

species of Mesalia, three New World Danian species are

known from the Gulf Coast of the southeastern United

States. They are from the Clayton Formation (Palmer

and Brann, 1966), which is of earliest Danian age (Dock-

ery, 1986). The three species are: Mesalia silenkene nsis

(Aldrich, 1894: 246-247, pl. 13, figs. 4a, 4b, 6; Stenzel

and Turner, 1942: card 110); Mesalia hardemanensis

(Gabb, 1860: 392, pl. 68, fig. 15; Stenzel and Turner,

1942: card 116); and Mesalia pumila (Gabb, 1860: 392

pl. 68, fig. 14; Stenzel and Turner, 1942: card 118).

In suminary, our search of the bieranare revealed that

Mesalia most likely originated during either the Maas-

trichtian in northern Africa or the early Paleocene (Da-

nian) in northern Africa and western Iran. During the

Danian it spread quickly to the Gulf Coast of the Wned

States by means of westward-flowing ocean currents

emanating from the western Tethyan region. These cur-

rents, which existed during the Late Cretaceous (Gor-

don, 1973; Johnson, 1999) aad continued into the Paleo-

cene and Eocene (Saul, 1986; Squires, 1987), were part

of a circumglobal-tropical current that contributed to a

widespread dispersal of marine biota (Haq, 1981). By the

late Danian, it reached California and northern Baja

California, Mexico, as well as Belgium (Cox, 1930; Glib-

ert, 1973). By middle Paleocene, it reached Greenland

(Kollmann and Peel, 1983), and by the late Paleocene, it

reached Nigeria (Adegoke, 1977). During the Paleocene

and Eocene, Mesalia reached its peak div ersity and be-

came most widespread, with occurrences mainly in the

Old World western Tethys Sea region. We did not de-

tect, however, any reported occurrences in Australia,

New Zealand, Japan, or Antarctica. The Paris Basin of

France (see Cossmann and Pissarro, 1910-1913), south-

western Nigeria (Adegoke, 1977), and the Gulf Coast of

the United States (Stenzel and Turner, 1940, 1942:

Palmer and Brann, 1966) are the principal areas in which

numerous species of Mesalia have been anlage

Some species became very widespread. For example, a

mentioned earlier, Mesalia fasciata ranged trom ae

Paris Basin, France to Pakistan (Eames, 1952). After the

warm greenhouse conditions that existed during the

Eocene, Mesalia was much reduced in its distribution

and mainly occurred in what is now the Mediterranean

Sea region (Cossmann, 1912).

MODERN MESALIA

Mesalia is extant and review of the scant literature, as

well as use of the intemet (note: <http:/Avww.alboranshells

.con/turritellidae> was particularly helpful), revealed as

many as possibly six species. The they are the ae

M. mesal (Adanson, 1757), M. brevialis Lamarck, 829.

M. varia Kiener, 1843; M. opalina Adams and See

1850; M. freytagi von Maltzan, 1884; and M. flammifera

Locard, 1897, which includes the subspecies M. flam-

mifera flammifera Locard, 1897 and M. flammifera sim-

plex Locard, 1897. There is much confusion as to exactly

how many species there are, and potential synonyms

need to be resolved. For example, some workers (e. o.,

Smith, 1915; Bowles, 1939) equated M. mesal with M.

brevialis, but other workers (e.g., Advovini and Cos-

signani, 2004) separated them. Bowles (1939) gave a

thorough review of the nomenclatural history of Mesalia

brevialis.

Page 6

THE NAUTILUS, Vol. 121, No. 1

A comprehensive malacological study of the modern

species of Mesalia is greatly needed. Because of the un-

certainties stemming from the poorly y known sy! stematics,

it is confusing to try to determine which species is found

where. We were able to establish with certainty (see

references below), however, that modern Mesalia is only

found in the Atlantic coastal areas of southern Portugal,

southwestern Spain, Morocco, Canary Islands, Western

Sahara, Mauritania, Senegal, and Guinea, as well as in

the westernmost RicAnereanean Sea, particularly in the

Alboran Sea (i.e., Strait of Gibraltar to southerm Spain on

the north and Morocco on the south) and the Aegean Sea

region of western Turkey.

Mesalia mesal and M. brevialis have the widest distri-

bution. Mesalia mesal has been reported from the Al-

garve region of southern Portugal, the Algeciras region of

southwestern Spain, and the “‘Alboran Sea ( (Poppe and

Goto, 1991), the Canary Islands (Macedo and Borges,

1999), Senegal (Bouchet, 1977: fe nae and Cossignani,

2004), and western Turkey (Demir, 2003). Me salia bre-

vialis has been reported from the Algarve region of

southern Portugal (Afonso et al., 2000; Alves et al., 2003),

southwestern Spain and the Alboran Sea (Hidalgo,

1917), Senegal (Ardovini and Cossignani, 2004), and

Guinea (Pasteur-Humbert, 1962). Mesalia opalina has

been reported from the Canary Islands and Morocco

(Poppe and Goto, 1991), as well as from Mauritania (Ar-

dovini and Cossignani, 2004). The other modern species/

subspecies of Mesalia are apparently restricted to the

northwestern coast of Africa (Ardovini and Cossignani,

2004).

Mesalia melanioides Reeve, 1849, was reported

(Smith, 1915) to be from West Australia, but this species

is now the type species of Neodiastoma Cotton, 1932

which differs from Mesalia by having axial sculpture on

the early spire. Marwick ( 1957) summarized the system-

atics of Neodiastoma and classified it as a pareorine.

Mesalia is found today on both muddy and sandy bot-

toms in coastal waters ranging in depth from lower in-

tertidal to 20 m (Hidalgo, 1917; Pasteur-Humbert, 1962:

Poppe and Goto, 1991; Afonso et al., 2000; Demir, 2003;

Alves et al., 2003). Bouchet (1977) reported that M. me-

sal, although not common there, can be found in the

seaward part of mz WgrOve-sw, saad systems along the coast

of Senegal. Specimen LACM 17 316 ( (Figures 3 3-7) of M.

mesal was collected in pees 5m depth, on sand

and rubble in Senegal.

Large numbers of M. mesal have been reported

\fonso et al., 2000) as almost always being partially in-

faunal (with their apices pointed upward) when found on

exposed low-tide mud flats on the inner lagoon sides of

islands within the Rio Formosa coastal-lagoon system of

southern Portugal.

The modern ecological parameters mentioned above

are not totally reliable for fossil Mesalia because prefer-

ences for substrate and depth of water might have pos-

sibly changed over time. In addition, the fossil occur-

rences of Mesalia had a pan-Tethyan distribution,

whereas the modern occurrences have contracted pri-

marily to the Iberian Peninsula, Alboran Sea, and north-

western Africa.

SYSTEMATIC PALEONTOLOGY

Superfamily Cerithioidea Fleming, 1522

Family Turritellidae Lovén, 1S47

Discussion: Allmon (1996: 9-12, table 1) thoroughly

reviewed the history of the dlasaheaton of ‘umitellid

gastropods and listed the five subfamilies and all the

genera/subgenera within each subfamily. These subfami-

lies are: Turritellinae Loven, 1847; Protominae Marwick,

1957; Pareorinae Finlay and Manwick, 1937: Vermicu-

lariinae Lamarck, 1799; and Turritellopsinae Marwick,

1957. Bouchet et al. (2005) included the first four of

these subfamilies, but removed Turritellopsinae. Instead,

they included subfamily Orectospirinae Habe, 1955.

Subfamily Pareorinae Finlay and Marwick, 1937

Discussion: Pareorine turritellids are characterized

from the other subfamilies of family Turritellidae by hav-

ing an aperture obliquely effuse over the anterior end of

the columella and forming a sinus (short spout), with the

adapical margin of the sinus usually making a spiral ridge

on the eolamells | Marwick, 1957).

Mesalia can be readily identified if its aperture is in-

tact, but when it is missing, workers have commonly

misassigned it to the similar looking genus “Twrritella”

Lamarck 1799, sensu lato, a group comprising at least 35

genera and subgenera names (Allmon, 1996), all of which

are turritellines whose apertures do not have a sinus

(short spout) at the anterior end of the aperture nor have

a spiral ridge on the columella. In addition, according to

Smith (1915), the cormeous operculum of Mesalia is pau-

cispiral and not multispiral, as in “Turritella,” but this

later distinction is not useful when studying fossil spe-

cles.

Ten pareorine genera were listed by Marwick (1957),

who also provided i illustrations of the growth- line traces

of some of these genera. Comparative information about

the str: itigraphic range, grow th-line details, whorl protile,

and protoconch shape of a of these genera was given

by Allmon (1996; table 1

Genus Mesalia Gray (nomen nudum, 1840), 1847

Type Species: Cerithiwm mesal Adanson, 1757 [=Tur-

ritella mesal Deshayes, 1843], by original designation;

Recent, southern Portugal, southwestern Spain, Alboran

Sea, Canary Islands, Seneval, and western Turkey.

Description: Small to large (wp to approximately 95

mm shell height), turritelliform, slender to conical ro-

tund. Pleural angle ranges from 15° to 41°. Protoconch

conical, small, smooth, and approximately two whorls.

Protoconch to teleoconch transition gradual. Teleoconch

whorls up to 16, whorl sides convex to flattish/concave.

Sculpture on early juvenile teleoconch whorls variable,

ranging from ne varly smooth or with very fine, unicostate,

bicostate, or tricostate spiral lirae; sculpture on adult

R. L. Squires and L. R. Saul, 2007

Page 7

whorls highly variable, ranging from smooth to numer-

ous, weak to moderately strong closely spaced spiral ribs,

but less commonly with fewer and more prominent spiral

ribs. Growth lines parasigmoidal on last whorl (including

iene, lateral sinus variable in amount of concavity (fex-

ure). Aperture with shallow effuse spout, ranging from

cee constrained to broad. Adapical edge of spout

usually forms weak spiral ridge that continues onto col-

umella.

Discussion: Mesaliopsis Thiele, 1929 [type species:

Mesalia opalina (Adams and Reeve, 1850)], Recent, was

reported by Wenz (1939) to be a subgenus of Mesalia,

but future work ah show it to be synonymous with

Mesalia.

Mesalia somewhat resembles Lithotrochus Conrad,

1855, of Jurassic age from Chile, South America. Coss-

mann (1912: 125) reported Lithotrochus to be a junior

synonym of Mesalia, but Wenz (1938: 280, fig. 596) and

Cox (1960: 1245-1249, fig. 159,11) believed Lehatroe hus

to be a trochid. It is an extraordinarly large gastropod

(height 125 mm) with a wide pleural angle, deme d upper

spire, turritelliform shape, anteriorly carinate whorls, and

relatively few spiral ribs. Details of its aperture are un-

known.

Cossmann (1912: 125) also reported Lithotrochus to

be a junior synonym of Arcotia Stoliczka, 1868, whose

OPE species, Arcotia indica Stoliczka (1868: 215, 469, pl.

16, figs. 12, 12a; pl. 19, fig. 6) is from Upper Cretaceous

(Trichinopoly Group) strata near the town of Alundan-

apooram, southern India. According to Sundaram et al.

(2001; fig. 3), this town’s name is also referred to as

Alundalippur and, from information in their map, this

town is underlain by the Kulakkalnattam Formation of

Turonian age. Wenz (1939) synonymized Arcotia with

Mesalia. Finlay and Manwick (1937) reviewed the mor-

phology of Arcotia and reported that, based on_ its

straight growth lines and open umbilicus, this genus is

not a synonym of Mesalia. They reported, furthermore,

that Aroctia appears to be a mathildid. Bandel (2000)

came to the same conclusion.

Mesalia is similar to the pareorine Woodsalia Olsson

1929. whose type species, Woodsalia negritosensis Ols-

son (1929: 13-15, pl. 4, figs. 5, 6) is from lower Eocene

rocks in northwestern Peru. Woods (1922: 78-79, pl. 7

figs. 5-7; pl. 8, figs. 1-3) and Wenz (1939: 651, fig. 1852,

two views) also illustrated this species. The full shape of

the aperture of this Peruvian gastropod, however, is not

known.

Genus Sigmesalia Finlay and Marwick, 1937, was

originally erected to accommodate a group of Eocene

gastropods from the Paris Basin, France that were pre-

viously identified as Mesalia. There has been no consen-

sus as to whether or not Sigmesalia is a distinct genus.

Marwick (1957) reported it to be a separate genus, as did

Le Renard (1994). Eames (1952) reported it to be a

subgenus of Mesalia, and Palmer and Brann (1966) re-

ported Sigmesalia to be synonymous with Mesalia.

Various views of representative specimens of the type

species of Mesalia are illustrated in Figures 3-9, and

various views of a representative specimen of the type

species of Sigmesalia are illustrated in Figures 10-13. Its

type species, Sigmesalia koeneni Le Renard, 1994 [new

name for Turritella sulcata Lamarck, 1804 (original des-

ignation), non Bosc, 1801], is of middle Eocene (Lu-

tetian) age and from Paris Basin, France. Finlay and

Marwick (1937) stated that the aperture and growth lines

of the type species of Mesalia seem to be generically

different than those of the Paris Basin shells, but the ry did

not provide any details. Davies (1971: 312, figs. 677a,

677b) mentioned that the growth lines of Sigmesalia

have a more flexed outer lip sinus than does Mesalia.

According to Marwick (1957: 163), Sigmesalia differs

from Mesalia by usually having a wider pleural angle.

The following paragraphs deal with our observations con-

cerning these proposed diagnostic features of Sigmesalia.

Inspection of representative specimens of several of

the Eocene Paris Basin species, including the type spe-

cies of Sigmesalia, stored in the LACMIP collection, as

well as inspection of photographs of 17 Paris Basin spe-

cies (see Cossmann and Pissarro, 1910-1913: pl. 21, figs.

126-1 to 126-15), revealed variability in the shape of the

aperture of Sigmesalia. For example, the aperture of Sig-

mesalia incerta (Deshayes, 1832; Cossmann and Pissarro,

1910-1913: pl. 21: fig. 126-4, two views) is similar to that

of M. mesal, in that the spout is broad and not well

constrained. The aperture of Sigmesalia koeneni how-

ever, is better developed (Cossmann and Pissarro, 1910—

1913: pl. 21, fig. 126-15).

The amount of flexure of the outer lip sinus is basically

similar in all the Eocene Paris Basin species, although

Mesalia solida (Deshayes, 1861) does show some vari-

ability. The amount of flexure of this feature is variable in

M. mesal and can be similar to the amount seen on

Eocene Paris Basin species. Variability in the amount of

flexure for both the Eocene and modem Mesalia shells

can also occur in proximity of growth checks and break-

ages of the outer lip incurred during the life of the gas-

tropod.

The pleural angle of the Eocene Paris Basin shells is

quite variable, ranging from 21° to 41°, but the low end

of this range [e.g., Mesalia ecki (Laubriére, 1881; Coss-

mann and Pissarro, 1910-1913: pl. 21, fig. 126-2)] is very

close to the value (16° to 18°) on M. mesal. Sigmesalia

koeneni has one of widest pleural angles (41°; see Figures

10 and 12). For meee purposes, an illustration

(Figure 14) is provided for Sigme salia solida. It has a

relatively narrow pleural angle of 25°, more like that

found on M. mesal (Figures 3, 4, and 7)

Other morphologic features that are variable on the

Eocene Paris Basin shells are strength and number of

spiral ribs, pattern of development of sculpture on the

early juvenile teleoconch whorls, and degree of indenta-

tion of the suture. Even the strength of "he spiral ridge

(Figure 10) on the columella is variable. Mesalia mesal pos-

sesses a spiral ridge on the columella, as do most specimens

of Sigmesalia koeneni (compare Figures 8 and 11)

Page 8

THE NAUTILUS, Vol. 121, No. 1

A few species of Sigmesalia are similar to Mesalia in

having a relatively narrow pleural angle and bicostate

sculpture on the ‘early juvenile where but not on the

adult whorls. They are the following: Sigmesalia instabi-

lis (Briart and Cornet, 1873: 86, pl. 12, figs. 9a-9b) of

early Paleocene (Danian) age from Belgium; Sigmesalia

salvani (Adegoke, 1977: 86-88, pl. 14, figs. 10-16) of

Paleocene age from Nigeria; and Sigmesalia fasciata

(Lamarck, 1804: 217) from Eocene strata in France, Bel-

gium, and Pakistan (Cossmann and Pisarrro, 1910-1913:

Cox, 1930; Eames, 1952); Sigmesalia pagoda ( (Cox, 1930:

160-161, pl. 18, figs. Ga—b, 7a—b) from Eocene strata in

Pakistan; Sigmesalia biplicata (Bowles, 1939: 328, pl. 34,

figs. 6, 8) from Paleocene strata in Alabama; and Sigme-

salia gomin (Bowles, 1939: 326-327, pl. 33, fig. 9) from

Paleocene strata in South Carolina.

The protoconchs of Mesalia mesal and Sigmesalia

solida are very similar (compare Figures 5 and 15); both

are small, smooth, have essentially tie same shape, and

the transition to the teleoconch is gradual.

In summary, we found that the morphologic features

of the Eocene Paris Basin shells are variable. We could

find no reliable, constant morphologic characters to dis-

tinguish Mesalia from Sigmesalia; hence, we regard them

as congeneric.

Mesalia martinezensis (Gabb, 1869)

(Figures 16-23)

a martinezensis Gabb, 1869: 169-170, 228, pl. 25, fig.

1; Dickerson, 1914a: pl. 13, fig. 10; Waring, 1917: 100, pl.

7 fig. 5.

Turritella maccreadyi Waring, wes 783; Waring, 1915: fig.

[not fig. 14]; Waring 1917: S7-SS, pl. 12, fig. 10.

Mesalia maccreadyi (W ering), uedes-Mejia, 1989: 176-177,

pl. 3, figs. 3-6.

Mesalia martinezensis (Gabb).—Cossmann, 1912: 126; Stew-

art, 1927: 353-354, pl. 25, fig. 1; Schenck and Keen, 1940:

pl. 20, fig. 5; Merriam, 1941: 127-128, pl. 39, figs. 1-5, 7;

Zinsmeister, 1974: 11S-119, pl. 12, figs. 5, 9; Zinsmeister,

1983: pl. 2. fig. 14: Paredes-Mejia, 1989: 173-176, pl. 3,

figs. 7-LO; Saul, 1983a: text-fig. 2, pl. 1, fig. 2.

Mesalia clarki (Dickerson).—Zinsmeister, 1983: pl. 2, fig.

13.

Description: Large (up to approximately 95 mm

height). Tarritelliform: Pleural angle approximately 20°.

Protoconeh unknown. Teleoconch up to 12 whorls, in-

creasing rapidly in size from the apex. Suture slightly

impressed. Sculpture consisting only of spiral ribs of dif-

fering strength but dominate d by carina located anteri-

orly; ribs generally becoming stronge r with growth; spiral

threads on inte rspaces and on carina surface, Carina usu-

ally strongly angulate but can be rounded or even sub-

dued. Posterior to carina, several widely spaced spiri al

ribs of variable strength occur, ranging from tertiaries to

primaries: two ribs on uppermost spire, three to four on

upper spire and one to three on lower spire. Anterior to

carina, several spiral ribs of variable strength occur, rang-

ing from tertiaries to primaries: approximately five ribs

On uppel spire and one to two ribs (both occasionally

quite prominent) on lower spire. Last whorl with three

primaries, both posterior and anterior to carina. Base

with three secondaries, anteriormost one weak: ribs ob-

solete on short neck. Aperture relatively small, D-

shaped; columella relatively broad, smooth; spout effuse

and short with anterior end projecting slightly; growth-

line trace of last whorl (including base) par asigmoidal,

with lateral sinus flexure strongest in vicinity of carina.

Holotype:

mim.

ANSP 4344, height 57 mm, diameter 23

Type Locality:

not given).

Martinez, northern California (details

Geologic Range: Late early Paleocene to early late

Paleocene (near the Danian-Selandian bounde ury to early

Thanetian),

Distribution: DANIAN = Turritella peninsularis qua-

leyi Zone: lower San Francisquito Formation, Warm

Springs Mountain, Los Angeles County, southern Cali-

fornia (new stratigraphic occurrence, LACMIP. loc.

21581). NEAR THE DANIAN-SELANDIAN BOUND-

ARY = Turritella peninsularis qualeyi Zone transitional

with Turritella peninsularis Zone: Martinez Formation,

Herndon Creek east of Lower Lake, Lake County, north-

ern California (Stanton, 1896 [faunal list]; Dickerson,

1914a; Merriam, 1941); upper Las Virgenes Sandstone,

Simi Hills, Ventura County, southern California (Waring,

1917; Nelson, 1925 [faunal list]; Merriam, 1941; Zins-

meister, 1983; Saul, 1983a). PROBABLY NEAR THE

DANIAN-SELANDIAN BOUNDARY: Reworked

specimens in Santa Susana Formation, Poison Oak Can-

yon, north side Simi Valley, Los Angeles County, south-

erm California (new stratigraphic occurrence, LACMIP

loc. 21554); Reworked specimens in Stokes Canyon

Breccia Member of the middle Miocene Calabasas For-

mation, Stokes Canyon, Santa Monica Mountains, Ven-

tura County (new stratigraphic occurrence, LACMIP

loc. 25281). SELANDIAN = Turritella peninsularis

Zone: Lower Vine Hill Sandstone, Martinez area, Contra

Costa County, northern California (Weaver, 1953 [faunal

list]); lower San Fri mcisquito Formation, Finvou Ridge

east of Big Rock Creek, Valymero area, Ante lope Vv alley,

Los Ange sles County, southern California (Dichensen:

1914b anal list]; Merriam, 1941; Kooser, 1980 [faunal

list]); lower Santa Susana Formation (= “Martinez ma-

rine member” of Nelson, 1925 [faunal list]), Simi Hills,

Ventura County, southern California (Kew, 1923 [faunal

list]; Nelson, 1925 [faunal list]; Zinsmeister, 1983; Saul,

1983a). LOWER THANETIAN = Turritella infragranu-

lata Zone: Upper Vine Hill Sandstone, Martinez area,

Contr re . County, northern California (Weaver, 1953

; upper Santa Susana Formation, Palisade 2S

Highl: ads. inta Monica Mountains, Los Angeles

County, southern California (new stratigraphic occur-

rence, LACMIP locs. 7060 and 11717); Sepultura For-

mation, Mesa San Carlos, northern Baja California,

Mexico (Paredes-Mejia, 1989),

R. L. Squires and L. R. Saul, 2007

Page 9

Figures 16-23.

Discussion: The largest specimens of this species oc-

cur in the lower San Francisquito Formation, Pinyon

Ridge east of Big Rock Creek, Valymero area, Antelope

Valley, Los Angeles County, southern California.

There is considerable variability in the strength of the

spiral ribs on M. martinezensis. Most specimens are Cari-

nate on all whorls, including the last whorl. On some

specimens. however, the carina becomes weaker on the

later whorls as the other spiral ribs become stronger,

giving these whorls a convex shape (Figures 19-21), like

Mesalia martinezensis (Gabb, 1869). Specimens coated with ammonium chloride. 16-18. Hypotype LACMIP

13399, height 55 mm, diameter 25.4 mm. 16. Apertural view. 17. Oblique apertural view. 18. Abapertural view. 19. Hypotype

LACMIP 13400, LACMIP loc. 22557, apertural view, height 36.3 mm, diameter 14.1 mm. 20. Bhai LACMIP 13401, LACMIP

loc. 21607, abapertural view, height 38.3 mm, diameter 19.3 mm. 21. Hypotype LACMIP 13402, LACMIP loc. 22698, abapertural

view, height 33.9 mm, diameter 21.6 mm. 22. Hypotype LACMIP 13403, LACMIP loc. 26897, apertural view, height 10.6 mm,

diameter 6.5 mm. 23. Hypotype LACMIP 13404, LACMIP loc. 22330, base, diameter 17.2 mm.

shells misidentified by some workers as Turritella ma-

creadyi Waring, 1914.

The overall felecuantel morphology of the 10 mm-high

tip of Mesalia martinezensis superficially resembles that

of the 15-mm high mathildid Carinathilda diminuata

(Perrilliat, Vega, aaa Corona, 2004) illustrated by Kiel et

al. (2002: 329-330, fig. 2.4) from the lower Maastrichtian

of the Mexcala Formation, Guerrero, southern Mexico.

Carinathilda diminuata is definitely a mathildid because

it has a heterostrophic protoconch. The resemblance be-

Page 10

THE NAUTILUS, Vol. 121, No. 1

tween these two gastropods, nevertheless, provides evi-

dence that the Late Cretaceous mathildids and lower

Paleogene turritellids can have similar looking adult

shells.

Mesalia martinezensis resembles “Mesalia” virginiae

Stilwell et al. (2004: 29-30, pl. 5, figs. 6-10) from lower

Paleocene (Danian) rocks on Seymour Island, Antarctic

Peninsula, but M. martinezensis has a subtle effuse spout

rather than the longer and more distinct, twisted narrow

anterior canal that “M.” virginiae possesses. In addition,

M. martinezensis has stronger ribs and a parasigmoidal

growth line, rather than an opisthocyrt one on the last

whorl. In our opinion, the aperture of “M.” virginiae is

unlike that of Mesalia.

Gabb (1869) mentioned that the broadly expanding

whorl of martinezensis approaches that seen on Turri-

tella robusta Gabb (1864: 135, pl. 21, fig. 74; not = T.

(Haustator) robusta Grzybowski, 1899), but Merriam

(1941: 128) reported that the Late Cretaceous T. ro-

busta, which is represented by a single poorly preserved

type specimen, is probably not a Me salia. This type speci-

men has an umbilicus, therefore it is not a turritellid. It

is from the Redding area, northern California, and not

from Tuscan Springs, as erroneously reported by Mer-

riam (1941). Jones et al. (1975: pl. 1, fig. 19) identified

this specimen, which is of Turonian age, to be Glauco-

nia? robusta (Gabb, 1864).

Merriam (1941: 10, 116) stated that mainly in profile

the Pacific slope Miocene Turritella temblorenesis

Wiedey, 1928, might readily be confused with Mesalia

martinezensis. The latter also resembles the Pacific slope

Miocene Turritella temblorensis tritschi Hertlein, 1928,

and Turritella ocoyana Conrad, 1857. The latter, how-

ever, has a different growth line. In addition, T. martin-

ezensis strongly resembles Turritella fredeai Hodson,

1926, of Miocene age from northern Colombia and

northern Venezuela. None of these above-mentioned

Miocene species, however, has the effuse spout of Me-

salia.

Mesalia clarki (Dickerson, 1914a)

Mores 24-39

(Figures 24—32)

Mesalia clarki (Dickerson).—Merriam, 1941; 128, pl. 39, fig. 6

Zinsmeister, 1983: table 1, pl. 2, fig. 14

Turitella {sic| clarki Dickerson, 1914a: 142-143, pl. 13, fig. 8.

Description:

mm height). Turritelliform. Pleural angle approximately

21 to 22°. 12

whorls, consisting of two whorl shapes: flatish rounded

Medium small (up to approximately 31

Protoconch unknown. Teleoconch up to

and anteriorly aneulate. Sutural area indente - Sculpture

consisting only of spiral ribs, variable in number,

strength, and spacing. Flattish to rounded shen shape:

upper spire aah one or two secondaries on posterior part

and two (bicostate) primaries on anterior part; lower

spire and last whorl with seven to eight nearly equal

strength primaries (anteriorly located ribs can be some-

what ‘angulate); spiral threads on all interspaces; poste-

riormost rib part of broad band; ribs on anterior part of

whorl tend to be slightly stronger than posteriorly located

ribs: base of last whorl with thee ribs. Angulate whorl

shape: upper spire with one secondary on posterior part

and two (bicostate), well dev eloped, f lat-topped prima-

ries on anterior part; lower spire and last whorl with

three primaries on posterior part and two stronger pri-

maries, with one secondary in between each, on anterior

part: spiral pas on all interspaces. Base (including

short neck) of last whorl with approximately seven,

evenly ied ies interspaces and ribs covered by spiral

threads. Aperture small; columella narrow with thin cal-

lus, occasionally with single, weak fold, slight twist on

anterior end of columella. Spout effuse, short, and nar-

row. Growth-line trace on last whorl (including base)

parasigmoidal, with lateral sinus flexure strongest medi-

ally.

Holotype: UCMP 11936, height 25 mm, diameter

16.5 mm.

Type Locality: UCMP loc. 1540

Geologic Range: Late Paleocene = Turritella infra-

granulata Zone.

Distribution: “Martinez” Formation, northeast side of

Mount Diablo, Contra Costa County, northern Califor-

nia (Dickerson, 1914a; Merriam, 1941; Zinsmeister and

Paredes-Mejia, 1955 [faunal list]; upper Santa Susana

Formation, Trailer and Quarry canyons, Los Angeles

County, Santa Monica Mountains, southern California

(Strathearn et al., 1988 [faunal list]; Squires and Saul,

1998: 1025).

Discussion: Mesalia clarki is abundant in the upper

Santa Susana Formation at LACMIP loc. 10508, in the

Santa Monica Mountains, Los Angeles County, southern

California. The anterior ends of the shells are very frag-

ile, and nearly all the specimens have incomplete aper-

tures. None of the specime ns has retained their proto-

conch, and most specimens are missing their upper spire.

Growth lines are hard to discern, usually visible only on

a single whorl (typically the penultimate whorl), and

were rarely preserved on the base of the last whorl. Some

of the specimens appear to have a wider pleural angle

(23°) than normal, but these particular specimens have

been crushed. A few of the specimens (five percent) have

naticid boreholes, and a few other specimens are en-

crusted, in part, by bryozoans. Rare spe cimens are

coated by calcareous algae.

All previous workers assigned Mesalia clarki to various

genera without knowledge of the shape of the aperture.

Our cleaning of representative specimens of Dickerson’s

species revealed it to have a short, shallow effuse spout

(Figures 24-25) and bicostate sculpture on the juvenile

whorls (F igure 30). There is considerable variation in the

sculpture and the shape of the whorls on M. clarki. Some

R. L. Squires and L. R. Saul, 2007 Page 11

28 be 20

Figures 24-32. Mesalia clarki (Dickerson, 1914). Specimens coated with ammonium chloride. All from LACMIP loc. 10508. 24.

Hypotype LACMIP 13405, apertural view, height 21.5 mm, diameter, 19.6 mm. 25. Hypotype LACMIP 13406, slightly oblique

apertural view, height 23.1 mm, diameter 10.8 mm. 26-27. Hypotype LACMIP 13407, height 22.6 mm, diameter 9.4 mm. 26,

Abapertural view. 27. Oblique apertural view. 28. Hypotype LACMIP 13408, apertural view, height 23.2 mm, diameter 9.2 mm. 29.

Hypotype LACMIP 13409, abapertural view, height 21.5 mm, diameter 11 mm. 30. _Hypotype LACMIP 13410, abapertural view,

height 23.6 mm, diameter 9.7 mm. 31. Hypotype LACMIP 13411, base, diameter 9.5 mm. 32. Hypotype LACMIP 13408, base of

same specimen shown in Figure 28, diameter 8.9 mm.

specimens have nearly uniform sculpture and flattish Zinsmeister (1983: pl. 2, fig. 14), Zinsmeister and

whorls (Figure 24), others have carinate whorls (Figure Paredes-Mejia (1958: table 1), ), and Paredes- Mejia (1989:

26). whereas others have uniform sculpture with convex table 3) reported M. clarki from the Santa Susana For-

whorls (Figure 29). mation in the Simi Hills, southern California. These re-

Page 12

THE NAUTILUS, Vol. 121, No. 1

ports, however, were based on the misidentification of a

specimen of Mesalia martinezensis that happens to lack a

strong anterior carina on the otherwise convex whorls.

Mesalia clarki resembles Motyris aralica

(Michailovski, 1912; Wenz, 1939: 652, fig. 1856) from

upper Eocene rocks in the Aral Sea region. Motyris

Eames, 1952, was formerly known as Tomyris

Michailovski, 1912. See Marwick (1957: 162-163) for

more taxonomic information pane Motyris. Me salia

clarki differs from M. aralica by not having tabulate

whorls with strongly indented sutures. The full aperture

of M. aralica is enlnowel, and details about its apical

whorl development are wanting. The only other species

of Motyris that we are aware of is gee ‘is pseudoaralica

Eames (1952: 30-31, pl. 1, fig. 15; pl. 2, figs. 58a, b) from

Pakistan, but its aperture is unknown. We believe that

when the great variability of Mesalia is taken into ac-

count, Motyri ‘is will prove to be congeneric.

ACKNOWLEDGMENTS

Earl Brabb (U. S. Geological Survey, Menlo Park) pro-

vided very useful information regar ding the stratigraphy

of the beds in the vicinity of the type iscality of Me salia

clarki. Lindsey T. Groves (LACM, Malacology Section)

kindly provided key literature dealing with the ecology of

modem Mesalia. The manuscript benehied from the re-

views by Warren D. Allmon (Paleontological Research In-

stitute, Ithaca, New York) and Steffen Kiel (University of

Leeds, England and Department of Paleobiology, Smith-

sonian Institution’s National Museum of Natural History).

Steffen Kiel also provided us with an important hard-to-

find reference and very useful stratigraphic information.

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APPENDIX

LOCALITIES CITED

Localities are LACMIP, unless otherwise noted. All

quadrangle maps are U. S. Geological Survey maps.

7060. Elevation 1427 ft., on ridge between Temesal and Santa

Ynez canyons at edge of fire road on top of ridge, Topanga

Canyon ‘Quadrangl e (7.5 minute, 1952, photorevise od

1981), Los Angeles County, southern California. Pale-

ocene. Santa Susana Formation. Coll.: H. D. B. Wilson,

June 1, 1941.

10508. North slope of Trailer Canyon near top of ridge be-

tween Quarry and Trailer canyons at approximately 1325

ft. elevation and just west of saddle, just below coralline-

algal beds in limy siltstone west of small f fault, road cut

north side of unpaved road 5600 ft. north of San Vicente

y Santa Monica Grant boundary, 10,400 ft. east of Los

Angeles City boundary, Topanga Quadrangle (7.5 minute,

1952, photor vised 1981), east of Santa Ynez ¢ Janyon, Pali-

sades Highlands, Santa Monica Mountains, Los Angeles

County, southern California. Lower upper Paleocene

(lower Thanetian). Santa Susana Formation. Coll.; ¢

Strathearn and others, fall, 1982.

11717. Float at about 1600 ft. elevation in bottom of south-

flowing gully joining Quarry Canyon at about 1410 ft. =“

evation: 1500 ft. SW of hill 2036, Topi nga Quadrangle (

minute, 1952, photorevised 1981), Los Angeles C a

southern C ali tonmie Paleocene. Santa Susana Formation.

Coll J.M Alderson, November 11, 1980.

21581. Black nodular shale and conglomerate on road 1.1 mi.

east from Cienaga Camp at Fish Canyon forks toward

Warm m Springs summit: on northwest side of ravine: north

side East Fork Fish Canyon, T. 6 N, R. 16 W, approximately

at. ft. north, 750 ft. east of bench mark 2205, Warm

Springs Mountain Quadrangle (7.5 minute, 1958, photo-

revised 1974), Los Angeles County, southern California. Pa-

leocene ( (upper Danian). San Francisquito Formation. Coll:

R. W. Webb and E. H. Quayle, jane 1941,

21607. South 1/2, SE 1/4, section 25, T. 2 N, R. 18 W, Cala-

basas Quadrangle (7.5 minute, 1952, photorevised 1967),

Ventura County, southern California. Lower upper Pale-

ocene (lower Thanetian). Santa Susana Formation. Coll:

Kinney and Sherman, date unknown.

21554. Reworked fossil boulders in conglomerate, in second

spur canyon off Poison Oak Canyon east of Las Llajas

Canyon, 2800 ft. up canyon (north) from Poison Oak Can-

yon; on west slope 25 ft. above bottom of canyon, Santa

Susana Quadrangle (7.5 minute, 1951, photorevised 1969),

north side Simi Valley, Ventura County, southern Califor-

nia. Paleocene. Santa Susana Formation, Coll.; P. L. Gold-

man, date unknown.

22330, Beds cropping out on nose of spur on west side of Meier

Canyon, approximately 600 ft. north of second “n” in

“Meier Canyon,” Santa Susana Quadrangle (7.5 minute,

1951, photorevised 1969), south side of Simi Valley, Simi

Hills, Ventura County, southern California. Lower middle

Paleocene (Selandian) = Turritella peninsularis Zone.

Santa Susana Formation, “Martinez marine member.”

Coll.: W. P. Popenoe, April 3, 1946.

22557. Sandstone bed below small waterfall [dry] west of road

going south through Barclay Ranch, 10,162 ft. south and

5660 ft. wet of junction of Souther Pacific railroad and

Los Angeles Ave. about 0.25 mi. east of Santa Susana,

Santa Susana Quadrangle (7.5 minute, 1951, photorevised

1969), Simi Hills, Ventura County, southern California.

Paleocene. Santa Susana Formation. Coll: M. Murphy,

spring, 1950.

22698. On first large ridge; trending southwest to west of ridge

trending south of hill 2150. Bearing from the nioihowest

corner of the Calabasas Quadrangle is S14°E; distance

12.210 ft., Calabasas Quadrangle (7.5 minute, 1952, pho-

torevised 1967), Simi Hills, Ventura County, southern

California. Paleocene. Santa Susana Formation. Coll.: J. H.

Fantozzi, June 1, 1951,

25281. Sandstone at elevation of 1000 ft., about 400 ft. south

aiid LO0O0 ft. west of northeast corner of section 5, T. 1S,

R. 17 W, Malibu Beach Quadrangle (7.5 minute, 1950,

photorevised 1967), on west side of northern tributary to

Stokes C anyon, western Santa Monica Mountains, Los ‘Ane

geles County, southerm California. Reworked Paleocene

(Selandian) fossils in middle Miocene Calabasas Forma-

tion, Stokes Canyon Breccia Member. Coll.: J. Stark and

T. Susuki family, May 5, 1965.

26897. Gully west side of Temesal Canyon opposite 2nd ‘e’ of

Temesal at about 1475 ft. elevation; approximately 1082 m

(3550 ft.) south; 533m (1750 ft.) east of hill 22036; San

Vincente and Santa Monica Grant, Topanga Quadrangle

(7.5 minute, 1952, photorevised 1967), Santa Monica

Mountains, Los Angeles County, southern California.

Middle upper Paleocene (middle Thanetian). Santa Su-

sana Formation, Coll: ]. M. Alderson, March 9, 1980.

UCMP. 1540. Elevation 1000 ft., 1 mi. south of Stewartville

(site), northeast corner of NW 1/4, section 15, T. 1 N,

R. 1 E, Antioch South Quadrangle (7.5 mimute, 1973, pho-

torevised), 300 ft. south of basal Tejon conglomerate and

600 ft. north of Chico-Martinez contact, northeast side of

Mount Diablo, Contra Costa County, northern California.

Upper middle Paleocene (Selandian) = Turritella infra-

granulata pac hecoensis ae “Martinez” Formation,

lower member. Coll: | ). Dickerson, circa 1912.

THE NAUTILUS 121(1):17-28, 2007

Page 17

Vertigo malleata, a new extreme

alcifuge land snail

(Gastropoda: Vertiginidae) from the Atlantic and Gulf coastal

plains of the USA

Brian F. Coles

Mollusca Section, Department of

Biodiversity

National Museum of Wales, Cathays Park

Cardiff, CF10 3NP ,

GREAT BRITAIN

[email protected]

Jeffrey C. Nekola

Department of Biology

University of New Me xIiCO

Albuque rque, NM $7131 USA

[email protected]

ABSTRACT

Vertigo malleata new species is an extreme calcifuge land snail

widely distributed in the Atlantic and Gulf costal plains of the

eastern USA. This species appears to have gone undetected

because of its small size and restriction to low pH = sites—

—— bogs, Atlantic white cedar (Chamaecyparis as S

L.) BSP) swamps, pocosins, and pine woodlands—which,

has been assumed, harbor little or no molluscan diversity. Ver r

tigo malleata is distinguished from other members of the genus

by the strongly pustulose surface of the body whorl, which gives

the shell a malleated appearance at low to moderate mé vonifi-

cation. While the major apertural lamellae/folds (parie tal, col-

umellar, and palatal) of this species are typical for Vertigo, the

strongly pustulose shell sculpture, occurrence of an infrapari-

etal lamella, and frequent development of subcolumellar and

basal lamellae in the absence of an angular lamella appear

unique. Although V. malleata is an abundant snail within its

range, the common use of short-return fire regimens to manage

forests of the eastern USA appears to be artificially limiting its

distribution to wet, less frequently bumed sites.

Additional key words: Bothriopupa, Nesopupa, biogeography,

fire ecology, community ecology, eastern North America

INTRODUCTION

Acidic and lime-poor habitats have long been thought to

support depauperate molluscan community abundance

and richness (Boycott, 1934; Baker, 1939; Kerey and

Cameron. 1979; Burch and Pearce, 1990). Consec quently,

little molluscan survey work has been attempted in acidic

sites even though they can re present a substantial frac-

tion of the landscape. However, such areas should not be

ignored for terrestrial gastropod biodiversity because

base-poor habitats can be as speciose as base-rich habi-

tats on a per-individual basis (Schilthuizen et al., 2003;

Corresponding author: Jeff Nekola

email: [email protected]

Pokryszko and Cameron, 2005), and because some spe-

cies like the European Vertigo ronnebyensis (Wester-

lund, 1871) and Zonitoides excavatus (Alder, 1830) are

restricted to or more frequent in bea sites (Kemey

and Cameron, 1979),

During land snail studies in eastern North America

(Nekola, 2002a; Nekola and Coles, 2004; Coles and

Nekola, unpublished data) we found that acidic habitats

often supported substantial populations of land snail taxa

that have been little reported since their original descrip-

tions; e.g., Vertigo cristata (Sterki, 1919), Vertigo nylan-

deri Sterki, 1909, Vertigo alabamensis Clapp, 1915, and

Vertigo perryi Sterki, 1905. In tact, V. alabamensis and

V. perryi were each previously known from only two sites

worldwide (Pilsbry, 1948: Hubricht, 1985). In the course

of these acid-habitat surv eys, we examined Saco Heath,

an undisturbed domed ombotrophic Sphagnum bog in

the Atlantic coastal plain of York County, Maine. At this

site we located a species of the genus Vertigo that was

strikingly different from all previously inoue taxa. This

form was subsequently found to represent the most com-

mon land snail of highly acid, mesic to wet habitats of the

Atlantic and Gulf coastal plains of the eastern USA.

Here, we describe this taxon as Vertigo malleata, new

species, document its biogeography and ecology, and

briefly discuss its relevant conservation issues.

MATERIALS AND METHODS

Site Selection: Approxiini itely 130 sites were surveyed

along the Atlantic and Gulf coastal plains of the eastern

USA from Maine to western gba i, including penin-

sular Florida south to Gainesville. These sites encom-

passed the entire soil base-status and moisture gradient

of the region and covered a total geographic extent of

2400 kin. Thirty sites represented base-neutral to base-

rich habitats (i.e.,

forests, and limestone outcrops), while the remaining

were base-poor (2.e., pine barrens, pine-wiregrass Sda-

calcareous wetlands, marl banks, rich

Page 18

THE NAUTILUS, Vol. 121, No. 1

vanna, heaths, Atlantic white cedar swamps, bay forest,

Sphagnum bogs, and pocosins).

Field Methods: Latitude and longitude of each site

was determined using a hand-held GPS. Terrestrial gas-

tropod faunas were documented from a representative

100-1000 m2 area within each site by hand collection of

larger taxa and litter sampling for smaller taxa. Litter

sapling was used as the primary method of collection

because it provides the most complete assessment of site

faunas (Oggier et al., 1995; Cameron and Pokryszko,

2005). As suggested by Emberton et al. (1996), collec-

tions were medé at places of high micro-mollusk density

such as loosely compacted leaf litter lying on top of

highly compacted damp soil or humus. This loose litter

was removed by hand and aggressively sieved in the field

using a shallow sieve of 2 mm mesh nesting loosely inside

a sieve of 0.6 mm mesh. The procedure consisted of

throwing handfuls of litter onto the coarser mesh accom-

panied by vigorous shaking, tapping, or other agitation.

The process was continued for 15-60 minutes, a time

interval that yielded 50-500 ml of fine material (0.6—2.0

mm). In general, sites were sampled in parallel (but in-

dependently) by each of the authors, although several

sites were sampled by only one worker (see Table 1)

Laboratory Procedures: Samples were slowly and

completely ‘dried at room te mperature and then passed

through an ASTME #30 sieve (0.6 mm mesh) with frac-

tions be sing hand-picked against a neutral background.

All shells, shell fragments, and slug plates were removed,

and all identif fable shells from each site were assigned to

species using the authors’ reference collections and vari-

ous museum collections (see below). The total numbers

of shells per species per site were recorded, as were the

number of unidentified immature individuals.

Comparisons: The new species was compared with

specimens of all eastern North American and western

Eurasian species of Vertigo, and to representative taxa in

the related genera Nearctula of western North America,

Nesopupa of the Old-World tropics, and Bothriopupa of

the neotropics. Comparative material consisted of the

authors’ extensive reference collections, the collections

of the gees Museum of Natural History, Gainsville,

): the Field Museum of Natural History, Chi-

cago, . (FMNH), the Carnegie Museum of Natural

History, Pittsburgh, PA (CM); the

Wales—Zoology, Cardiff, U K (NMW.Z): the Natural His-

tory Museum, London, UK, and the Queensland Mu-

seum, Brisbane, Australia. Additional comparisons were

made with material presented by Pilsbry (1920; 1948).

Imaging:

ing a digital camera attached to a stereomicroscope. Ap-

proximately 12 separate 1388 x 1040 pixel images were

made of each specimen with the image focal lengths

Shells were imaged at 40x magnification us-

positioned at 120%m increments from the front to back

of the shell. CombineZ5 freeware (ht ttp:/Avww. hadleyweb

p.blueyonder.co.uk/CZ5/ combinez5.htm) was used

see ese

National Museum of

to assemble a final image from the well-focused parts of

each separate image, The body whorl surface of the new

taxon was also imaged at 150x with 60 images positioned

at 5 jum focal length increments and assembled into a

single image using CombineZ5. These separate images

were imported into Adobe Photoshop, where brightness

and contrast were optimized and the background made

uniformly black. These images were then compiled into

a single plate.

Community Ecology: — Analysis of co-occurring terres-

trial gastropod species and abundance was determined

using data for sites sampled by the second author (ie., all

. s with accession numbers prefixed by JON in Table

). These analyses were performed for the whole dataset

of 49 discrete sites and also by geographic sub-region—

New England (Maine, Massachusetts), New Jersey, the

North and South Carolina coastal plain, and the Gulf

coastal plain—to allow for documentation of composi-

tional gradients across the range of the new species. The

physical habitat and plant community from each site was

also noted.

Nomenclature: Taxonomic nomenclature follows that

of Turgeon et al. (1998) with updates from Nekola

(2004). Apertural lamellae and fold nomenclature follows

that of Pilsbry (1945: 869, fig. 469), i.e., parietal “teeth”

are referred to as “folds” and all other “teeth” are termed

“lamellae”, whatever their form.

SYSTEMATICS

Class Gastropoda

Subclass Pulmonata

Order Stylommatophor l

Family Vertiginidae

Genus Vertigo Miiller, 1773

Vertigo malleata new species

(Figures 1-15, 20, Tables 1-2)

Diagnosis: Minute; shell ovoid, similar in size and

shape to Vertigo ventricosa (Morse, 1565) but. distin-

cuished by malleated appearance of the body whorl at

low to moderate (10-40x) magnification; upper whorls

fine ‘ly rib-striate, minutely decussated by spiral lines; ap-

erture with parietal and columellar lamellae, a small in-

fraparietal lamella (occasionally absent), and two palatal

folds; one or more subcolumellar-basal lamellae usually

present; angular lamella absent.

Description: Shell 1.5—2.1 mm tall x 1.25-1.4 mm

wide (holotype 1.98 x 1.36 mm), ovoid to ovoid-conical,

inflated, approximately 44.5 whorls, with deep suture;

translucent, olive-ye low to brown in color; body whorl

approximately 66% of total he ight. Protoconch and

neanic whorls minutely papillose pre fine spiral stria-

tion; subsequent whorls finely rib-striate: striae most dis-

tinct on penultimate whorl where they are minutely de-

cussated by fine spiral lines; on body whorl the se ulpture

degenerates into an irregularly pustulose surface (Figure

B. F. Coles and J. C. Nekola, 2007

Page 19

4) which at low to moderate (10-40x) magnification

takes on a malleated appearance as it appears hammered

with small depressions; behind the aperture the sculp-

ture takes the form of coarse, irregular rib-striae (Figures

2. 11). Aperture rounded, approximately 40% of shell

height; lip reflexed but not thickened, peristome usually

dark blackish-olive: sinulus moderate-weak: basally the

aperture abruptly inflates to form a rounded swelling,

but not a distinct crest (Figures 2, 8). Umbilicus closed

(Figure 3). Aperture typically with six lamellae and folds

(Figures 1, 5, 7, 9, 12-15): a strong, slightly sinuous pa-

rietal lamella (Figures 1,5, 9, 15); a shelf-like columellar

lamella that spirals internally around the columella for

approximately one whorl; two palatal folds of approxi-

mately equal length that extend approximately 0.2 whorls

into body whorl, the lower slightly more immersed than

the upper (Figures 1, 5-7, 10, 15), both highest at mid-

length (Figures 1, 5, 6, 15); a nodular infraparietal

lamella usually present (Figures 1,5, 9, 12-15), occasion-

ally vestigial or absent (Figures 6, 10); angular lamella

absent: presence of a nodular subcolumellar lamella and

nodular subcolumellar-basal lamella variable (Figures 1,

6, 7, 9, 10). Apertural ends of the palatal folds coincide

with abrupt inflation of basal aperture (Figures 7-8), in

consequence appearing to be raised on a weak crest

when viewed within the aperture but not associated with

any internal shell thickening; externally shell only slightly

impressed over palatal folds. Body of animal grey vat

several organs of a brown or cream color visible through

the upper whorls of shell. All dissected individuals bias e

proven to be aphallic (Beata Pokryszko, personal com-

munication), hence the genitalic anatomy is unknown.

Holotype (Figures 1-4): NMW.Z.2005.011.03830,

USA North Carolina, Pender County, Holly Shelter

Game Land, Brian Coles, 1 April 2003.

peas (Figures 5-15): || NM\W.Z.2005.011.03531-

03839, figured material, see Figure legends for details;

NMW.Z.2005.011.02118-02120, approximately 5100 in-

dividuals (split into three approximately equal lots) from

type locality; UF 348143, approximately 700 individuals

from type locality; CM 73971, 143 individuals from type

locality; NMW Z,.2005.011.02597, 90 specimens, Wells

Heath, York County, Maine (43°20'2” N, 70°38'24” W),

Brian Coles: NMW.Z.2005.011.02591, 26 specimens,

Skunknett Audubon Preserve, Barnstable County, Mas-

sachusetts (41°38'59” N, 70°22’31” W): NMW.Z.2005.

011.02585, 170 specimens, Peterson Swamp Wildlife

Management Area, Plymouth County, Massachusetts

(42°0'37" N, 70°49'4” W), Brian Coles: NMW.Z.2005.

011.02514, 122 specimens, Stafford Forge Wildlife Man-

agement Area, Ocean County, New Jersey (39°42'44" N,

74°22'10" W), Brian Coles: NMW.Z.2005.011.02197,

250 specimens, Lewis Ocean Bay Preserve, Horry

County, South Carolina (33°47'16" N., 78°50'56" W.),

Brian Coles; NMW.Z.2005.011.03035, 42 specimens,

Collins Bay, Ware County, Georgia (31°5'12” N.,

§2°36'56" W.), Brian Coles: NMW.Z.2005.011.03065,

107 specimens, Wilma Station, Liberty County, Florida

(30°9'34" N., $4°57'39" W.), Brian Coles; NMW.Z.2005.

011.03079, 162 specimens, Pond Creek, Conecuh Na-

tional Forest, Covington County, Alabama (31°6'12" N.,

86°32'3" W.), Brian Coles.

Type Locality: Holly Shelter Game Land (34°31'57"

N, 77°44'41” W), Pender County, North Carolina, USA;

ae dense scrub of mesic bay/pine forest at pocosin

margin, individuals sieved from deep bracken fern and

pine needle litter, collected by Brian Coles, 1 April 2003.

Other Material (Table 1): Sixty additional lots col-

lected by Brian Coles are deposited i in the Coles Collec-

tion of the National Museum of Wales. Fifty three lots

representing 3133 individuals collected by Jeff Nekola

are deposited in the Nekola collection (JCN).

Etymology: The specific name malleata refers to the

hammered appearance of the body whorl at low to mod-

erate magnification.

Variation: Vertigo malleata was rather constant in

general appearance along its 2400 km range, although

some variation in shape, size, color, sculpture, and de-

velopment of the apertural lamellae was observed. Varia-

tion in size and shape has been noted above. In addition,

the most southern populations (Georgia, Alabama, and

Florida) tended to be darker in color and showed the

most strongly developed shell sculpture (Figures 9-12).

While the parietal lamella, columellar lame: and the

palatal folds varied little, the infraparietal lamella varied

from strong (Figures 1, 5, 19) to weak (Figure 9) to

occasionally warty (Figures 6,10). The eae

and nodular basal lamella although usually distinct (Fig-

ures 1, 5, 7, 13) were also occ: isionally absent ( Figure 5).

Multiple subcolumellar-basal lamellae of eS place-

ment were also noted most frequently in Gulf Coast

populations (Figures 9, 10). However, such trends were

not distinct enough to support the designation of geo-

graphical races, w ith most of this observ ed morphological

variation occurring within local regions or populations.

Comparison with Other Species of Vertigo and of

Related Genera: Vertigo malleata differs from all

other Vertigo species by its strongly pustulose body

whorl sculpture and possession of an infraparietal and

subcolumellar-basal lamellae while lacking an angular

lamella. Because of these unusual characteristics, we do

not feel assignment of this taxon to a particular subgenus

to be prudent at this time. Additional data, possibly

based on DNA sequence information, will be required to

accurately determine its closest relatives.

On casual inspection, Vertigo malleata could be taken

for a member of the V. gouldii group (e.g. Vertigo cris-

tata; see Pilsbry, 1948: 958, figs. 4, 5, 8; 967, figs. 1-16)

because of its shell color, striated upper whorls, and silky

luster. Like V. malleata, V. cristata has four prominent

lamellae and strong striation on the penultimate whorl

(Pilsbry, 1948: 967, figs. 4-5, 973, fig. 520; Nekola, 2001)

Page 20

THE NAUTILUS, Vol. 121, No. 1

Table 1. Vertigo malleata: sites, brief habitat descriptions, collection dates, accession numbers, and total number of specimens

taken.

Site

State/County # Site: Habitat! Coordinates Date Accession Nufmber Specimens

Alabama

Covington 1 Pond Creek seep (Conecuh — 31°06'12" N, May 5 2005 NMW.Z.2005.011.03079 162

NF); Tlex-Smilax-bay 86°32'03" W JON 12365 LU

scrub on seep margin

2 Moccasin Branch (Conecuh 31°06'42" NJ May 5 2005 JCN 12371 3

NF); old pine-bay-heath 86°35'53" W

forest

3 Bear Bay (Conecuh NF): 31°6'29" N, May 5 2005 = NMW.Z.2005.011.03068 ]

heath-dominated 86°38'54" W

scrub on wetland

margin

Mobile 4 Grand Bay Forever Wild 30°25'07" N, May 1 2005) NMW.Z.2005.011.03019 3

Preserve; wet bay and $8°19'35" W

mixed forest

Florida

Columbia 5 Impassable bay (Osceola 30°23'31" N, Jan § 2005 NMW.Z.2005.011.02849 740

NF WMA); wet holly-bay — $2°30'05" W May 2 2005 JCN 12280 71

scrub

6 Osceola National Forest 30°22'30" N, May 2 2005) NMW.Z.2005.011.03026 213

WMA; wet Pinus- 82°32'04" W JON 12285 75

Lyonia-Vaccinium

savanna

7 ~~ Osceola National Forest 30°22'39" N, Jan § 2005 NMW.Z.2005.01 1.02845 38

WMA: wet Acer- 82°31'42" W May 2 2005 | NMW.Z.2005.011.03024 23

Taxodium-Lyonia forest May 2 2005. JCN 12283 6

Leon S Wolf Trap Bay 30°22'04" N, Jan 7 2005 NMW.Z.2005.011.02513 ~100

(Apal: achicola NF); tall S4°34'11" Wo May 4 2005) NMW.Z.2005.011.03054 132

pine-holly-bay forest May 4 2005 JCN 12324 37

9 Wolf T rap Bay 30°21'46" N Jan 7 2005 NMW.Z..2005.01 1.02516 44

(Apalachicola NF): $4°34/23" Wo May 4 2005 NMW.Z.2005.011.03050 7

wet-mesic pine-holly- May 4 2005 JCN 12321 1]

heath forest

10 Otter camp (Apalachicola 30°20'20" N, Jan 7 2005 NMW.Z.2005.011.02520 ~50

NF); regenerating mesic S4°36'41" Wo May 4 2005) NMW.Z.2005.011.03056 4

pine-holly heath

Liberty 1] Wilma Station: mesic, old 30° °09'34" N, May 4 2005 = NMW.Z.2005.011.03065 127

pine-magnolia-bay forest °57'39" W JON 12344 30

12 Juniper Creek Islands 30°03'15" N, May 4 2005) NMW.Z.2005.011.03062 65

(Apalachicola NF); old 4°45'40" W JCN 12337 41

growth pine-holly-bay

forest

13° Juniper Creek Islands 30°04'46" N, May 4 2005) NMW.Z.2005,011.03059 5

(Apalachicola NF); S4°45'41" W JON 12333 75

white cedar-pine-holly

forest

I4 Juniper Creek Islands 80°02'07" N, May 4 2005 NMW.Z.2005,011.03064 ~40

(Apalachicola NF); 84°49'38" W JON 12539 4S

pine-red maple-white

cedar forest

15 Carr Bridge (Apalachicola 30°07'26" N, May 4 2005. JCN 12340 12

NF); wet-mesic Ilex 84°53/31"” W

forest

Wakulla 16 South of Otter Camp 30°16'55" N May 4 2005) NMW.Z.2005.011.03057 69

(Apalachicola NF); $4°36'54" W May 4 2005 JCN 12327 53

wet-mesic pine-holly

forest

17 W Branch Sopchoppy R 30°15/05" No May 4 2005 NMW.Z.2005.011.03058 5

(Apalachicola NF); $4°37'30" W JON 12328 36

pine cy press-bay-holly

forest

B. F. Coles and J. C. Nekola, 2007

Page 21

Table 1. Continued

Site

State/County # Site; Habitat! Coordinates Date Accession Number Specimens

Georgia

Ware IS Collins Bay; wet holly-wax 31°05'12" N, May 2 2005 NMW.Z.2005.011.03035 42

myrtle-bay forest §2°36'56" W JON 12300 58

19 Dixon State Forest; wet 31°05'36" N, May 3 2005 JON 1230] 4

Pinus-Gordonia forest 82°16'13" W

20 Dixon State Forest; 31°06'49" N, May 3 2005 NMW.Z.2005.01 1.03035 ]

wet-mesic Quercus- 82°16'16" W JON 12302 2

Ilex-Gordonia forest

Maine

York 2) Saco Heath 1 (TNC): sedge 43°32'42" N Oct 14 2002 NMW.Z.2005.01 1.01550 3

and heath litter on 70°28'33" W Aug § 2004 NMW.Z.2005.01 1.02567 l4

Sphagnum bog NMW.Z.2005.011.02577 52

JON 12092 10

JON 12099 88

JON 12101 3

Oct O1 2004 NMW.Z.2005.011.02614 55

NMW.Z.2005.011.02616 40

22 Saco Heath 2 (TNC); low 43°32'50" N, Aug OS 2004. NMW.Z.2005.011.02571 45

forest with Carex 70°27'32” W JEN 12095 59

groundcover ;

23. Wells Heath (TNC); under 43°20'02" N, Oct 01 2004. NMW.Z.2005.011.02597 90

heath scrub on 70°38'24" W

Sphagnum bog

Massachusetts

Barnstable 24 Skunknett Audubon 41°38'59" N, Aug 13 2004. NMW.Z.2005.011.02591 26

Preserve 2; Myrica- 70°22'31"” W JCN 12180 22

Chamaecyparis bog

margin

Bristol 25 Noquochoke WMA: 41°39'35" N, Aug 12 2004. NMW.Z.2005.011.02592 2)

Chamaecyparis-Cryilla T1°O1'07" W JON 12168 17

swamp forest

26 Noquochoke WMA: 41°39'39" N, Aug 12 2004 NMW.Z.2005.01 1.02590 4

leatherleaf island in acid 71°01 12" W JCN 12164 12

sedge fen

27 Pine Swamp Brook; 41°55'57" N, Aug 10 2004. NMW.Z.2005,011.02586 9

leatherleaf fringe of open 71°03'49" W JCN 12149 5

acid bog

Plymouth 28 Peterson Swamp WMA; 42°00'37" N, Aug 10 2004. NMW.Z.2005.011.02585 ~170

Chamaecyparis-Acer 70°49'04" W JON 12145 137

rubrum forest

Worcester 29. Tom Bog; Sphagnum bog 42°30'46" N, Oct 05 2004. NMW.Z.2005.011.02760

with serub 72°12'43”" W NMW.Z.2005.011.02761 ~170

New Jersey

Atlantic 30 Park Road (Wharton SF): 39°42'58" N, May 22 2004 = NMW.Z.2005,011.02516 18

moist Ilex-Gaylussacia- 74°44'10" W JON 12050 54

Kalmia scrub

Burlington 31 Swan Bay WMA; low 39°35'13" N May 20 2004 =NMW.Z.2005.011.02479 107

Nyssa-llex-. Acer rubrum 74°30'50" W JEN 11983 103

forest

32 Lebanon State Forest; open — 39°52’28" N, — May 19 2004. =NMW.Z.2005.011.02469 2]

heath-Smilax scrub 74°30'57" W May 21 2004 JCN 12026 l4

33. Roberts Brook; low Nyssa- 39°47'07" N, May 19 2004 =NMW.Z.2005.011.02466 20)

Chamaecyparis-heath 74°39/26" Ws May 21 2004. JCN 11989 5

forest

34 Brendan T Byrne State 39°53'07" N, May 22 2004. NMW.Z.2005.011.02499 |

Forest: dry 74°30'22" W

Chamaecyparis-bog

Camden 35 2 miles WSW of Delette: 39°46'32"” N, May 19 2004 = NMW.Z.2005.011.0246] 6

moist bank with pine, 74°48'21" W May 21 2004 = NMW.Z.2005.011.02454 28

oak, wax myrtle 8

JON 11995

THE NAUTILUS, Vol. 121, No. 1

Table 1. Continued

Site

State/County # Site; Habitat! Coordinates Date Accession Number Specimens

Gloucester 36 Winslow WMA: heath scrub 39°37'08" N, May 23 2004 = NMW.Z.2005.011.02518 12

in abandoned blueberry 74°53'43" W JON 12054 14

field

Ocean 37 Colliers Mill WMA: acid 40°05'35" N, May 22 2004. NMW.Z.2005.011.02508 ~150

bog with Ilex, 74°25'58" W JEN 12036 73

Chamaedaphne, Aronia

38 Stafford Forge WMA; moist —39°42'44" N. May 22 2004 = NMW.Z.2005.011.02514 122

Kalmia-Pinus forest 74°22'10" W JON 12045 54

39°53'34" N May 22 2004 = NMW.Z.2005.011.02511 3

74°19'58" W JEN 12039 15

North Carolina

Bladen 39 sas Mill Bay (Bladen eas N Jun 02 2003 | NMW.Z.2005.011.02204 ~100

uakes SF); pocosin with 31/33” W JEN10613 142

ae yparis

Brunswick 40 Green Swamp (TNC); 34°06'14" N. fun O01 2003) NMW.Z.2005.011.02193 50

Chamaecyparis-bay 78°18'35" W JON 10615 94

forest

41 Green Swamp (TNC); 34°05'42" N, Jun O01 2003) | NMW.Z.2005.011.02194 20

medium pocosin 78°17'48" W JON 10617 19

42 Prospect Ridge; mature 34°03'48" N, Jun OL 2003 NMW.Z.2005.011.02196 25

pine-bay forest 78°20'52" W JCN 10622 2

Carteret 43 Millis Road (Croatan NF): 34°46'16" N, Feb 24 2003) NMW.Z.2005.011.02128 82

wet pocosin with 76°58'39" W JCN 10624

leatherleaf

Craven 44. Sheep Ridge (Croatan NF); 34°56'07" N, Feb 24 2003) NMW.Z.2005.011.02132 ~600

medium pocosin 77°04' 14" W NMW.Z.2005.011.02130 30

JON 10693 48]

JON 10708 10

45 Catfish Lake South 34°55'39" N. Feb 24 2003. =NMW.Z.2005.011.02126 ~400

(Croatan NF); low, wet 77°05'05" W JON 10675 257

pocosin

46 Catfish Lake South 34°55'10" N, Feb 24 2003 JCN 10668 l

(Croatan NF): roadside 77°05'24" W

ditch in medium pocosin

47 Neusiok Trail North 34°54'03" N, Jun 01 2003) NMW.Z.2005.011.02190 20

(Croatan NF); wet-mesic 76°49'06" W JCN 10686

pine-bay forest

ones 48 Catfish Lake Wildemess 34°55'07" N, Feb 24 2003) NMW.Z.2005.011.02125 40

(Croatan NF); mature 77°10'43" W JCN 10713 64

bay-pine forest

Moore 49 Pinebluff; bay forest in 35°06'14" N, Jun 03 2003 JON 10746 is)

gulley along US 1 79°28'28" W

Pamlico 50 Goose Creek Game Land: 85°15'14" N, May 31 2003 | NMW.Z.2005.011.021S8 10

pine straw under scrub 76°35'52" W

Pender 51 Holly Shelter game land; 8157" N, Apr O1 2003) NMW.Z.2005.011.03830 Holotype

edge of mature mesic 76: 4’ 11" W NMW.Z.2005.011.02119 ~1700°

bay/pine forest NMW.Z.2005.011.02118 ~1700°

NMW.Z.2005.011.02120 ~1700°

CM73971 143°

UF348143 ~700°

52 Holly Shelter game land; 34°32'57" N, Apr O1 2003) | NMW.Z.2005.011.02428 ~600

dense pocosin scrub 77°46'54" W

53 Holly Shelter game land; 34°33'06" N, Apr O01 2003 NMW.Z.2005.011.02117 115

dense bay scrub 77°47'37" W

54 Lanier Quarry (TNC); 34°37'49” N, Jun OL 2003 NMW.Z.2005.011.02192 ~SO

Shrubs bordering 77°40'27" W JON 10753 64

pime-wiregrass Savanna

rrell 55 Pocosin Lakes NWR; low 35°42'30" N, Apr 03 2003) NMW.Z.2005.011.02122 ~900

pocosin 76/11" W NMW.Z.2005.01 1.02123 ~9O0

May 31 2003 JCN 10824 92

B. F. Coles and J. C. Nekola, 2007

Table 1. Continued

State/County # Site; Habitat!

Coordinates Date

Accession Number Specimens

56 Pocosin Lakes NWR:

maple-oak-pine woodland

35°40'19" N,

76°12'16" W

Apr 03 2003 NMW.Z.2005.011.02121 50

57. Frying Pan Landing 35°48'03" N, Apr 03 2003 | NMW.Z.2005.011.02174 |

(Pocosin Lake NWR); 76°06'00" Wo May 31 2003 JEN 10828 l

pine pocosin ;

South Carolina

Horry 58 Lewis Ocean Bay Preserve; 33° ee N, Jun 02 2003 NMW.Z.2005.011.02201 ~200

medium pocosin on 78°50'36" Wo JON 10955 300

roadside

59 — Lewis Ocean Bay Preserve; 33°47'16" N, Jun 02 2003 NMW.Z.2005.01 1.02197 ~250

mesic pine-bay forest 78°50'56" W JON 10960 123

60 Lewis Ocean Bay Preserve; 33°47'33" N, Jun 02 2003 NMW.Z.2005.011.02200 95

mesic longleaf pine forest — 78°51'02” Ww JCN 10964 26

' Abbreviations used are: NF National Forest, NWR National Wildlife Refuge, SF State Forest, TNC The Nature Conservancy,

WMA wildlife management area.

However, V. cristata has striate (not pustulose) sculpture

on the body whorl, has a weak crest (rather than a basal

inflation), lacks an infi raparietal lamellae, and has a nodu-

lar (not shelf-like) columellar lamella. These two species

were found co-occurring in several New England loca-

tions (Table 1, sites 21, 92, 23. and 29), where they could

readily be distinguished under low m: agnification.

Vertigo mulled also resembles Ve rtigo ventricosa

(Morse, 1865) and Vertigo perryi Sterki, 1905 with re-

spect to the ovoid shape, large aperture vs. shell height

ratio, reflected lip (Pilsbry, 1948: 958. figs. 1-3, 7), and

basal apertural inflation (Coles and Nekola mpnen

data); V. perryi also has a dark colored peristome (Pils-

bry, 1948: 968). However, these species cannot easily be

confused because V. ventricosa and V. perryi have glossy

shells with only weakly developed striae, lack an Ear

parietal lamella, and have a peg-like columellar lamella.

Although not previously reported in the genus Vertigo,

the pustulose sculpture ‘of the body whorl in Vertigo

malleata is not unique to this species; Vertigo iE ee

sis and Vertigo conecuhensis (Pilsbry, 1948: 949, fig. 510,

9, 12-14: 950, fig. 511) of Ree ee North America

also weakly exhibit this trait (Figures 18, 19). At low

magnification, the sculpture of V. malleata also some-

what resembles the pitted or granular surface of mem-

bers of the Nesopupinae. However, members of the

Nesopupinae commonly have an angular lamella (i.e.,

Nesopupa, Sterkia), while none are aown to have an

intraparietal lamella. Supertficially, V. malleata also ap-

pears similar to the neotropical genus Bothriopupa (Pils-

bry, 1948: 1011, fig. 539). However, with respect to

shape, color, nature of the surface sculpture and configu-

ration of the major apertural lamellae and folds, V. mal-

leata much more closely resembles other members of the

genus Vertigo (Figures 16, 17).

Geographic Distribution: Vertigo malleata occurs

from southern Maine to southeastern Georgia along the

Atlantic coastal plain to the west side of Mobile Bay

along the Gulf coastal plain, apparently excluding pen-

=acuiler Florida (Table 1, Figure 20), This distribution

includes a number of regions of particular ecological in-

terest and conservation concern, e.g. es Pine Barrens of

New Jersey (sites 30, 32-35, 37-38), the sandhills and

pocosins of the Nou Carolina ie South Carolina

coastal plain (sites 39-48, 51-60), the Okefenokee

Swamp of southeastern Georgia (sites 19, 20), and the

Appalachicola sand plain of western Florida ( (site s 8-17).

It seems likely that its distribution extends into the Gulf

coastal plain of Mississippi and eastern Louisiana. Al-

though it is not yet known whether the species range

extends beyond the eastern USA, given the known ranges

of Atlantic coastal plain plant species (Sorrie and Weak-

ley, 2001) the sand plains of southern Nova Scotia would

appear to be an appropriate location for future surveys.

Preferred Habitats: Vertigo malleata occurred in ap-

proximately two-thirds of all surveyed acid habitats. In

southern Maine and Massachusetts it was found in damp

and lightly compacted leaf litter on Sphagnum bogs un-

der a dense cover of ericaceous and other acidophile

shrubs (e.g., Gaylussacia, Vaccinium, Kalmia, and

Myrica). In this region it was also present in Atlantic

white cedar bogs, where it occurred in leaf litter accu-

mulations on mossy hummocks, In the New Jersey Pine

Barrens V. malleata was found in dense leaf litter under

tall heath (Vaccinium, ach pas ia, Kalmia), Myrica, and

Ilex scrub at the edges of bogs, Atlantic white cedai

swamp forest, and mesic microsites in upland pine-oak

forest. Populations in North and South Carolina were

primarily located under dense heath, bay, holly, and wax

myrtle scrub in pocosins, bay forest, wet-mesic pine

woodland, and pine-wiregrass savanna. At Pocosin Lakes

National Wildlife Re fuse, for example, V. malleata was

abundant in leaf litter on scrub se cee within

flooded pond pine woodland (Table 1, site 55), absent in

adjacent broadleaf woodland, and is sent only in rela-

tively low numbers at the transition zone (site 56). Popu

THE NAUTILUS, Vol. 121, No. 1

Figures 1-19. Vertigo malleata and related taxa. 1-4. Vertigo malleata. Holotype, NMW.Z.2005.011.03530, Holly Shelter Game

Lands, Pender County, North Carolina, 34°31'57" N, 77°44'41" W; 1. Apertural view. 2. Abapertural view. 3. View showing parietal

and upper palatal lamellae. 4. Sculpture on body whorl surface, width of detail is 0.25 mm. 5. Vertigo malleata, second specimen from

the type locality, NMW.Z.2005.011.03831, showing more conical shell shape. 6. Vertigo malleata, NMW.Z.2005.011.03832, Stafford

Forge WMA, Ocean County, New Jersey, 39°42'44" N., 74°22'10" W, showing small size and lack of infraparietal and subcolumellar-

basal lamellae. 7-8. Vertigo malleata, NMW.Z.2005.011.03533, Wells Heath, York County, Maine, 43°20'2"” N, 70°38'24" W. 7.

\pertural view. 8. View from apex showing apical whorls and the basal apertural dilation. 9. Vertigo malleata,

NMW.Z,2005.011.03834, Wilma Station, Liberty County, Florida, 30°9'34" N, 84°57'39" W, showing strong shell sculpture, a series

of subcolumellar-basal lamellae, and a weak infraparietal lamella. 10, 11. Vertigo malleata, NMW.Z.2005.011.03835, Pond Creek

seep, Covington County, Alabama, 31°6'12"” N, 86°32'3" W. 10. Apertural view showing subcolumellar and basal lamellae, an

indistinct nodule below the columellar lamella, and lack of an infraparietal lamella. 11. Abapertural view, 12. Vertigo malleata,

NMW.Z.2005.011.03536, Collins Bay, Ware County, Georgia, 31°05'12" N, $2°36'56" W, showing elongate shape, fused sub-

columellar and basal lamellae, and distinct sinulus. 13. Vertigo malleata, NMW.Z.2005,011.03837, Skunknett Audubon Preserve 2,

Barnstable County, Massachusetts, 41°38'59" N, 70°22'31" W, showing light shell color and basal lamella only. 14. Vertigo malleata,

NMW.Z.2005.011.03838, Peterson Swamp WMA, Plymouth County, Massachusetts, 42°00'37" N, 70°49'4"” W, showing small size,

vestigal infraparietal, and reduced basal lamellae. 15. Vertigo malleata, NMW.Z.2005.011.03839, Lewis Ocean Bay Preserve, Horry

County, South Carolina, 33°47'16" N, 75°50'56" W, showing bi-lobed basal lamella. 16. Bothriopupa tenuidens (C.B. Adams, 1845),

FMNE 106420, Louis Brand Collection, Columbia University. 17. Bothriopupa conoidea (Pfeiffer, 1853), FMNH 119055, Kyk-

Over-All Island, Kartabo, British Guiana. 18. Vertigo conecuhensis, JON 12364, Pond Creek seep, Covington County, Alabama,

31°6'12" N, 86°32'3" W. 19. Vertigo alabamensis, JCN 10781, Lanier Quarry, Pender County, North Carolina, 34°37'49" N, 77°40/27" W

lations in Georgia, Florida, and Alabama were found pri-

marily in bay scrub along swamp margs small water

courses, and seepage Zones within pinelands Populations

were also rarely encountered in mesic pine forest frag-

ments that had escaped frequent fire management (see

below): again, individuals were restricted to humid litter

tumulations. Throughout its range, Vertigo malleata

ppeared to avoid even moderately less acidic habitats

uch a sedge meadows (Maine, Massachusetts), cattail

swamps and marshes (Maine, Massachusetts, New Jer-

sey), and bottomland bald cypress/water tupelo/

sweetguin forests (North and South Carolina, Georgia,

Florida, and Alabama)

Associated Land Snails and Community Composi-

tion: Across all 49 analyzed sites (Table 2), Vertigo

malleata constituted 35% of total individuals. This frac-

tion appeared to be inversely correlated with latitude,

B. F. Coles and J. C. Nekola, 2007

g2°W ee 88°W

84°W

96°W g2°W 88°W

igure 2

Figure 20.

80°W 76°W

84°W

Distribution of Vertigo malleata in eastern North America. Black circles represent sites supporting populations and

72°W 68°W

42°N

38°N

34°N

26°N

80°W 76°W 72°W

open circles represent inventoried sites that do not harbor this species.

ranging from 17% in the New Jersey Pine Barrens to

32% in New England, 35% in the Carolina coastal plains,

and 76% in the Gulf coastal plain. Population de ioe s of

Vv _malle vata were frequently observed to exceed 500 per

m*, with an estimated de ue of the order of 2000 per m-

at the type locality (Table 1, site 51). These densities

range among the highest re ets ed for any land snail spe-

cies (Frest and Johanne s 1995, Cameron 2003).

A total of 34 terrestrial mollusk taxa and 5886 indi-

viduals were observed from these sites (Table 2). The

average number of co-occurring taxa was 3.69 + 0.37,

and ranged from 0-9. Throughout its range, the ten most

frequently co-occurring taxa were: Striatura milium

17% of all other individuals), Strobilops texasiana

(15%), Vertigo milium (13%), Punctum minutissimnaun

(11%), Vertigo alabamensis (9%), Striatura meridionalis

(8%), Gastrocopta pentodon (8%), Euconulus trochulus

(4%), Euconulus chersinus (2%), and Gastrocopta con-

tracta (2%). The most frequent co-occurring taxa varied

by region: Striatura milium and Punctum minutissiu-

mum in New England: Striatura meridionalis, Striatura

milium, Punctum minutissimum, and Gastrocopta pent-

odon in the New Jersey Pine Barrens; Strobilops texasi-

ana, Vertigo milium, and Vertigo alabamensis along the

Carolina coastal plain; and Vertigo alabamensis, Stria-

tura meridionalis, and Gastrocopta pentodon along the

Gulf coastal plain. Co-occurring Vertigo taxa also varied

by region: Vertigo cristata, Vertigo perryi, and Vertigo

Page 26

THE NAUTILUS, Vol. 121, No. 1

Table 2. Frequency of co-occurring species across the range of Vertigo malle ata!

{

Number of co-occurring individuals (%)

New Jersey

Carolina coastal plain Gulf coastal plain Total

Taxon New England

Vertigo malleata sp. nov. 308

Striatura milium (Morse, 1859) 480

Strobilops texasiana Pilsbry & Ferris, 1906

Vertigo milium (Gould, 1840)

Punctum minutissumum (1. Lea, 1541) 113

Vertigo alabamensis Clapp, 1915

Striatura meridionalis (Pilsbry & Ferris, 1906)

Gastrocopta pentodon (Say, 1821) 6

Euconulus trochulus (Reinhardt, 1883)

Euconulus chersinus (Say, 1821)

Gastrocopta contracta (Say, 1822)

Glyphyalinia solida (H. B. Baker, 1930)

Glyphyalinia sp.

Vertigo oralis Sterki, 1898

Hawaiia miniscula (A. Binney, 1840)

Glyphyalinia luticola Hubricht, 1966

Vertigo conecuhensis Clapp, 1915

Gastrocopta tappaniana (C. B. Adams, 1842) 4

Zonitoides arboreus (Say, 1816) 12

Euconulus fulvus (Miiller, 1774) 20

Vertigo ovata Say, 1822

Neohelix solemi Emberton, 1988 ]

Ventridens cerinoideus (Anthony, 1865)

Vertigo cristata (Sterki, 1919) 13

Nesovitrea electrina (Gould, 1541) 6

Vertigo perryi Sterki, 1905 5

Gastrocopta riparia Hubricht, 1978

Helicodiscus parallelus (Say, 1817)

Troidopsis soelneri (J. B. Henderson, 1907)

Deroceras sp.

Glyphyalinia indentata (Say, 182° |

Vertigo rugosula Sterki, 1890

Vertigo ventricosa (Morse, 1865) 2

Striatura ferrea Morse, 1864 1

Triodopsis hopetonensis (Shuttleworth, 1852)

Total co-occurring individuals 664

Co-occurring species richness 13

340, 1S07 779 3234

407 119 1006

61 $35 ll 907

765 768

398 164 675

494 64 558

413 25 46 484

258 156 45 465

216 216

120 3 123

109 109

15 64 19 98

28 63 91

61 6 67

53 53

28 12 3 43

39 39

28 32

5 14 31

20) 20

1S 19

19 19

3 3

2 2

| 2

2 2

| 1

1613 3359 250, 5886

9 26 LO 34

' Data taken from 49 discrete sites of the junior author collection (lots prefixed by JCN in Table 1),

* Juveniles and young adults of unclear identity.

ventricosa were sympatric in New England, while Ver-

tigo milium, Vertigo alabamensis, Ve rtigo oralis, Vertigo

conecuhensis, and Vertigo rugosula were sympatric in the

Carolina and Gulf coastal plains.

CONSERVATION IMPLICATIONS

The data presented here show that Vertigo malleata is a

characteristic component of the base-poor biota of the

Atlantic and Gulf coastal plains, having been found in

66% of surveyed acid sites, and accounting for up to 75%

of all mollusks in these sites. The sundance ancl wide-

spread occurrence of V. malleata would seemingly sug-

gest that it is not of immediate conservation concern.

However, it appears that the species is in fact under

threat because of the widespread use of fire as a man-

agement tool. Many coastal plain habitats, including

those of V. malleata (i.e., pine woods, pine-wiregrass sa-

vanna, and pine barrens), have come to be wiewed by

many plant ecologists as pyrogenic (Myers, 1985; Chris-

tensen, L98S) \ and are being typically managed by anthro-

pogenic fire return inte eal of <5 years, with mz uny areas

being burned annually. However, such high- freque ney

fire man: wement policies have been tas to exact a

pan negative impact on total biodiversity, including

Le pidopter: i, Homoptera, Hymenoptera, Araneae, ( ‘ol-

lembola (Swengel, 1996, 1998: Harper et al., 2000), and

terrestrial Mollusca (Ne kola, 2002b).

The impact of fire on Vertigo malleata is illustrated by

its distribution in the Appi lachicola uplands of F lorida.

B. F. Coles and J. C. Nekola, 2007

Page 27

We were unable to find V. malleata in forest that had

been burned within three years, however, the presence

of substantial populations in a tiny unburned inholding of

mesic pine forest (Table 1 site 11), unburned mesic pine-

red maple-Atlantic white cedar forest (Table 1 site a

and unburned mesic margins of wetlands (Sites 8,

12-13, 15-17) suggest that while it is not ay a

restricted to wetlands, it has become lar gely limited to

these sites simply because they remain unburned. While

these observations require further inv estigation, we esti-

mate that at least 95% of the \

the Apalachicola National Forest has been eliminated by

management practices. Conversely, the presence of 14

malleata in mesic bay-pine forest that had regenerated

after burn (Table 1 site 10) shows that, apart from its

intrinsic interest as an extreme calcifuge, this snail can

potentially be used to monitor recovery from over-

burning.

ACKNOWLEDGMENTS

We thank Beata Pokrysko (Museum of Natural History,

Wroclaw University, Wroclaw, Poland) for comments

concerning the identity of Vertigo malleata as an unde-

scribed species, and to Jochen Gerber for loans of Both-

riopupa material from the FMNH collection. José H.

Leal, Barry Roth, and John Slapcinsky all provided useful

comments on earlier drafts. The following U. S. Federal

Government agencies, U.S. State Gov ernment agencies,

private organizations and individuals have helped during

this study by providing access to lands and information

on sites: Alabama Department of Conservation and

Natural Resources, Forever Wild Program (Eric Soe-

hren); Maine Department of Inland Fisheries and Wild-

life. Wildlife Resource Assessment Section (Beth

Swartz): Massachusetts Natural Heritage and Endan-

gered Species Program (Tom French and Jennifer

Loose); New Jersey Department of Environmental Pro-

tection, Division of Fish and Wildlife, Endangered Spe-

cies Program (Dave Golden); New Jersey Department of

Environmental Protection, Division of Parks and Forests

(Thomas Keck, and the Superintendents of Belleplain

State Forest-W. Scott Mauger, Brendan T. Byrne State

Forest-Christian Bethmann, and Wharton State Forest—

Lynn Fleming); North Carolina Division of Forestry Re-

sources, Bladen Lakes State Forest (Michael Chesnutt);

North Carolina Wildlife Resources Commission

(Stephen Hall and Randall Wilson); South Carolina De-

partment of Natural Resources, (Lewis Ocean Bay Pre-

serve Manager—Jamie Dozier); The Nature Conservancy,

North Carolina Chapter (Dan Bell and Bruce Sorrie):

The Nature Conservancy, Southern Maine Field Office

(Parker Schuerman and Keith Fletcher); U. S. Forest

Service. Apalachicola National Forest, Wakulla District

(Marcus Beard): U.S. Fish and Wildlife Service, Pocosin

Lakes National Wildlife Refuge (Wendy Stanton).

V. malleata population of

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Boycott, A. E. 1934. The habitats of land mollusea in Britain.

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Cameron, R. A. D. 2003. Life-cycles, molluscan and botanical

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tropoda, Pulmonata: Vertiginidae). Heldia. 5: 95-110.

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species richness and composition of land mollusc commu-

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Emberton, k. C., T. A. Pearce, and R. Randalana. 1996. Quan-

titatively sampling land-snail species richness in Mad: agas-

can rainforests. Malacologia 38: 203-212.

Frest, T. J. and E, J. Johannes. 1995. Interior Columbia Basin

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Hubricht, L. 1985. The distributions of the native land molluscs

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Kerney, M. P. and R. A. D. Cameron. 1979. Field guide to the

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Collins Press, London, 288 pp.

Myers, R. L. 1985, Fire and the dynamic relationship between

Florida sandhill and sand pine scrub vegetation. Bulletin

of the Torrey Botanical Club 112: 241-252.

Nekola, J. C 2001. Distribution and ecology of Vertigo cristata

(Sterki, 1919) in the western Great Lakes region. Ameri-

can Malacological Bulletin 16: 47-52.

Nekola, J. C. 2002a. Distribution and ecology of terrestrial gas-

tropods in northwestern Minnesota. Final Report, Minne-

sota Department of Natural Resources, St. Paul, 200 pp.

Nekola, J. C. 2002b. Effects of fire management on the rich-

ness and abundance of central North American grassland

land snail faunas. Animal Biodiversity and Conservation

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Nekola, J. C. 2004. Terrestrial gastropod fauna of northeastern

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American Malacological Bulletin 1S: 21-44.

Nekola, J. C. and B. F. Coles. 2004. Eastern Massachusetts

Vertigo perryi survey. Final Report, Massachusetts Divi-

sion of Fisheries and Wildlife, Westborough, 20 pp

Oggier, P., S. Zschokke, and B. Baur. 1998. A comparison of

three methods for assessing the gastropod community in

dry grasslands. Pedobiologia 42: 348-357.

Pilsbry, H. A. 1920. Manual of Conchology, structural and sys-

tematic. Pupillidae. Vols. 24-25. Ac ade smy of Natural Sci-

ences, Philadelphia.

Pilsbry, H. A. 1948. Land Mollusca of North America (North of

Mexico). Volume 2, part 2. Academy of Natural Sciences,

Philadelphia, pp. 521-1113 + xlvii.

? ere

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Pokryszko, B. M. and R.A.D. Cameron. 2005. Geographical

variation in the composition and richness of forest snail

faunas in northern Europe. Records of the Western Aus-

tralian Museum, Supplement 68: 115-132

Schilthuizen, M., H. N. Chai, T. E. Kimsin, and J. J. Ver-

meulen. 2003. Abundance and diversity of land snails

(Mollusca: Gastropoda) on limestone hills in Borneo.

Raffles Bulletin of Zoology 51: 35-42.

Sorrie, B. A. and A. S. Weakley. 2001. Coastal plain vascular

plant endemics and phytogeographic patterns. Castanea

66: 50-82.

Swengel, A. B. 1996. Effects of fire and hay management on

the abundance of prairie butterflies. Biological Conserva-

tion 76; 73-85.

Swengel, A. B. 1998. Comparisons of butterfly richness and

abundance measures in prairie and barrens. Biodiv ersity

and Conservation 7: 639-659. °

Turgeon, D. D., J. F. Quinn Jr., 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. Scheltema, F. G.

Thompson, M. Vecchione, and J. D. Williams. 1998. Com-

mon and Scientific Names of Aquatic Invertebrates from

the United States and Canada, Mollusks, 2nd edition.

American Fisheries Society, Special Publication 26.

American Fisheries Society, Bethesda, ix + 526 pp:

THE NAUTILUS 121(1):29-36, 2007

>a0e@ 29

Page 29

Population dynamics of the fingernail clam Sphaerium

occidentale (Lewis, 1856) Genes Sphaeriidae) in an

intermittent pond

Meghna Roy

D. Dudley Williams

Department of Life Sciences

University of Toronto at Scarborough

1265 Milit: uv Trail, Scarborough

Ontario MIC 1A4, CANADA

ABSTRACT

A population of Sphaerium occidentale (Lewis, 1856) was stud-

ied over a two-year period in an intermittent freshwater pond

in southern Ontario, Canada. Sub- populations in control areas

of the pond showed marked differences between the two years,

which appeared to be related to different hydroperiods (34 days

in 2001 vs. 94 in 2002), water temperature, and density of

suspended bacteria. In the first year, there was recruitment of

young into the population in April/May, but there appeared to

be very little or no Reproduction in the second year. Resource

(decaying riparian leaves) addition and remoy al experiments

performed in the second year had an impact on some but not

all of the physico-chemical parameters measured in the pond,

and also affected bacterial densities. However, there were no

significant overall effects of these manipulations on sub-

populations of S. occidentale, although there was a trend to-

ward greater density and biomass in the resource removals

during the first half of the hydroperiod. Sphaerium occidentale

appears to be well adapted for survival in such harsh environ-

ments via a specialized physiology ry and, reproductively, employ-

ing a bet-hedging strategy However, populations may occa-

sionally be reduced to le vale that may result in local extinction.

To counter the latter, sphaeriids have dispersal mechanisms

that allow recolonization from metapopulations.

INTRODUCTION

Sphaeriid clams are widely distributed and many specie S

inhabit temporary fres shwater habitats. They are impor-

tant components of the benthic communities of such

habitats where they act as water clarifiers and organic

nutrient sinks (Thorp and Covich, 2001). Sphaeriids are

known to be simultaneous hermaphrodites, with the abil-

ity to self-fertilize. a trait well suited to founding and

maintaining populations in temporary waters (W sThanae:

2006). Sphaerium occidentale (Lewis, 1856), commonly

Corresponding author: Meghna Roy

email: [email protected]

known as Herrington’s fingernail clam, is unique among

the Sphae sriidae in that it is exclusive to temporary waters

(McKee and Mackie, 1951). Along with all other sphaeri-

ids, individuals of this species brood its direct-developing

young within special sacs in the immer demibranchs of the

gills, from where they are eventually released as benthic

juveniles (Mackie et al., 1974). Although many sphaeriids

are synchronous brooders, species of ‘Sphae rium are se-

quential brooders—that is, several sets of embryos, each

ina separate stage of ontogeny, are present within the

brood sacs, together, each the product of a separate

spawning (Mackie, 1978). Presumably, this trait is a bet-

hedging strategy conducive to releasing a subset of viable

young rapidly at the beginning of a hy droperiod of un-

certain length, while maintaining a reserve should the

first subset ie lost (Stearns 1992). Although the popula-

tion dynamics of Sphaerium occidentale are not well

known, McKee pe Mackie (1981) observed that the

species completed its life cycle in 24 weeks when main-

tained under a stable hydroperiod in the laboratory, as

opposed to three years in an intermittent pond. This

finding supports the hypothesis of Thorp and Covich

(2001) that seasonal variation in water level drives bivalve

life history traits. The purpose of the present study was to

determine any relationship between hy droperiod length

and the popul ition dynamics of S. occidentale in an in-

termittent pond, and to explore other possible influences

through field manipulation experiments.

MATERIALS AND METHODS

The study population lives in an intermittent pond in

Vandorf, southern Ontario, Canada. The pond is devoid

of fish and has an area of approximately 1000 mn at the

beginning of hydroperiod. It is surrounded by a hay field

ane wieed deciduous woodland and supports emergent

vegetation comprising mainly Phalaris grass. The pond

substrate is primarily muddy and homoge neous through-

out. The population was studied over a two year pe riod

Page 30

THE NAUTILUS, Vol. 121, No. 1

in 2001 the pond held water for 34 days and had a maxi-

mum depth of 66 cm; in 2002 it held water for 94 days

and had a maximum depth of 48 cm.

The populations were studied in six randomly chosen

areas of the pond that were enclosed with watertight,

circular galvanized sheet-metal walls installed prior to

snowmelt in 2001. The walls were embedded 10 cm into

the substratum and rose well above the water surface,

forming corralled homogeneous sections of the pond

each 2.4 m in diameter. Two of the enclosures were

chosen at random to serve as controls. In the autumn of

2001, the locations of the control enclosures within the

pond were changed in order to avoid carry-over effects,

and the other four enclosures were manipulated to either

receive additional riparian leaf litter (two each received a

mixture of dry maple and beech leaves |~3.0 kg] that had

been dried for 24 hours at 100°C), or have litter removed

(~1.540.04 kg of leaves and other vegetation from each

of the remaining two). It might be argued that the use of

imperforate galvanized metal enclosures may have ren-

dered the treatments uninhabitable (i.e., produced a

container effect) for S. occidentale, but that is highly

unlikely as clams are virtually immobile and non-

selective filter feeders. Further, evidence from other taxa

(e.g., ciliates) in this pond showed that the enclosures

resulted in higher species richness and abundance (An-

drushchyshyn et al., 2006).

The environments within the enclosures were sampled

on a weekly basis beginning as soon as the pond basin

filled in 2001 and again in 2002. The following param-

eters were measured: water depth, with a one-meter

stick, and dissolved oxygen, pH, temperature, and con-

ductivity with a portable Hach Kit spectrophotometer

(DR2000: Hach Company, Loveland, Colorado, USA).

Turbidity, ammonia, nitrate, and phosphorus were mea-

sured using a Hydrolab multiprobe (Hydrolab Corpora-

tion, Austin Texas, USA), and chlorophyll ad was mea-

sured using the acetone extraction method (American

Public Health Association, 1995). To assess bacterial

density, a column of water extending from the bed to the

surface was collected using a plastic tube; this was done

at two locations within each enclosure and then com-

bined in a bucket from which a subsample of 30 cm Was

removed and stained with acridine orange. These stained

samples were then filtered through a 0.2 wm Sartorius

filter (Cat. No. 13007) and the bacteria counted (on ~20

fields with a concentrations of ~200 bacteria per field of

view) under an epifluorescence microscope, following

the method of Sorokin (1999).

( Q uns Were Se imple d using a calvanized-steel box sam-

pler (area 0.1 m~) which was inserted 2 cm into the pond

Bed. Two such bee samples were taken at regular

intervals from random locations in the control enclosures

in 2001 and from the control and treatment enclosures in

2002. All samples were put in large Ziploc bags, labeled,

and preserved in 4% formalin in the field. In the labo-

ratory, clams were removed by sieving and handpicking

inder a magnifying lens from samples collected on 7

April, 3 May, 10 July, and 21 August in 2001, and 7 April,

6 May, 3 June, 11 July, and 6 September in 2002. All

specimens were counted, blotted on filter paper to re-

move excess water, and weighed to the nearest 0.0001 ¢

using a digital balance. All specimens proved to be

Sphaerium occidentale, and identification was confirmed

by Dr. Gerald Mackie (University of Guelph, Canada).

Clam le ngths (anterior to posterior) were measured to

the nearest 0.1 mm using a dissecting microscope fitted

with an ocular micrometer, and specimens were assigned

to the following four length classes: <2.5, 2.5-4.0, 4.1—

5.5, and > 5.5 mm.

RESULTS

CONTROL POPULATIONS:

In 2001, the hy ‘drope riod of the Vandorf pond was short

(34 days, from 7 April to 17 May) and likely related to a

rapid 10° - warming of the pond i in the first week of May

(Figure la, b) and low rainfall. Associated with this

warming was a significant increase in the density of sus-

pended Thactenia [Figure le; F=7.14, p=0.02 . from a re-

peated-measures ANOVA], perhaps partly a concentra-

tion effect, and a marked increase in the growth of indi-

vidual clams (Figure 2a). From May to July, 2001, mean

biomass increased from 0.33+0.02 to 2.1+0.22 ¢ 0.1 m 2

of pond bed area (ANOVAR F=5.95, p=0.03). At the

times of sampling the dry pond bed in July and Septem-

ber, 2001, no additional growth was noted. In 2002, the

hydroperiod was considerably longer (94 days) and

started almost three weeks earlier (18 March to 19 June;

Figure 1b), While water temperature attained the same

maximum value as in 2001, its increase was more gradual

(Figure la). Bacterial densities throughout 2002 were

more stable, except for a minor increase in early June

(Figure le). Clam growth during 2002 was very different

from 2001 and was largely limited to shifts from size class

| to 2 in May-June (Figure 2b; ANOVAR F=12.88,

p=9.005). Biomass increase -d from 0.24+0.14 ¢ 0.1m in

May to 0.88+0.39 ¢ 0.1 m~ in July (ANOV AR F=15.13,

p=0.0001). a

Between-year comparisons suggest that while there

was recruitment of young into the population in April/

May of 2001 (Figure 2a), the similarity in population

structure between Se ptember, 2001 and April/May, 2002

and concomitant decline in density of juveniles (Figure

2b) indicate that re production did not occur im 2002.

MANIPULATED POPULATIONS:

The resource addition and removal experiments per-

formed in 2002 had an impact on some but not all of the

physico-chemical parameters measured in the Vandorf

pond. Largely unaffected, compared with the control en-

closures, were water temperature (although there was an

increase in the resource addition enclosures in May),

water depth, pH phosphorus, and chlo-

rophylla. Affected were dissolved oxygen levels (typically

(Figure 3 a,b,c)

M. Roy and D. D. Williams, 2007

Page 3]

oO —@— 2001

oo 15.4 ——m— 2002

Sos],

0) T 1 1 T [—— r T 1

March April May June

Month

Depth (cm)

March April May June

Month

Cc)

s 30000 -

w 25000 -

xe}

E 20000 -

5 10000 -

S 4

S 5000 2

(0) TT oer ee a oe

Mar-18 Apr-1st Apr-3rd May-1st May-2nd Jun-03 Jun-20

week week week week

Date

Figure 1. Seasonal variation in water temperature, depth, and bacterial density in the control enclosures in 2001 and 2002. Mean

values are shown for the control enclosures. Error bars for bacteria indicate +1SE (n = 2).

Page 32

THE NAUTILUS, Vol. 121, No. 1

Control - 2001

we sample

taken

(7) (40) (448) (363)

|r| ol}

Control - 2002

ive

(69) (39) (83) (87) (96)

Size class

rej

asx

—

(79) (54) (99) (167) (123)

Resource removal - 2002

tet dt

(128) (114) (188) (145)

ori May janet (aii guy ay) September ( (dry)

s/n) ol)

Figure 2. Seasonal length frequency a aneee of

Sphaerium occidentale in 2001 and 2002. Size classes 1, 2, 3,

and 4 represent clams < 2.5, 4, 5.5, and >

in length. Numbers in parentheses are sample Sizes.

5.5 mm, respe ative ly,

highest in the resource removal and lowest in the re-

source addition treatments) and nitrate (highest in the

resource additions early on in the hydrope wig)

4 ab). Bacterial densitie Ss were highest in the resource

Seen treatments, lowest in the resource removals, and

intermediate in the controls during the first half of the

hydroperiod, but more similar thereafter (Figure 4c)

Despite differences in the above parameters, there

Figure S

were no significant overall treatment effects on the den-

sities or biomass of S. occidentale, although there was a

trend towards greater density and biomass in the re-

source removals during the first half of the hydroperiod

(Figure 5a,b). Neither did there appear to be any notable

differences in the population structures in the addition

and removal enclosures compared with the 2002 controls

(Figure 2b,c,d).

DISCUSSION

a sharp rise in water temperature and rapid decline in

le pt th of the Vandorf pond in early May of 2001, accom-

panie od by low rainfall, likely promote 1d the sion hydro-

pe riod in that year. Bivi alve feeding filtration rates are

known to be proportional to the ambient temperature

| horp and ¢ Lame 8 2001)

ing of the pond fortuitously allowed

Thus the sudden 10°C warm-

occidentale to

benefit maximally from the high density of suspended

bacteria available at that time, and to grow rapidly. How-

er, and in spite of the longer (~3 times) hydroperiod in

2002, during this second year S. occidentale individuals

grew much more slowly—although a growth spurt did

occur during the higher temperatures "reached prior to

pond dry-up in June. Based on this two-year comparison,

it would appear that individual clam growth does not

necessarily benefit from a longer hy rope »riod—indeed a

shorter one may be better provided that food supply and

water temperature are favorable. Thomas (1963) noted,

however, that S. partumeium (Say, 1822) grew more un-

der a longer hydroperiod although its life span is 12-13

months compared with up to 3 years in S. occidentale.

The former species may well be predisposed to contin-

ued growth in that its semelp: wrous strategy commits it to

development and reproduction under sometimes less

than optimal pond conditions. S. occidentale appears to

demonstrate more plasticity in its growth rate (McKee

and Mackie, 1981), and its iteroparous strategy may allow

it, on occasion, to forego a less than favourable repro-

ductive opportunity,

Only adult (our size class 4) S. occidentale release off-

spring (McKee and Mackie, 1951). Some of the sub-

adults present in April 2001 grew rapidly, attained size

class 4, and produced large numbers of size class 1 and 2

young, as evident from the very high densities found in

July 2001. As S. occidentale is, as previously noted, a

sequential brooder it is conceivable that the high July

densities may have resulted from multiple release events

of young from ¢ April until the end of the hydroperiod.

The same size-class proportions present in the July and

September samples indicate that no further growth took

place during the period when the pond bed was dry,

which agrees with the findings of Mckee and Mackie

(1983) for this species. Further, the presence of the same

distribution of size classes in April (2002) shows that

there was no winter growth or container effect from

the use of imperforate metal enclosures. The substantial

drop in clam density between September 2001 and

April 2002 likely indicates winter mort: lity. Throughout

2002 no significant increase occurred in the control

population and this, together with modest shifts towards

the large size classes by September, suggests that the

population largely | failed to breed in this second year.

Unlike S. partumeium, which is more commonly found

in permanent waters (Clarke, 1973), and has a semelpa-

rous, univoltine life cycle where only juveniles aestivate

and the resulting adults breed and die the next spring

(Way et al., 1980), S. occidentale can aestivate in all size

classes and has the potential to breed several times and

over more than one hydroperiod as pond conditions al-

low.

Interestingly, the initial samples collected in a

2001 produced very few clams, and no live adults.

possible explanation for this is that when sphaeriids en

their terminal size, they do not survive a subsequent

dry phase (Way et al. 1980). It is like ly, then that the

majority of the previous year’s population had been

adults at the end of the 2000 hydroperiod, The popula-

tion was thus at a very vulnerable stage in early 2001, but

M. Roy and D. D. Williams, 2007 Page 33

—@— Resource addition

O ——l—— Resource removal

sill Control

-E

—

Mar- Mar- Mar- Apr- Apr- Apr- Apr- May- May- May- May- Jun- Jun- Jun-

18 25 31 7 16 22 29 6 13.20 27 03 10 17

Date

Depth (cm)

Mar- Mar-Mar- Apr- Apr- Apr- Apr- May- May- May- May- Jun- Jun- Jun- Jun-

18 25 31 7 16 22 29 6 13 20 27 £403 10 17, = 20

Date

= 2)

Mar- Mar- Mar- Apr- Apr- Apr- Apr- May- May- May- May- Jun- Jun- Jun-

18 25 31 7 16 22 29 6 13 20 27 03 10 17

Date

Figure 3. Seasonal variation in water temperature, depth, and pH in the treatment enclosures in 2002. Mean values are shown for

the control. resource addition, and resource removal enclosures.

Page 34 THE NAUTILUS, Vol. 121, No. 1

Dissolved 0 (mg/L)

Mar- Mar- Mar- Apr- Apr- Apr- Apr- May- May- May- May- Jun- Jun- Jun-

18 25 31 ig 16 22 29 6 13 20 27 #203 #10 = «17

Date

Nitrate (mg/L)

Mar- Mar- Mar- Apr- Apr- Apr- Apr- May- May- May- May- Jun- Jun- Jun-

18 25 31 7 16 22 29 6 13 20 27 #03 10 17

Date

a) —@— Resource addition

= ——™—— _ Resource removal

é Control

19)

fan)

Mar-18 Apr-7 Apr-22 May-06 May-20 Jun-03 Jun-17

Date

Figure 4. Seasonal variation in oxygen, nitrate, and bacterial densities in the treatment enclosures in 2002. Mean values are shown

for the control, resource addition, and resource removal enclosures. Error bars for bacteria indicate +1SE (n = 2).

was quite quickly restored by July, only to suffer sub- ated with an increase in dissolved oxygen and a lower

stantial winter mortality in all size classes later that same concentration of suspended solids, although improved

yeal water clarity did not affect chlorophyll @ production. Re-

Removing resources from the enclosures was associ- source addition reduced oxygen levels and increased tur-

M. Roy and D. D. Williams, 2007

x Control

= Resource addition

— Resource removal

—

7p)

ase,

April May

jo)

™~

ie)

jaa)

April May

June

July (dry) September (dry)

Month

July (dry)

September (dry)

Month

Figure 5. Mean values of Sphaerium occidentale densities in numbers 0.1 m~ and biomass in g 0.1 m~ in the treatment enclosures

in 2002. Mean values are shown for the control, resource addition, and resource removal enclosures. Density and biomass bars

indicate +1SE (n=2) and +1SD (n=2), respectively.

bidity, likely as a result of suspension of more litter

breakdown products. While it might be expected that an

increase in suspended food particles (including bacteria

in the first half of the hydroperiod) would have benefit-

ted the S. occidentale population (as shown for other

detritivores, by Richardson, 1991), it is known that too

high a concentration of suspended materials can inter-

fere with the filtering and respiratory mechanisms of

freshwater bivalves—although sphaeriids are known to

be very tolerant of hypoxia (Thorp and Covich, 2001).

Further, sphaeriids can also feed on pond bed deposits

and thus a direct response to food particle manipulation

may not be detectable. Despite the observed differences

in the physico-chemical environments and food levels

between the treatments and the controls, there were no

apparent differences in the structure (size and growth) of

the sub-populations. Lack of a differential response may

be a reflection of the fact that intermittent ponds are

normally subject to large within-vear and between-year

fluctuations in environmental variables (due to dilution,

. and thus their inhabitants may not

show population response patterns typical of those

evaporation, etc

known from permanent ponds (Brénmark and Hansson,

1998S).

The responses of the natural and manipulated sub-

populations of S. occidentale in the Vandorf pond suggest

that length of the hydroperiod was the major driving

force on population dynamics. Further, the species ex-

hibits a number of important adaptations that enable

population survival despite large inter-year variations

in its habitat, although populations may occasionally

be reduced to dangerously low numbers that may result

in local extinction. Sphaeriid clams are known, however,

to be able to colonize/recolonize these small ponds

via transportation on the bodies of more-mobile pond

inhabitants (e.g., Fryer 1974). It is likely that such re-

colonization events from a regional metapopulation are

common.

ACKNOWLEDGMENTS

We thank the Natural Sciences and Engineering Re-

search Council of Canada for funding, and Katarina

Magnusson for collecting the samples

Page 36

THE NAUTILUS, Vol. 121, No. 1

LITERATURE CITED

American Public Health Association. 1995. Chlorophyll. In:

Franson, M. A. H. (ed.) Standard Methods for the Exami-

nation of Water and Wastewater. American Public Health

Association, American Water Works Association and Wa-

ter Environment Federation, Washington, pp. 10.17—

10,24.

Andrushchyshyn, O. P., A. kK. Magnusson, and D. D. Williams.

2006. Responses of intermittent pond ciliate populations

and communities to in situ bottom-up and top-down ma-

nipulations. Aquatic Microbial Ecology 42: 293-310.

Brénmark, C. and L. Hansson. 1998. The ‘biology of ihe and

ponds. Oxford University Press, New York, 216 pp.

Clarke, A. H. 1973. The freshwater molluses of the Canadian

Interior Basin. Malacologia 13: 1-509.

Fryer, G. 1974. Attachment of bivalve molluscs to corixid bugs.

Naturalist 28: 18.

Mackie, G. L. 1978. Growth dynamics in natural populations of

Sphaeriidae clams (Sphaerium, Musculium, Pisidiwm). Ca-

nadian Journal of Zoology 57: 441-456.

Mackie, G. L., S. U. Qadri, and A.H. Clarke. 1974. Develop-

ment of brood sacs in Musculium securis Bivalvia: Sphaeri-

idae. The Nautilus 88: 109-111.

Mckee P. M. and G. L. Mackie. 1981. Life history adaptations

of the fingernail clams Sphaerium occidentale anil Muscu-

lium securis to ephemeral habitats. Canadian Journal of

Zoology 59: 2219-2299,

McKee P. BE and G. L. Mackie. 1983. Respiratory adaptations

of the fingernail clams Sphaerium occidentale and Muscu-

lium securis to ephemeral habitats. Canadian Journal of

Fisheries and Aquatic Sciences 40: 783-791.

Richardson J. S. 1991. Seasonal food limitation of detrivores in

a montane stream: an experimental test. Ecology 72: 873—

SS7.

Sorokin, Y. I. 1999. Aquatic microbial ecology—a textbook for

students in environmental science. 99. Backhuys Publish-

ers, Leiden, 248 pp.

Stearns, S.C. 1992. The Evolution of Life Histories. Oxford

University Press, Oxford, 249 pp-

Thomas, G. J. 1963. Study of a population of sphaeriid clams in

a temporary pond. The Nautilus 77: 37-43.

Thorp J. H. and A. P. Covich. 2001. Ecology and Classification

of North American Freshwater Invertebrates. 2°¢ Edition.

gry Press, New York, 1056 pp.

Way, C. M., D. J. Hormbach, and A. J. Burky, 1980. Com-

pe a life history tactics of the sphé veriid clam Muscu-

lium partumeium (Say), from a permanent pond and a

temporary pond. American Midland Naturalist 104; 319—

397

Williams, D. D. 2006. The Biology of Temporary Waters. Ox-

ford University Press, Oxford, 337 pp.

a SSS SSS SSS SSS SSS SSS

THE NAUTILUS 121(1):37-39, 2007

Page 37

Research Note

Paleoenvironmental significance of the eastern mud snail,

Ilyanassa obsoleta (Say, 1822

), from a microtidal coastal

sequence of southern New Eneland

Ilya V. Buynevich

Woods Hole Oceanographic Institution

Geology and Geophysics Department,

MS#22

Woods Hole, MA 02543 USA

[email protected]

Reconstruction of former sea-level positions is one of the

key issues in Quaternary paleoenvironmental research.

Along with high-marsh peat that has been widely used

for regional sea-level reconstructions due to its narrow

and robust elevation range, intertidal mollusks inhabiting

protected coastal embayments must also be eonsideied

as compleme ntary reference points for the water levels.

This paper presents evidence for potential use of the

eastern (Atlantic) mudsnail Iyanassa obsoleta (Say,

1822), as a paleoenvironmental indicator in middle Ho-

locene deposits of southern New England.

The eastem mudsnail occupies tidal flats along the

Atlantic seaboard, often occurring in dense populations

on the sediment surface of intertidal mudflats or mixed

flats (Brenchley, 1980; Whitlatch, 1952; Culbert and Ra-

leigh, 2001). Their feeding habits include scavenging,

predation, algal foraging, por deposit-feeding (Weiss,

1995: Kelaher et al., 2003), with feeding svthans shown

to be correlated with tidal cycles ( Goberson: 1979). Na-

tive along the Atlantic seaboard, Ilyanassa obsoleta has

been recently under stress of competitive exclusion and

egg predé ition by invasive gastropods, such as Littorina

littorea (Brenchley, 1982: Carlton, 1992) and has itself

become an invasive species in some parts of the West

Coast (Race, 1982). Although these snails are known to

burrow into the substrate or migrate into shallow subtidal

depths during the winter (Batchelder, 1915; Dexter,

1961; Brenchley, 1980), the depth of migration is likely

to be limited bv tidal range and wave energy. For ex-

ample, in a microtidal coastal setting (mean tidal range:

0-2 m) where tidal flats are fronted by wave-dominated

coastal barriers, the vertical habitat range of I. obsoleta

will be relatively narrow (Whitlatch, 1982). This fact,

combined with occurrence of the eastern mud snail in

post-glacial deposits (past 10,000-15,000 years; Carlton,

1992), makes this species a potential indicator of former

sea level.

The vertical range of a particular plant or animal spe-

Corresponding author: Ilya Buynevich

email: [email protected]

cies preserved in a geological record and its position

relative to a specific tide len el (indicative meaning; van

der Plassche, 1986; Donnelly et al., 2004) can he esti-

mated based on modem ecological response of that or-

ganism to tidal inundation. Therefore, comparison of in-

situ shells of I. obsoleta with an adjacent reliable sea-level

indicator, such as high-marsh peat, is the first step in

assessing its paleoenvironmental significance. The aim of

this paper is to use similar ages of L. obsoleta and high-

marsh peat in submerged banlianrer deposits offshore

Martha's Vineyard wane, Massachusetts, as evidence of

a potential use of this gastropod as an independent sea-

level indicator.

The microtidal barrier coastlines of southern New En-

gland and Long Island have been formed in a regime of

post-glacial marine transgression, with Holocene sedi-

mentary sequences now submerged on the inner conti-

nental shelf (R ampino and Sanders: LOSO: FitzGerald et

al., 1994: Schwab et al., 2000). The low mean tidal range

(0.7 m) and unlimited fetch offshore Martha’s Vineyard

island have combined to produce a high-energy, wave-

dominated environment (Figure 1). Furthermore; the

existence of a large glacial sand source has been condu-

cive to the formation of coastal barriers, which at lower

stands of sea level protected muddy coastal bays and

fringing saltmarshes (Oldale, 2001). A recent geophysical

study of the seafloor offshore Martha’s Vineyard (Goff et

al., 2005) offered an opportunity for study of the sub-

merged Holocene coastal deposits that contained re-

mains of shallow-water macrofauna, including I. obsoleta

High-resolution seismic imaging of the seafloor and

vibracores, ranging in depth of penetration from 0.5 to

approximately 2.0 m, were used to delineate a large sub-

merged pe aleo-valley (Figure 1; Buynevich et al., 2002).

The main valley (width: 300-500 m: de »pth: 1.5-7.0 m)

the offshore extension of Edgartown Great Pond, one of

many proglacial spring-sapping valleys incise sd into the

late Pleistocene glacial outwash deposits (Uchupi and

Oldale, 1994). All of the sediment cores that penetrated

marine sands and sampled organic-rich mud facies are

confined to the paleo-valley. These deposits contain vari-

Page 38 THE NAUTILUS, Vol. 121, No. 1

(east valley mean tidal range

margin)

13 1.0

ca. 6,200 years ago

44 + 2.0

depth

below

present

MHW

(m) 16

seafloor

Martha’s

Vineyard

6,230+60 calBP

5km paleo-valley

aa transect

1com

2.0

eS coarse sand (variable gravel content)

eS fine-medium marine sand

ee black backbarrier mud with in-situ shells

high-marsh peat

1400 2100 3100 3900

distance from shoreline (m)

Figure 1. Geological section along the axis of the main valley offshore Martha’s Vineyard (see inset for location) showing the

occurrence of black, organic-rich backbarrier muds underlying marine sands. These facies contain in-situ gastropods and biv alves and

have been sampled to a depth of at least 1S m below present sea level. Photographs: A) Iyanassa obsoleta shells encased in black

mud (one shell was used for dating). Saltmarsh peat recovered in the adjacent core 1 has a similar age and both dates are used to

constrain the position of mean high water ca. 6,200 years ago. B) cleaned I. obsoleta shell from core 13 (sample depth: ~15 m below

present se a level)

able amounts of organic material, occasional small bur- constrained portion of the regional sea-level history, but

rows, as well as whole shells of I. obsoleta (Figure 1, fit well within the regional sea-level envelope of Oldale

photos) and shallow-water bivalves. One core through and O'Hara (1980), as well as New York and New Jersey

the valley margin retrieved a short section of saltmarsh shelf data (Stuiver and Daddario, 1963; Rampino and

peat, which suggests a low-energy backbarrier setting Sanders, 1980). The similarity between the ages of high-

with saltwater access through a tidal inlet. marsh peat and that of eastern mud snail, along with the

The age of an in-situ I. obsoleta shell from core 2.4 (14 geological context and ecology of I. obsoleta, demon-

m below present mean high water [MHW]) was deter- strates the potential of this species as a paleo-sea-level

mined with accelerator mass spectrometry (AMS) radio- indicator. Although the elevation of muddy sediments

carbon dating and compared to that of a mee ee peat and macrofaunal remains relative to contemporaneous

from core | (13.5 m below MHW). A date of 6,230 + 60 sea level is less constrained than that of high-marsh peat,

cal BP (calibrated years before present; 20 error) i‘ the their association with peat and great cross-shore extent

shell provides an age for the minimum elevation of mean make them additional points of reference for the position

high tide level in this part of the valley fill The age of of sea level. Substantial compaction by loading and vi-

6,245 + 75 cal BP on the saltmarsh peat (8'°C= —15.4%o) bracoring action is unlikely due to relatively small thick-

further constrains the paleo-MHW elevation at this lo- ness of overlying sediments and basal positions of dated

cation (Figure 1). Both dates plot in a relatively poorly material in both cores, respectively,

SSS SSS SSS SS SSS

I MV: Buynevich, 2007

Page 39

This study demonstrates that in areas where occur-

rence and thickness of peat are limited, such as in mi-

crotidal settings, I. obsoleta-bearing muds may prove to

be reliable indicators of past water levels, energy condi-

tions, and former shoreline positions. In particular, in-

tertidal and shallow subtidal portions of submerged val-

ley fills provide an ideal geological setting for preserving

a nearly continuous sequence of back barrie ccameal

that can be used for paleoenvironmental and sea-level

reconstruction.

ACKNOWLEDGMENTS

The study was funded by the Office of Naval Research

and the Coastal Ocean Institute of the Woods Hole

Oceanographic Institution. Author wishes to thank John

Goff, Roy Wilkens, Rob Evans, Peter Traykovski, Chris

Jenkins, and the crew of R/V Care HENLOPEN for assis-

tance with coring operations and Heidi Fuchs for eco-

logical insight. Michael Savarese provided helpful com-

ments on the manuscript.

LITERATURE CITED

Batchelder, C. H. 1915. Migration of Ilyanassa obsoleta, Lit-

torina littorea, and Littorina rudis. The Nautilus 29: 43-46.

Brenchley, G. A

Ilyanassa obsoleta in Barnstable Harbor. Biological Bulle-

tin 159: 456-457.

Brenchley, G. A. 1982. Predation on encapsulated larvae by

adults: effects of introduced species on the gastropod Iya-

nassa obsoleta. Marine Ecology Progress Series 9: 255-262.

Buynevich, I. V, R. L. Evans, S. Schock, R. Wilkens, J. A. Goff,

P. A. Traykovski, C. Jenkins, G. Quentin, P. Beaujean, J.

Wulf, C. Vaczo, H. Nieto, and H. Gittings. 2002. Geom-

etry and sedimentary characteristics of a submerged pro-

glacial spring-sapping valley, offshore Martha’s Vineyard,

Massachusetts. Eos Transactions, AGU, 83(47), Fall

Meeting Supplement, Abstract OS61A-0156.

Carlton, J. T. 1992. Introduced marine and estuarine mollusks

of North America: an end-of-the-20'"-century perspective

Journal of Shellfish Research 11: 489-505.

Culbert. \W. and L. Raleigh. 2001. The Ecology of Coastal Salt

Ponds: A Pilot Study at Long Point Wildlife R efuge, West

Tisbury and Chilmark, Martha’s Vi ineyard. The Trustees of

Reservations, V ineyard Haven, Massi ichusetts, 74 pp.

Dexter, R. W. 1961. Mass movement of a colony of mud snails,

Ilyanassa. The Nautilus 75: 85-S6.

Donnelly, J. P.. P. Cleary, P. Newby, and R. Ettinger. 2004.

iG oupling instrumental and geological records of sea-level

change: evidence from southern New England of an in-

. 1980. Distribution and migratory behavior of

crease in the rate of sea-level rise in the late 19"" century.

Geophysical Research Letters 31, L05203: 4 pp.

FitzGerald, D. M., P. S. Rosen, and S. van Heteren. 1994. New

England Barriers. In: Davis, R. A., Jr. (ed.) Geology of

Holocene barrier island systems. Springer-Verlag, pp.

3 eae ‘

Goff, J. A., L. Mayer, P. A. Traykovski, I. V. Buynevich, R

Wilkens. R. Raymond, G.G lang, R. L. Evans, H. Olson,

and ©. Jenkins. 2005. Detailed inve stigation of sorted bed-

forms, or “rippled scour depressions”, within the Martha's

Vineyard Coastal Observatory, Massachusetts. Continental

Shelf Research 25: 461-484.

Kelaher, B. P. J. S. Levinton, and J. M. Hoch, 2003. Foraging

by the mud snail, Myanassa obsoleta (Say), modulates spa-

tial variation in benthic community structure. Journal of

Experimental Marine Biology and Ecology 292: 139-157.

Oldale, R. N. 2001, Cape Cod, Martha's Vineyard, and Nan-

tucket: The Geologic Story, 2"4 Edition, On C Jape Publi-

cations, 224 pp.

Oldale, R. N. and C. J. O'Hara. 1980. New radiocarbon dates

from the inner continental shelf off southeastern Massa-

chusetts and local sea-level rise curve for the past 12,000

years. Geology 8: 102-106,

Race, M. S. 1982. Competitive displacement and predation

between introduced and native mud snails. Oecologia 54:

337-347.

Rampino M. R. and J. E. Sanders. 1980. Holocene transgres-

sion in south-central Long Island, New York. Journal of

Sedimentary Petrology 50: 1063-1080.

Robertson, J. R. 1979. fivdeave for tidally correlated feeding

rhythms in the eastern mud snail, Myanassa obsoleta. The

Nautilus 93: 3540.

Stuiver, M. and J. J. Daddario. 1963. Submergence of the New

Jersey coast. Science 142: 941.

Schwab, W. C., E. R. Thieler, J. R. Allen, D. S. Foster, B. A.

Swift, and F. Denny. 2000. Influence of inner-continental

shelf geologic frame work on the evolution and behavior of

the barrier-island system between Fire Island Inlet and Shin-

necock Inlet, Long Island, New York. Journal of Coastal Re-

search 16: 408-422.

Uchupi, E. and R. N. Oldale. 1994. Spring sapping origin of the

enigmatic relict valleys of Cape Cod, Martha’s Vineyard

and Nantucket Islands, Massachusetts, Ge omorphology 9:

$395.

van der Plassche, O. 1986. Introduction. In: Van der Plassche,

O. (ed.) Sea-level Research: A Manual for Sample Collec-

tion and Evaluation of Sea Level Data. Norwich, Geo

Books, 1-26.

Weiss, H. M. 1995. Marine animals of Southern New England

and New York: Identification keys to common nearshore

and shallow water macrofauna. Bulletin 115, State Geo-

logical and Natural History Survey of Connecticut, 709 pp.

Whitlatch, R. B. 1982. The ecology of New England tidal flats:

a community profile. U.S Fish and Wildlife Service, Bio-

logical Services Program, Washington, D.C., FWS/PBS-

81/01, 125 pp.

THE NAUTILUS 121(1):40, 2007 Page 40

Errata

In the last issue of The Nautilus, in the article by Thomas J. DeVries (2006), please substitute the specific epithet

meleani for stucchii:

On page 139, left-hand column, second paragraph, second line;

On page 140, legend of Figure 2, first line;

On page 146, left-hand column, sixth paragraph (“Remarks”), first line.

Due to an editorial lapse, the word stucchii was used unintentionally in these three instances.

LITERATURE CITED

DeVries, T. J. 2006. The Neogene history of Prisogaster Mérch, 1850 (Gastropoda: Turbinidae) in South America. The Nautilus 120:

139-149.

THE NAUTILUS 121(1):41, 2007 Page 41

Notice

THE 2007 R. T. ABBOTT VISITING CURATORSHIP

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The Curatorship, established originally in accordance with the wishes of the late Dr. R. Tucker Abbott, Founding Director of the

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percentage of our holdings have been catalogued through a computerized database management system; part of the catalogue is

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Interested malacologists are invited to send a copy of their curriculum vitae together with a letter detailing their areas of taxonomic

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Applications for the 2007 Visiting Curatorship should be sent electronically to the above e-mail address no later than May 30, 2007,

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Tae NAUTILUS

Volume 121, Number 2

June 27, 2007

ISSN 0028-1344

A quarterly devoted

to malacology.

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Volume 121, Number 2

June 27, 2007

x T 9Q_72

CONTENTS ISSN 0028-1544

Ellen E. Strong On the anatomy and systematics of Juga from western North America

Terrence J. Frest (Gastropoda: Cerithioidea: Pleuroceridae) .. 0... ee 43

Timothy A. Pearce Discriminating shells of Gastrocopta pentodon (Say, 1822) and G.

Marvin C. Fields tappaniana (C. B, Adams, 1842) (Gastropoda: Pulmonata) with an example

Kayoko Kurita from the Delmarva Peninsula, eastern USA... 0. 66

M. G. Harasewych Upper Jurassic Pleurotomariidae (Gastropoda) from

Steffen Kiel southwestern Madagascar ... 1... ee 76

M. G. Harasewych Sassia melpangi, a new ranellid species (Gastropoda: Tonnoidea) from

Alan G. Beu the Central Pacific 6... ee eee 90

José Carlos N. de Barros A new species of Microcancilla (Gastropoda: Cancellariidae) from the

Richard E. Petit continental slope off northeastern Brazil... 0.00. 95

Silvio Felipe B. de Lima A new species of Gerdiella (Gastropoda: Cancellariidae) from the South

José Carlos N. de Barros Atlantic Ocean off Brazil with discussion of an undescribed species ...... . 99

Richard E. Petit

Book: Review? .(¢ 5.6 4 4 ced oe b4.h4. bs Sameer d ob ew Oe el embed Cwregd e544 G89 phew. ea sede 5 104

Sponsored in part by the State of Florida, Department

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THE NAUTILUS 121(2):43-65, 2007

Page 43

On the anatomy and systematics of Juga from western North

America (Gastropoda: Cerithioidea: Pleuroceridae)

Ellen E.

Department of Invertebrate Zoology

National Museum of Natural History

Smithsonian Institution

P.O. Box 37012

Washington, DC 20013-

[email protected]

Strong

Deixis Consultants

7012 USA

Terrence J. Frest

2517 NE 65th Street

Seattle, WA 98115 USA

[email protected]

ABSTRACT

The family Pleuroceridae is a speciose and ecologically impor-

tant family of limnic gastropods in North America and eastern

Asia. Juga is the only native pleurocerid genus that occurs in

Pacific drainage systems of Western North America, but has

only recently been accepted as independent from other North

American genera and may have affinities to Asian pleurocerids.

As such, this genus represents a key piece to the puzzle of

pleurocerid systematics. However, published accounts of Juga

anatomy are limited to the reproductive system, Consequently,

the anatomy of three species is describe d herein: these three

taxa represent the type species of the three extant subgenera:

Juga (Juga), J. (Calibasis), and J. (Oreobasis). A lectotype is

designated for Goniobasis acutifilosa Stearns, 1890, the type

species of Calibasis; Melania newberryi, the type species of

Oreobasis, is here removed from the synonymy of Juga bulbosa.

This analysis confirms that Juga shares many anatomical fea-

tures with other North American and Asian pleuroce srids, but is

clearly set apart from eastern North American pleurocerids in

features of the ovipositor pore, radula, midgut, kidney, and

pallial gonoduct. Juga is distinct from all other limnie cerithio-

ideans Teno thus far in the form of the midgut crescentic

ridge, the configuration of prostate glands, and an evagination

of the kidney wall separating the main chamber and bladder.

Based on information currently available, unlike molecular

data, there is no morphological feature unambiguously linking

Juga to Asian pleurocerids. Anatomically, Ore obasis is strikingly

similar to Juga sensu stricto and is synonymized with it, whereas

Calibasis is retained as a valid taxon.

INTRODUCTION

The Pleuroceridae Fischer, 1585, is a speciose and eco-

logically important family of limnic gastropods occurring

in North America and Eastern Asia. Des spite their im-

portance. understanding of their systematics is discour-

agingly incomplete. The current classification of limnic

lineages within the Cerithioidea Fle ming, 1822, is rooted

in the works of Thiele (1928, 1929). ake subdivided the

heterogeneous Melaniidae Children, 1523 (an invalid

name for Thiaridae Gill, IS71) into six subfamilies in-

cluding the Pleurocerinae. An alternative classification

advanced by Morrison (1954) distinguished only three

limnic families: (i) the Pleuroceridae distributed

throughout the Americas, Africa, and Asia, (ii) the Mel-

anopsidae in Europe, and (iii) the pantropical Thiaridae.

While promoting the important notion of several inde-

pendent lineages, this concept heavily weighted plesio-

morphic (ovipositor and oviparity) and homoplastic

(brooding) features and resulted in a highly polyphyletic

Pleuroceridae—a view that persisted for over three de-

cades (e.g. Ponder and Warén, 1988). Recent work has

resolved some of this confusion and supports the distinc-

tiveness of many of Thiele’s groupings (e.g. Glaubrecht,

1996, 1999; Ly deard et al., 2002: Kohler and Glaubrecht,

2001, 2003; Strong and Glaubrecht, 2002, 2003; Kohler

et al., 2004; von Rintelen and Glaubrecht, 2005). How-

ever, molecular data (Lydeard et al., 2002) do not sup-

port monophyly of the Pleuroceridae as currently de-

fined (e.g. Bouchet and Rocroi, 2005). The analysis of

Houbrick (1988) based on morphological data did not

include sufficient taxon sampling to adequately assess

monophyly of the family.

In North America, ple urocerid diversity is highest east

of the continental divide where they are represented by

seven genera (Athearnia Morrison, 1971, Elimia H. and

Adams, 1854, Io Lea, 1831, Leptoxis Rafinesque,

1819, Lithasia Haldeman, 1840, Pleurocera Ratinesque,

1818, and the extinct Gyrotoma Shuttleworth, 1545) and

72 aaa ‘ly 148 species currently considered valid

(Johnson et al., 2005). Juga H. and A. Adams, 1554, with

11-12 estimated valid species (Burch, 1989; Turgeon et

al., 1998; Johnson et al., 2005),

rocerid genus that occurs in the Pacific and Interior

drainages from central California to central Washington

(Figure 1). Established on the basis of early teleoconch

shell sculpture, four subgenera are recognized (Tayloi

1966; Burch, 1989)—three are extant: Juga H. and A

is the only native pleu-

Page 44

THE NAUTILUS, Vol. 121, No. 2

Figure 1. Black line indicates currently

Distribution of juga. B

known area of contiguous distribution. Black circles represent

isolated sites bey ond main distribution. Black squares are loca-

tions of extant subgeneric type localities and of material used in

this investigation.

Adams, 1554 (with plicate early sculpture), Calibasis

Taylor, 1966 (with lirate early sculpture), and Oreobasis

Taylor, 1966 (with weak to no early sculpture). Idabasis

Taylor, 1966 (with plicate and lirate early sculpture) is

known only from fossils. Some east Asian species have

also been assigned to Juga, but more recently these have

been reassigned to Parajuga nate a and Staroboga-

tov, 2004 ( Starobogatov et al., 2004): however, this name

is unavailable from a hanes standpoint, as no

type species was designated.

Juga has no discrete conchological feature that distin-

guishes it from eastern North American genera and has

often been synonymized with Goniobasis Lea, 1862 (a

na synonym of Elimia) (e.g. Tryon, 1865, 1873; Pils-

bry, 1 899: Walker, 1918: Henderson, 1935a, b: Goodrich,

1942). Taylor (1966) was the first to restore Juga as a

valid genus in the modern literature, primarily on the

basis of its disjunct biogeographic distribution and the

presence of a distinctive ovipositor (see Discussion, be-

low). Based on recent molecular (Holznagel and Ly-

deard, 2000; Lydeard et al., 2002) and morphological

findings (Prozorova and Raschepkina, 2004; Strong,

2005). Juga is supported as distinct from other North

\merican pleurocerids and may have ties to those from

eastern Asia (see Discussion, below). As such, this genus

represents an important and intriguing part of the puzzle

of pleurocerid systematics.

Thus, the goal of this analysis is to establish the ana-

tomical or ganization for the type species of the current

extant subgeneric subdivisions within Juga:

Juga H. ‘and A. Adams, 1854: type species (by subse-

quent designation Baker, 1963) Melania silicula Gould,

1847.

Calibasis Taylor, 1966: type species (by original des-

ignation) Goniobasis acutifilosa Stearns, 1890.

Oreobasis Taylor, 1966: type species (by original des-

ignation) Melania newberryi Lea, 1560.

This also represents the first comprehensive anatomi-

cal investigation of any Juga species; the only information

published thus far concerns female reproductive

anatomy (Prozorova and Raschepkina, 2004). In the con-

text of ongoing morphological and molecular phyloge-

netic analyses of pleurocerids and cerithioideans in gen-

eral, this information is vitally important for assessing Fe

affinities and relationships of taxa currently placed in the

Pleuroceridae.

MATERIALS AND METHODS

Specimens for morphological study were collected by

hand or dip net, stored cold until evening, relaxed with

menthol in shallow water from the same spring or stream

in a broad, open container overnight, and transferred to

dilute cold 4-5% formalin the next morning. Specimens

were maintained in formalin for 48 hours then trans-

ferred to buffered 70% ethanol/10% glycerin/20% water

for longer term storage. Voucher wiateral’ is deposited in

the National Museum of Natural History in Washington,

DC, (USNM) and with Deixis Consultants in Seat-

tle, WA.

Specimens were examined using a Leica MZ 12,5 bin-

ocular microscope with camera lucida: visualization of

structures was enhanced with aqueous toluidine blue.

Typically 2 to 4 specimens were examined for each organ

system investigated, particularly for complex structures

(i.e. midgut, nerves), as well as to assess intra-specific

and/or cenconel variation in reproductive anatomy.

A comprehensive anatomical account is provided for

Juga (Juga) silicula—the type species of the genus. Only

discrete differences are detailed for J. ( (Calibasis) ) acuti-

filosa and J. (Oreobasis) newberryi with comparative re-

marks highlighting qualitative differences. As far as can

be detenmiied: near ae material was used for this

investigation (see details below). A thorough systematic

treatment of each species is not provided as a phyloge-

netic analysis and comprehensive revision of the genus

are forthcoming (Frest et al., unpublished data).

Geographic names, road names and numbers, and

land ownership data were confirmed using the DeLorme

Mapping Washington, Oregon, and Northern California

Atlas and Gazetteer, the late »st available USGS 7.5’ series

topographic maps, and National Geographic TOPO!

2006. Universal Transverse Mercator (UTM) grid coor-

E. E. Strong and T. J. Frest, 2007

Page 45

dinates are based on NAD27 (1927 North American Da-

tum). Locality descriptions have been downloaded from

Deixis Consultants MolluscDB!'™'. Collector abbrevia-

tions are as follows: EJ, Edward J. Johannes; TF, Ter-

rence J. Frest.

Institution codes cited in the text are: USNM: Na-

tional Museum of Natural History, Smithsonian Institu-

tion, Washington, DC; MCZ: Museum of Comparative

Zoology, Harvard,

RESULTS

Pleuroceridae Fischer, 1885

Juga (Juga) silicula (Gould, 1847)

Type Material: Three syntypes (USNM 12137) (cited

as MCZ 12137 in Graf, 2001) (Figures 2-4). Johnson

(1964) indicated that the largest (Figure 3) might be the

specimen illustrated by Gould ( 1852, 1856, pl 10, figs.

164, 164a); however, in tinct of whorls and overall

size, the figured specimen is most likely the smallest

syntype (Figure 2). As the figured specimen represents

the smallest syntype, and the largest specimen mostly

lacks the distinguishing axial ornament characteristic of

juvenile shells, a lectotype is not here selected.

Type Locality: Indicated as “Nisqually, Oregon”

(Gould, 1847) (see Figure 1). These specimens likely

ae have been Gollented by the Wilkes Expedition,

1838-1842, before the separation of Washington Terri-

tory from the larger Oregon Territory in 1853. This ex-

pedition started from Fort Nisqually, which was located

in the current Washington State (still part of Oregon

Territory in 1547). \. There was no rail station or city with

the name “Nisqually” at that time. Thus, the river or the

fort would be indicated. Johnson (1964) reported the

type locality as “near present site of Tacoma, Washing-

ton” (also perpetuated by Graf, 2001), which is loosely

true but unnecessarily vague and misleading as Tacoma

lies outside the northern range of Juga, w hich ends just

north of the Nisqually River (Pierce-Thurston Co.

line)—by no mere coincidence near the approximate

southem terminus of Late Wisconsinan glaciation.

The Nisqually River is a glacial flour stream originat-

ing on Mount Rainier with a depauperate fre shwate oT

mollusk fauna, despite various literature ascriptions, Its

tributaries are comparatively rich, however, especially as

they approach Puget Sound. “Nisqually” thus likely re-

fers to Fort Nisqually *, one of two trade outposts of the

Hudson Bay oa founded in 1833 on the Nisqually

Reacl 1 dive -ctly south of Sequalitchew Creek (Hitchman,

1 Puget Sound tributary northeast of the

de River. In 1843 the operation was moved about

Dupont, Washington (Phillips, 1997), roughly in the

same drainage. For quite some time, this was the only

settlement in the area, so that the origin of Gould’s speci-

mens could be Sequalitchew Creek or one of the other

nearby tributaries of the Nisqually River. Attempts to

2 miles northeast, to approximately Ny present site of

collect Juga from the boggy Sequalitchew Creek have

been unsuccessful (Frest, unpubl. data). However, ma-

terial from nearby McAllister Creek, also a Puget Sound

tributary just west of the Nisqually River, has young

specimens most closely resembling the types found thus

far.

Remarks: Evidently a valid species (see also Good-

rich, 1942: Burch and Tottenham, 1980: Burch, 1982a, b,

1989; Graf, 2001) but perhaps with a rather restricted

range, confined to a few streams on the southeastern end

of Puget Sound, near the Nisqually River.

Material Examined: Washington: Thurston County:

McAllister Creek at eine Road bridge, near McAl-

lister Creek Hatchery, depth 0-1.22 m (Zone 10

520890E 5210540N [122°43'34”"E, 47°02'59’N],

Nisqually 1994 7.5! quadrangle, elevation 1.5 m), Deixis

Consultants locality #5709, 29 Sep. 2005, collected by

TF, EJ (USNM 1100657) (9 specimens dissected) (see

Figures 7-9).

External Anatomy: Operculum ovate, corneous, dark

reddish brown in color, with 3.5 whorls; paucispiral with

eccentric nucleus of approximately 3 whorls (Figure 16).

Nucleus comprising slightly less than one half of total

length (~45%).

Head- foot dark gray to black in color, with lighter gray

snout tip and and pale. foot sole; in females, ovipositor

groove pale in color. Foot broad with wide propodium

and long anterior pedal gland along anterior margin (Fig-

ure 17, ap). Ciliated egg groove forming longitudinally

groovy ed tract extending fon anterior tip of palliz il gon-

oduct (go) and broadening continuously into shallow: tri-

angular shelf on side of neck below right cephalic ten-

tacle (ovp). Ovipositor surface groovy ad. with folds di-

rected medially into pore. Shallow grooved tract

extending from pocket to edge of foot, “short distance

back from anterior pedal gland. Extensible snout (Fig-

ures 17, 18, sn) broad, squarish, with short triangular

cephalic tentacles (t); tentacles also short in relaxed

specimens.

Ctenidium (Figures 18, 19, 26, et) extending from pos-

terior end of mantle cavity to near mantle edge, anteri-

orly curving toward the left. Osphradium forming simple

ridge alongside efferent branchial vessel, sometimes with

anterior tip markedly curving toward the left; osphra-

dium undulating slightly at anterior and posterior ends,

aan aes (os). Hypobranchial gland moder-

ately well developed with transverse ridges, especially at

posterior end of mantle cavity (Figures 1S, 19, hg).

aca | System: RADULA: Radula comprising ~98

rows (n = 2) (Figures 20-25). Rachidian bro: idly rectan-

gular, rr than tall, with smoothly rounded v-shaped

lower margin and single small basal denticle at each

lower, outer corner (Figure 22). Upper margin slightly

concave with cutting edge bearing one large central tri-

angular, spatulate cusp, and three stout, conical denticles

on each side. Lateral teeth (Figures 20, 21, 23) with

moderately short lateral extensions (slightly more than

THE NAUTILUS, Vol. 121, No. 2

1S56 pl 10 figs 164. 164a

Shells of Juga species. 2-4. Juga (Juga) silicula (USNM 12137, syntypes). Figure 2 is likely the figured specimen

uga (Calibasis) acutifilosa (USNM 60596 lectotype Stearns, 1S90 pl 15. fig. 9). 6.

Juga (Oreobasis) newberryi (USNM 118961, lectotype). 7-15. Material examined in morphological analysis. 7-9. Juga (Juga) silicula

USNM 1100659). 10-12. Juga (Calibasis) acutifilosa (USNM

Sc cl

lf the length of lateral cutting edge), and single promi-

itulate cusp flanked by two inner and two to three

nticl Outermost denticle weakly developed

nr shape ize, and position (Figure 23

Figures 24, 25) with broadly rounded

lon lender shafts. Narrow flanges

1100658). 13-15. Juga (Oreobasis) newberryi (USNM_ 1100660

developed on outer edges of marginal teeth shatts along

distal one half to two thirds. Outer flange much wider

and longer on inner marginal tooth; outer flange only

slightly wider on outer marginal tooth. Inner marginal

teeth with six and outer marginal teeth with seven flat

tened denticles

E. E. Strong and T. J. Frest, 2007 Page 47

‘int

OOO NET ek Or

tA UWI

0s-\————

Eee)

Figures 16-19. External anatomy of Juga (Juga) silicula (USNM 1100659). 16. Operculum. 17. Ovipositor and egg groove. Right

lateral view of head-foot. 18-19. External view of organs in visceral mass. Dotted line indicates extent of pericardium under main

kidney chamber (Figure 15). Abbreviations: ag, albumen gland; ap, anterior pedal gland; b, bladder; eg, capsule gland; em,

columellar muscle: et, ctenidium; dg, digestive gland; f, foot; go, pallial gonoduct; hg, hypobranchial gland; int, intestine; kd, main

kidney chamber; me, mantle edge; op, operculum: os, osphradium; ov, ovary; ovp, ovipositor; per, pericardium; sn, snout; ss, style

sac: sto, stomach: t, cephalic tentacle. Scale bars = 1 mm.

Page 45

THE NAUTILUS, Vol. 121, No. 2

Figures 20-25. Radula of Juga (Juga) silicula (\USNM 1100659). 20. Section of anterior radular ribbon. Scale bar = 200 jm. 21.

Rachidian and lateral teeth. Scale bar =

50 pm. 22. Detail of rachidian teeth. Scale bar = 20 pm. 23. Detail of lateral teeth; note

weakly formed outer third denticle. Scale bar = 50 zm. 24. Marginal teeth. Scale bar = 50 wm. 25. Detail of cutting edge of marginal

teeth. Note unequal size of cusps on inner and outer teeth. Scale bar = 20 jm.

Forecut: Buccal mass short and robust (Figure 26, bm).

Odontophore occupying posterior one half to two thirds

of buccal cavity with small, glandular subradular organ

protruding before radula. Small jaws present at anterior

ends of dorsal folds; epithelium of buccal cavity between

dorsal folds glandular (stippled). Dorsal folds deeply cleft

along midline adjacent to odontophore; cleft receives

salivary gland ducts at posterior end and shallows ante-

riorly. Very shallow, non- -glandular buccal pouches ex-

page underneath Bosal folds adjacent to buccal gan-

lia (bg) at rear of buccal cavity. Radular sac (1s) shit

a 4.4 mm), curving upward behind base of buccal mass,

not passing through nerve ring, with tip resting against

right posterior Sad of buccal mass. Robust buéeal retrac-

tors (rt) inserting onto lateral walls of cephalic hemocoel

anterior to nerve ring. Short, glandular mid-ventral fold

forming small triangular ridge just behind odontophore

in anterior esophagus, fiesled by two ventro-lateral folds.

Ventro-lateral folds converging short distance behind mid-

ventral fold, forming rear-facing triangular depression sur-

rounding mid-ventral fold. Paired dorsal and ventral folds

continuing through long mid-esophagus (e) into posterior

esophagus. Epithelium of mid-esophagus between dorsal

and ventral folds longitudinally grooved; septate esophageal

gland lacking. Posterior esophagus narrow, with numerous

folds of approximately equal height. Long, tubular salivary

glands (sg) passing through circum- esophageal nerve ring,

extending . posterior esophagus.

Mipcut: Esophagus opening under ledge on left side of

midgut floor (Figure 27, e). Marginal fold (mf) extending

anteriorly from esophageal aperture alongside major

typhlosole (t1), then turing posteriorly, bordering right

margin of sorting area (Sa). Sorting area elongate, rect-

angular, tapering posteriorly; posterior tip curving

slightly to the left around crescent-shaped sorting area

pad (sap). Accessory marginal fold (amf) forming. weak

ridge paralleling marginal fold from near esophagus,

curving around posterior margin of sorting area; poste-

rior segment of fold variable, in some specimens inter-

secting straight longitudinal ridge along left posterior

end of sorting area (figured), in other specimens weakly

bifurcate (see e.g _ Figures s 37, 51). Fine parallel striations

extending anteriorly from esophagus up face of major

typhlosole (Figure 27, tL). Midgut roof to the left of

sorting area coarsely folded and cuticularized (eu). Gas-

tric shield (gs) broadly concave; shield continuous with

cuticle of adjacent oe of stomach roof and floor.

Glandular pad (gp) large, rounded posteriorly, with

lightly textured surface. Slight ov i: lip of glan-

delar pad forming shi low pocket ( y behind gastric

shield. Crescentic. ridge (er) ee wide, shallow

crescentic groove. Proximal end of ridge posteriorly bor-

dering deep pouch that receives multiple openings of

digestive gland (dd); distally, ridge fusing to right, pos-

terior end of glandular pad. Size of opening to digestive

eland duct ve stibue variable. Single, weak, irregular lon-

citudinal fold (ef) along floor opposite caecum. Promi-

nent ioe fold (uw) extending from beneath right

side of style sac lip (ss), along flog. to ridge extending

from base of major ty phlosole: fold bounding u- -shaped

depression below lip of style sac. Style sac large, com-

municating along entire length with intestinal eroove;

intestine forming prominent protuberance at distal tip of

E. E. Strong and T. J. Frest, 2007 Page 49

Figures 26-29. Intemal anatomy of Juga (Juga) silicula (USNM 1100659). 26. Mantle cavity and anatomy of cephalic hemocoel.

Dorsal view, anterior is below. 27. Midgut anatomy. Dorsal view, anterior is uppermost. 28. Kidney anatomy. Internal view of

bladder. Lateral view, anterior is uppermost. Roof of bladder (below intestine) cut open to reveal interior; adjacent to incision,

stippling indicates intersection of Ory tubules with roof of bladder. Arrow indicates opening in outpocketing of wall between

bladder and main kidney chamber. 29. Circum-esophageal nerve ring. Frontal view on the left, right lateral view on the right. Arrow

indicates connective to supra-esophageal ganglion. Abbreviations: amf, accessory marginal fold; arv, afferent renal vessel: bg, buccal

ganglion: ec, caecum: ce, cerebral ganglion: cf, caecal fold: co, thickened connective between left pleural and sub-esophage sal vaneglia;

cr, crescentic ridge; et, ctenidium; cu, cuticularized region of stomach roof; d, dialyneury; dd, digestive gland duct vesuivule: e,

esophagus; go, pallial gonoduct; gp, glandular pad; gs, gastric shield; int, intestine; kd, main kidney chamber: me, mantle edge; mf,

marginal fold: np, nephropore: os, osphradium; pe, pedal ganglion; pl, pleural ganglion; r, rectum; rs, radular sac; rt, buccal

retractor muscle; sa, sorting area; sap, sorting area pad; sb, sub- -esophageal ganglion; sg, salivary gland; sp, supra-esophageal

ganglion: ss, lip of style sac: st, statocyst: t, cephalic tentacle; t1, major typhlosole; u, u-shaped fold; z, zygoneury. Scale bars = 1 mm

Page 50

THE NAUTILUS, Vol. 121, No. 2

style sac where it separates from the latter (not visible

dorsally; see e.g. Figure 50, ss). Crystalline style present.

HINDGUT: Proximal intestine (Figures 18, 19, int) pass-

ing below distal tip of style sac, then extending posteri-

orly in broad loop, partially overlying style sac, to main

gastric chamber (sto). Intestine continuing forward,

passing under posterior end of main indaey chamber

(kd), entering pallial roof between bladder (Figure 19, b)

and main kidney chamber (kd), to papillate anus near

mantle margin (Figure 26, r).

Reno-pericardial System: Kidney comprising two

interconnected chambers (Figure ‘ 28). Main chamber

(kd) lying along dorsal surface of body whorl, anteriorly

surrounding pericardium (Figure 18, per, dotted line),

crossing axis of body from right to left and extending

short distance into pallial ry at base of mantle cavity.

Main chamber occluded with excretory tubules anteri-

orly (within pallial roof), posteriorly, and along left mar-

gin. Central portion of main chamber with small, narrow

Tannen Second chamber (exposed chamber in Figure 28)

extending between pericardial chamber to right body

wall below intestine, forming small bladder (Figure 19,

b). Wall separating main chamber and bladder forming

large outpocketing; small aperture within wall, just in

front of afferent renal vessel, connecting main chamber

and bladder (Figure 28, arrow). Bladder io gely occluded

by vertical sheets of excretory tissue radiating from af-

ferent renal vessel (arv). Sheets of excretory tissue

branching and anastomosing, and fusing to right lateral

wall, floor and roof below intestine; sheets laterally en-

closing outpocketing of main chamber wall. Bladder

communicating to mantle cavity via large nephropore

(np); outpoc ke ting of main chamber wall exte nding into

ne Bee restricting communication with m: antle ¢ cav-

ity. Bladder penetrating connective tissue along right

ee of body, short distance into mantle cavity.

Nephridial gland absent.

Pericardium long and narrow (Figure 1S, per, dotted

line), extending to recurved intestinal loop.

Nervous System: Circum-esophageal nerve ring

(Figure 29) lying immediately behind buccal mass, well

behind base of cephalic tentacles. Cerebral ganglia (ce)

connected by short, stout commissure, each producing

seven nerves (optic, statocyst, tentacular, and 4 labial

nerves). Buccal ganglia (Figure 26, bg) lying ventro-

laterally at base of buccal mass, adjacent to retractor

muscles, at outer edges of esophagus where it emerges

from buccal cavity. Pleural ganglia (Figure 29, pl) ) lying

behind and below cerebral vanglia, connected to cere-

bral ganglia by short, thick connectives. Pedal ganglia

pe) with two prominent anterior nerves and variable

number of smaller accessory nerves (five to seven). Sta-

toeysts (st) with numerous statoconia present dorsally

alongside pedal ganglia behind pedal connectives, Sub-

esophageal ganglion (sb) joined to left pleural canglio

by thickened connective (co); connective producing on

small nerve. In addition to connectives to right and left

pleural and visceral ganglia, sub-esophageal ganglion

producing four small nerves. Zygoneury (z) formed be-

tween sub-esophageal and right pleural ganglia. Long

connective uniting right pleural and supra- esophageal

ganglia (Figures 26, 29, sp), the latter lying on left side of

mentie floor near midline of osphradium. Dialyneury

formed between pallial nerve of left pleural ganglion and

osphradial nerve of supra-oesophageal ganglion at junc-

tion of mantle roof and floor. Single visceral ganglion

present between pericardium and kidney at base of

mantle cavity.

Reproductive System: FEMALE: Gonad (Figures 1S,

19, ov) dorsally surrounding digestive gland from tip of

visceral mass to posterior end of midgut (sto). Oviduct

emerging ventrally from ovary. Renal oviduct (Figure 30,

ovi) entering glandular pe allial oviduct at base of mantle

cavity. Palliat oviduct with proximal albumen (ag) and

distal capsule glands ( (cg). Proximal albumen gland, be-

low pallial portion of bladder, forming u- shaped tube

with glands developed dorsally along axis of fold and

simple gonoductal groove between opposing flattened

surfaces of glands. Anteriorly, albumen and capsule

glands becoming highly glandular and thrown into com-

plex undulating folds: gonoductal groove compressed

and highly eonvohite d (gg). Albumen gland approxi-

mately one third the length of capsule | land. Pallial ovi-

duct communicating with mantle cavity through narrow

aperture along entire length (arrow), except for tubular

section of alleunien gland at base of mantle cavity. Above

aperture, along anterior ~one fourth of f oviduct, deep

sperm gutter (sg, dashed line) present within medial

lamina; gutter opening broadly to long, broadly rounded

spermatophore bursa ( (spb). Bursa broadening posteri-

orly and extending to tubular portion of aibeanen gland.

Behind opening to bursa, sperm gutter becoming shallow

abruptly and continuing posteriorly ( (dashed line) as shal-

low groove. Near posterior end of oviduct, shallow ridge

entering small rounded aperture, just inside ventral edge

of eden lamina, leading to small pouch-like seminal

receptacle (res). Prominent glandular protuberance

from opposing inner surface of lateral lamina extending

into receptacle aperture, completely filling narrow proxi-

mal portion. Thin, narrow glandular ridge extending an-

teriorly from protuberance along inner edge of lateral

lamina; glandular ridge extending to pallial oviduct tip

just inside seminal groove.

Mate: Narrow vas deferens (Figure 31, 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 det-

erens narrowing and curving dorsally to enter posterior

end of prostate (pr) at base of mantle c ‘avity. Prostate

glandular, opening to mantle cavity through narrow ap-

erture along entire length except for a short fused seg-

ment at base of mantle cavity (Figures 31, 33, arrow).

Glands of medial and lateral laminae closely appressed,

forming tightly interlocking tongue and groove arrange-

E. E. Strong and T. J. Frest, 2007

Page 5]

Figures 30-33. Reproductive anatomy of Juga (Juga) silicula (L

Anterior is to the left. Arrow indicates posterior extent of opening to gonoductal groove. “< indicates transition between albumen

and capsule glands. 31. External, left lateral view of prostate. Anterior is to the left. Arrow indicates posterior extent of opening to

gonoductal groove. 32. Internal view of prostate medial lamina. Anterior is to the right. 33. Internal view of prostate lateral lamina.

Anterior is to the left. Arrow indicates posterior extent of opening to gonoductal groove. Abbreviations: ag, albumen gland; eg,

capsule gland: gg, gonoductal groove; ovi, renal oviduct; pr, prostate; res, seminal receptacle; sg, sperm gutter; spb, spe rmatophore

bursa; vd, vas deferens. Scale bars = 1 mm.

ment (Figures 32, 33). Glandular tissue of medial os

ventrally forming elongate, flattened textured fold (Fig-

ure 32), corresponding to concave a surface of op-

posing fold in lateral lamina (Figure 33). Medial lamina

expanding to surround fold within fe ral lamina. Dor-

sally and posteriorly, glands of lateral and medial laminae

flaring to form flattened flange; size and shape of flange

variable between individuals. Apart from fold in medal

lamina, inner surface of prostate essentially smooth.

Juga (Calibasis) acutifilosa (Stearns, 1890)

Type Material: Lectotype (USNM 60596; figured

specimen in Stearns, 1890, pl. 15, fig. 9) (Figure 5), by

present designation, in order to e ananee the stability of

the nomenclature in accordance with article 74.7.3 of the

ISNM 1100659), 30. External, left lateral view of pallial oviduct.

ICZN; indicated as holotype on label. Fourteen paralec-

totypes (USNM 60596X) in secondary type collection of

USNM, indicated as paratypes on old Jabel and as syn-

types on newer label printed in 2001. Although Graf

(2001) stated that the “holotype” is by original de ssigna-

tion, Stearns figured the largest of the syntypes and pro-

vided the dimensions, but did not make an explicit : pe

designation (designation of holotype) in the text and in-

dicated that the description was based on examination of

approximately three dozen specimens. Collected by

H. W. Henshaw.

Type Locality: “Eagle Lake” (Stearns, 1890). Taylor

(1981) corrected this to head of Willow Creek, Lassen

Co., California (see Figure 1). There are no populations

in the Eagle Lake drainage which includes several small

THE NAUTILUS, Vol. 121, No. 2

tributaries that flow into the lake during winter and the

only outflow is through the remnants of ihe Bly Tunnel.

Initiated during the 1920’ s, the Bly irrigation project di-

verted water from Eagle Lake to Willow Creek at Mur-

rers Meadows sev eal kilometers away in the more arid

Honey Lake drainage. Today, the ‘tunnel is mostly

blocked except for an 8’(~20 cm) pipe that still allows

some outflow to Willow Creek. The headwaters of Wil-

low Creek are regarded as springs along Murrer’s Upper

Meadow and Bly Tunnel (Moyle et al., 1996). Graf

(2001) lists the type locality as “Eagle Lake, [Lassen Co.,]

California’.

Remarks: The species varies widely in shell morphol-

ogy, from populations with several strong persistent lirae

to those with lirae confined to the adapical whorls. Body

color also may differ drastically from population to popu-

lation. Few sites show much intrapopulation variance;

but adults can vary from population to population, with

those at some sites highly variable and others essentially

invariant. Very few populations i in onlya part of the range

are as strongly and completely lirate as some Willow

Creek (and Murrer’s Upper Meadow) populations.

These observations are consistent with preliminary re-

sults based on COI sequences that indicate the species,

as currently recognized, is highly polyphyletic (Frest et

al., unpublished data).

Material Examined: California: Lassen County: ie

low Creek collected off dirt road (to E.), 0.40 kin S.

Murrer’s Lower Meadow, depth 0.05-0.20 m (Zone if

695000E 4493820N [120°41'51"E, 40°34'26"N], Gall-

atin Peak 1989 7.5’ quadrangle, elevation 1509 m),

Deixis Consultants locality #1484, 10 Sep. 1993, col-

lected by TF, EJ] (USNM 1100658) (4 specimens dis-

sected) (see Figures 10-12); southern-most of three

springs at N. end of ae Lower Meadow, below

road on E. side of meadow, E. of Eagle Lake, depth

0-0.03 m (Zone 10 694995E 4495225N [120°41'49”E,

40°35'12"N], Gallatin Peak 1989 7.5’ quadrangle, eleva-

tion 1545 m), Deixis Counsultants locality #1287, 10 Sep.

1993, collected by TF, E] (USNM 1100659) (3 speci-

mens dissected).

External Anatomy: Operculum ovate, with slightly

angular tip (Figure 34). Nucleus comprising slightly less

than one half of total length (~41%).

Ovipositor pore rather deep (Figure 35, ovp) with

deep, highly grooved tract extending to edge of foot,

slight doa. back from anterior pedal gland ( (ap).

Mantle edge (Figure 35, me) cre mulated, cor respond-

ing to spiral teleoconch sculpture. Ctenidium (Figure 36,

ct) extending from posterior end of mantle cavity to mantle

edge. Hypobr: mchial gland weakly developed ( (hg).

{EMARKS: With the exception of minor individual and/or

preservational differences, the external anatomy is essen-

tially identical to Juga silicula. Juga acutifilosa differs

only in that the operculum is slightly more angular and

the nucleus ¢ omprise Si a slightly smé le or proportion of the

total length, the ovipositor and groove to the « dge of the

foot are significantly deeper and more grooved, the

groove opens to the foot nearer the end oF the anterior

pedal ¢ gland, the gill extends slightly nearer the mantle

edge, and the hypobranchial g gland j is much more weakly

developed.

Alimentary System: RaApuLA: Radula comprising

~104 rows (n = 2) (Figures 38-43). Rachidian basal mar-

gin concave bordering bluntly rounded median projec-

tion: basal denticles feline or only slightly developed

(Figures 39, 40). Cutting edge bearing one lar ge central

conical cusp, and two stout, conical dente les on each

side (Figure 40). Lateral teeth (Figures 38, 39, 41) with

short lateral extensions (slightly less than half the length

of lateral cutting edge), and single, prominent triangular

cusp flanked by two inner triangular denticles and three

to four outer denticles. Wealdy developed outermost

denticle may be present or absent (Figure 41). Marginal

teeth (Figures 42, 43) with broadly rounded cutting

edges ae long, slender shafts. Inner marginal teeth wai

five and outer marginal teeth with six fletiened denticles.

Forecut: Radular sac long (~6.7 mm), extending back

through nerve ring approximately one. half distanee to

supra- “esophageal ganglion, then curving anteriorly with

tip overlying nerve ring. Long, thin tubular salivary

glands passing through circum- esophageal nerve ring, ex-

tending to posterior “esophagus.

Mine uT: Sorting area elongately triangular (Figure 37,

Sa). Accessory marginal fold (amf) forming cereal ridge

eee mar ginal fold from near esophagus, curving

around posterior margin of sorting area, with weakly be

furcate posterior sad Glandular pad (gp) ) moderately

large, rounded else with coarsely textured sur-

face. Deep pocket (c¢) extending under elendular pad

behind gastric shield. Crescentic ridge (er) bounding

narrow, shallow crescentic groove. Sty le sac ee intes-

tine forming slight protuberance at distal tip of style sac

where it separates from the latter (Figure 36, ss).

REMARKS: In eae to Juga silicula, the denticles of

the rachidian and lateral teeth are typically more conical,

with fewer denticles present on the rachidian and mar-

ginals, but more outer denticles present on the lateral

teeth. The weakly developed outermost denticle on the

lateral teeth is variably developed, and only very small.

Rachidian basal denticles are lac ‘king, or only sli ghtly de-

veloped. The radular sac is exc eptionally | ong in this spe-

cies and the salivary glands appear narrower.

Within the midgut, in spite of the differences high-

lighted above, overall configuration and proportions of

features are very similar be tween fuga silicula and J.

acutifilosa, with the exception that the sorting area is

more elongately triangular in shape in J. acutifilosa.

No significant differences in the configuration of the

hindgut, with the possible exception that the hindgut

dramatically widens upon entering the mantle cavity in

the specimens examined for Juga acutifilosa. However,

this may be individual variation.

E. E. Strong and T. J. Frest, 2007 Page 53

Figures 34-37. Anatomy of Juga (Calibasis) acutifilosa (USNM 1100659, except when noted), 34. Operculum. 35. Ovipositor and

egg groove (USNM 1100638). Right lateral view of head-foot. 36. External view of organs in visceral mass (USNM 1100658). Dotted

line indicates extent of pericardium under main kidney chamber. 37. Midgut ani itomy. Dorsal view, anterior is uppermost. Abbre-

viations: amf, accessory marginal fold: ap, anterior pedal ; gland: e, caecum: ef, caece il fold: eg, capsule gland; er, crescentic ridge:

ct, ctenidium; cu, cuticularized region of stomach roof; dd, digestive gland duct vestibule; dg, digestive gland; e, esophageal

aperture; f, foot: gp, glandular pad; gs, gastric shield: hg, hypobranchial gland; int, intestine; kd, main kidney chambe r; me, mantle

edge: mf, marginal fold: op, pee eh os, osphradium; ov, Ovary; Ovp, ovipositor; sa, sorting area; sap, sorting area pad; sn, snout:

ss, style sac: sto, stomach; t, cephalic tentacle; t1, major typhlosole; u, u-shaped fold; v, ventricle. Scale bars = 1 mm

Page 54

THE NAUTILUS, Vol. 121, No. 2

Figures 38-43. Radula of Juga (Calibasis) acutifilosa (USNM 1100659). 38. Section of anterior radular ribbon. Scale bar = 100

um. 39. Rachidian and lateral teeth. Scale bar = 50 jm. 40. Detail of rachidian teeth. Scale bar = 50 pm. 41. Detail of lateral teeth.

Note very weak yale tes of tiny outer fourth denticle. Scale bar =

50 jum, 42. Marginal teeth. Scale bar = 20 ym. 43. Detail

of cutting edge of marginal teeth. Note unequal size of cusps on inner and outer teeth. ‘Seale bar = 20 wm.

Reno-pericardial System: REMARKS: Configuration

of features within the kidney and density of excretory

tissue essentially identical to Juga silicula, with the only

exception being that the central lumen within the main

chamber is slightly shorter.

Nervous System: Pedal ganglia (pe) with two promi-

nent anterior nerves and four smaller accessory nerves.

REMARKS: Configuration of nervous system, including

number of nerves produced by major ganglia, otherwise

identical to Juga silicula.

ey ane System: FEMALE: Proximal albumen

gland (Figure 44, ag), below pallial portion of bladder,

forming small, flattened rounded pouch with glands de-

veloped along dorsal axis: shape of pouch somewhat vari-

able. Along anterior ~one third of oviduct, es sperm

gutter (sg, dashed line) present within medial lamina:

gutter opening broadly to long, broadly rounded sper-

matophore bursa (spb). Behind opening to bursa, sperm

gutter shallowing abruptly to shallow groove (dashed

line), then rapidly becoming obsolete. Near posterior

end of oviduct, small rounde ‘d aperture, just inside ven-

tral edge of medial ee leading to small narrow semi-

nal receptacle (res). Low, flatly rounded glandular pro-

tuberance from oa inner surface of lateral lamina

slightly extending into rece oS aperture. Extremely

thin, ol undular ridge extending from protuberance to ovi-

duct anterior tip along inner “edge of lateral lamina op-

posite seminal groove; ridge thickening somewhat ante-

riorly.

MALE: Glands of medial and lateral laminae forming

loosely interlocking tongue and groove arrangement

(Figures 46, 47).

REMARKS: In contrast to Juga silicula, the proximal albu-

men gland forms a small, flattened, rounded pouch. The

height and shape of the pouch is variable in J. acutifilosa,

but is distinctly smaller and slightly less glandular than

the more u- shaped tube in J. cilia ula. Ov exall. the capsule

and albumen glands are similar in proportion, but the

albumen gland is slightly shorter and not as massive as

that in J. silicula. In addition, the seminal groove is

longer and deeper anteriorly but becomes obsolete be-

fore reaching the seminal receptacle, the lateral lamina

glandular ridge is much weaker and the protuberance

into the aperture of the seminal receptacle is more

bluntly rounded than in J. silicula.

Male reproductive anatomy is very similar to that of

Juga silicula in the morphology of the folds within the

prostate, but the folds are not as highly developed and

hence, do not tightly interlock to the same degree. In

addition, the ventral fold of the medial ian: is less

textured and the dorsal and posterior flange is narrower

in ]. acutifilosa.

Juga (Oreobasis) newberryi (Lea, 1860)

Type Material: Lectotype designated by Graf (2001)

(USNM 118961; figured specimen in Lea, 1863, pl. 37,

fig. 135) (Figure 6); indicated as holotype on label.

Twalvé parale sctotypes ( (USNM LIS9GLX) in secondary

type collection of USNM; indicated as paratypes on old

label and as syntypes on newer label printed in 2001. At

the time of publication, the lectotype designation of Graf

E. E. Strong and T. J. Frest, 2007

Page 55

GL rate

erik:

SIS

Figures 44-47. Reproductive anatomy of Juga (Calibasis) acutifilosa (USNM 1100659), 44. External, left lateral view of pallial

oviduct. Anterior is to the left. Arrow indicates posterior extent of opening to gonoductal groove. “<” indicates transition between

albumen and capsule glands. 45. External, left lateral view of prostate. Anterior is to the left. Arrow indicates posterior extent of

opening to gonoductal groove. 46. Internal view of prostate medial lamina. Anterior is to the right. 47. Internal view of prostate lateral

lamina. Anterior is to the left. Abbreviations: ag, albumen gland; eg, capsule gland: gg, gonoductal groove; ovi, renal oviduct; pr,

prostate; res, seminal receptacle; sg, sperm gutter; spb, spermatophore bursa; vd, vas deferens. Scale bars = 1 mm.

was invalid as it did not follow strict guidelines concern-

ing the language of lectotype designations after 1999,

However, under Declaration 44, Amendment of Article

74.7.3 of the ree Bulletin of Zoological Nomenclature

60(4) December 2003), Graf's lectotype designation is

now valid. Collected by J. S. Newberry.

Type Locality: Indicated as “Upper des Chutes

River, Oregon Territory” (Lea, 1560) (see Figure 1).

Most likely, the lectotype came from the Deschutes R.

near Bend, Oregon, even though pleurocerids are now

absent this far up the river. However, Juga newberryi

does occur in the lower Deschutes River.

Remarks: Lea’s (1860, 1562, 1563) description of Mela-

nia newberryi and 1863 illustration are quite accurate. As

noted and illustrated by Lea, the most frequent color

pattern is three yellow bands separated by three almost

black. Occasionally, the lower band may be divided into

two or more (Burch, 1989: fig. 452) or the dark bands

may be tan in color. Alternatively, the shell may be band-

less, in which case the color varies from yellowish-tan to

dark tan. Lea (1863: 301) does not seem to have had any

of the band-less form; recent field surveys have not re-

vealed any “pure” populations of the band-less form, on

the other hand, large popul itions are not likely to lack it

(Frest, unpublishe .d data)

This species is one of a small group of Juga with the

whorls smooth throughout ontogeny which led Taylor

(1966) to make it the type of Oreobasis. In the same

Page 56

THE NAUTILUS, Vol. 121, No. 2

publication, 7 Taylor suggested that J. newberryi is a

“probable” synonym of Melania bulbosa Gould, 1847. By

1977, Taylor (1977) had accepted this synonymy without

qualification, which was followed by many authors

(Burch and Tottenham, 1980; Burch, 1982a, b, 1989:

Graf, 2001). However, it is not clear why the two species

were considered so ee Comparison of the types in-

dicates that, while being similar in whorl height, ‘the two

differ in shape of the aperture, whorl profile and rate of

whorl expansion, particularly for the body whorl; J. bul-

bosa generally has more than three not corroded whorls

while J newberryi is often more severely corroded but

the early teleoconch of the latter is quite distinctive when

present. A search of major museum collections failed to

locate many specimens aside from the types; most mu-

seum lots ascribed to J. bulbosa or J. newberryi clearly do

not belong to either.

Tryon (1865) was apparently the first to reflect on the

similarity of the two, indicating the presence of bands in

Juga newberryi as the sole separating feature, but he did

not synonymize them. However, Tryon’s illustration of J.

bulbosa (e.g. 1873, fig. 496) seems to use his own speci-

men rather than Gould's types, even though he claimed

to have had them (1873: 255). In contrast to Gould's

types, the specimen figured by Tryon is rather large,

strongly corroded with Tes than 3 whorls, and the sur-

viving whorls appear strongly convex, with a very deep

suture. It is likely a band-less form of J. newberryi. Thus,

although long considered synonyms, the pe reception that

the two are “exactly similar in outline” ( Tryon, 1865: 246)

may be due to Tryon’s apparent confusion between J.

bulbosa and bandless forms of J. newberryi. The scarcity

of museum lots may also have contributed to the confu-

sion about the morphology and occurrence of both taxa.

Consequently, we here remove Juga newberryi from

the synonymy of J. bulbosa. At present, J. newberryi

occurs only in the lower Deschutes River, Oregon, from

about Troutdale to roughly 6 miles above the mouth,

where it is replaced by J. (Juga) plicifera. Juga bulbosa is

likely also a valid species, but attempts to recollect this

taxon anywhe re in the historically identified range have

been unsuccessful (Frest, un published data): thus, this

hypothesis cannot be tested 7 the moment.

Material Examined: Oregon: Jefferson County: Des-

chutes River at RM 97.2-97.4 on E side of channel,

~0.3-0.6 km S of US 26 bridge and mouth of Shitike

Creek, at Rainbow Landing, depth 0-0.41 m (Zone 10

640460E 4957320N M21°13'36"E. 44°45'26"N], Eagle

Butte 1962 7.5’ quadrangle, elevation 440 m), Deixis

Consultants locality #2472, 13 Aug. 2000, collected by

F, EJ (USNM 1100660) (

(10 specimens dissected) (see

Figures 13-15).

External Anatomy: Operculum ovate, with angular

tip (Figure 48). Nuc leus comprising about one third of

total length (~34%).

Rather shallow ovipositor pore (Figure 49, ovp) with

shallow, grooved tract extending to edge of foot, slight

distance back from anterior pedal oland (ap).

——

Ctenidium (Figure 50, et) extending from posterior

end of mantle cavity to mantle edge. Hypobranchial

gland (hg) moderate sly well dev oe with warty tex-

ture, particularly at posterior end of mantle cavity.

Remarks: The external anatomy of Juga newberryi is

almost identical to the two preceding species, and differs

in that the operculum is more angular in shape and the

nucleus is considerably smaller and more basal than ec-

centric; as in J. acutifilosa, the gill extends slightly nearer

the mantle edge and the ovipositor groove opens nearer

the end of ie anterior pedal gland, but the pore and

distal groove are considerably “deeper in J. acutifilosa

than in ihe other two species. The hypobranchial gland of

J. newberryi is unique in having a warty texture.

Alimentary System: RADULA: Radula comprising

~106 rows (n = 2) (Figures 52-57). Rachidian with

v-shaped lower margin; basal denticles lacking (Figures

53, 54). Cutting edge bearing one large central conical

cusp, and two stout conical denticles on each side (Fig-

ure 54). Lateral teeth (Figures 52, 53, 55) with short

lateral extensions (slightly ee than half the length of

lateral cutting edge), and single, prominent triangular

cusp flanked by three to four inner and three to four

outer denticles. Outermost denticle present, weakly de-

veloped, and variable in shape, size, and position (Figure

55). Marginal teeth (Figures 56, 57) with broadly

rounded cutting ne and long, slender shafts. Inner

marginal teeth with four to five and outer marginal teeth

with six to seven flattened denticles.

ForEGUT: Radular sac moderately long (~6.25 mm), ex-

tending back through nerve ring, then curving upward

be baie base of buccal mass with tip overlying left side of

nerve ring, Epithelium of mid-esophagus ‘between dorsal

and ventral folds irregularly te xtured.

Mipcut: Sorting area short, broadly triangular (Fig 51,

sa). Accessory marginal fold (amf) forming weak ridge

par alls ling marging i fold from near esophagus, curving

around posterior me ein of sorting area, with weakly te

furcate posterior end. Glandular pad (gp) moderately

large, rounded ae riorly, with lightly textured surface.

Crescentic ridge (er) bounding shi allow, broad crescentic

groove, Style sac small (Figure 50, ss); intestine forming

prominent protuberance at distal tip of style sac where it

separates from the latter.

ReMARKS: As in Juga acutifilosa, the denticles of the

rachidian and lateral teeth are typically more conical in J.

newberryi than in J. silicula; there are only two outer

denticles on each side of the central rachidan cusp, more

outer denticles on the lateral teeth, the rachidian basal

denticles are lacking, and the lateral extensions are pro-

portionally smaller. However, in contrast to J. acutifilosa,

J. newberryi and J. silicula share a similar lower margin

on the rachidian and similar development of the weak,

outermost denticle on the lateral teeth. Juga newberryi is

unique in possessing three to four inner denticles on the

E. E. Strong and T. J. Frest, 2007 Page 57

Figures 48-51. Anatomy of Juga (Oreobasis) newberryi (USNM 1100660). 48. Operculum. 49. Ovipositor and egg groove. Right

lateral view of head-foot. 50. External view of organs in visceral mass. Dotted line indicates extent of pericardium under main kidney

chamber. 51. Midgut anatomy. Dorsal view, anterior is uppermost. Abbreviations: amf, accessory marginal fold; ap, anterior pedal

gland: au, auricle: b, bladder; e, caecum; ef, caecal fold; er, crescentic ridge; et, ctenidium; eu, cuticularized region of stomach roof:

dd, digestive gland duct vestibule: dg, digestive gland; e, esophageal aperture; f, foot: gp, glandular pad; gs, gastric shield; hg,

hypobranchial gland: int, intestine; kd, main kidney chamber; me, mantle edge; mf, marginal fold; op, operculum; os, osphradium;

ovp, ovipositor: sa, sorting area; sap, sorting area pad; sn, snout; ss, style sac; sto, stomach; t, cephalic tentacle; te; testes; tL, major

typhlosole: u, u-shaped fold. Scale bars = 1 mm

Page 58

THE NAUTILUS, Vol. 121, No. 2

TRIN —

Figures 52-57.

Radula of Juga (Oreobasis) newberryi (USNM 1100660). 52. Section of anterior radular ribbon. Scale bar = 100

pm. 53. Rachidian and lateral teeth. Scale bar = 50 pm. 54, Detail of rachidian teeth. Scale bar = 20 pm. 55. Detail of lateral teeth;

note weakly formed outer fourth denticle. Scale bar = 50 jum. 56. Marginal teeth. Scale bar = 50 wm, 57. Detail of cutting edge of

marginal teeth. Note unequal size of cusps on inner and outer teeth. Scale bar = 20 jm.

lateral teeth. The radular sac is intermediate in length

relative to the buccal mass compared to J. silicula with a

very short sac, and J. acutifilosa with a very long sac.

Juga newberryi differs greatly from the two preceding

species in the relative proportions of features in the mid-

gut, which is significantly longitudinally compressed.

Consequently, the sorting area is broadly triangular

rather than elongate and thé glandular pad is dispropor-

tionately smaller. Possibly reflecting the smaller size of

the style sac, the opening of the style sac to the stomach

is also consistently smaller.

There are no significant differences in the configura-

tion of the hindgut between Juga newberryi and J. sili-

cula.

Reno-pericardial System: REMARKS: In contrast to

both preceding species, the bladder of Juga newberryi is

compressed and less voluminous, bringing the nephro-

pore closer to the afferent renal vecel and decreasing

the length of the wall between the main chamber and fhe

bladder. Additionally, the sheets of excretory tissue in the

bladder are more numerous, and more highly branched,

almost entirely occluding the lumen.

Nervous System: Pedal ganglia with two prominent

anterior nerves andl five to six smaller accessory nerves.

In addition to comnections to right and left pleural and

visceral ganglia, sub-esophageal ganglion producing

three small nerves.

{EMARKS: Apart from the differences highlighted above,

configuration of the nervous system is basically identical

to Juga silicula.

Reproductive System: FEMALE: Capsule gland (Fig-

ure 58, eg) comprising approximately anterior two thirds

of pallial oviduct. Mong anterior ~one third of oviduct,

shallow sperm gutter (sg, dashed line) present within

medial lamina; gutter opening narrowly to long, thin

spermatophore bursa (spb). Sperm gutter continuing

Figure 58.

Reproductive anatomy of Juga (Oreobasis) new-

berryi (USNM_ 1100660). Exte smal, left lateral view of pallial

oviduct. Anterior is to the left. Arrow indicates posterior extent

of opening to gonoductal groove, “<4” indicates transition be-

tween albumen and capsule glands. Abbreviations: ag, albu-

men gland; eg, capsule gland; gg, gonoductal groove; evi, renal

oviduct; res, seminal receptacle; sg, sperm gutter; spb, sper-

matophore bursa. Scale bar = 1 mm.

E. E. Strong and T. J. Frest, 2007

Page 59

posteriorly as shallow groove to small, narrow seminal

receptacle (res). Moderate glandular protuberance from

opposing inner surface of lateral lamina extending into

receptacle aperture. Glandular ridge extending from

protuberance to oviduct anterior tip along inner edge of

lateral lamina opposite seminal groove; ridge thinning

anteriorly.

Remarks: The proximal albumen gland forms a promi-

nent u-shaped tube that is larger than that in fuga sili-

cula. The spermatophore bursa is smaller than both pre-

ceding species, but as in J. acutifilosa, the glandular ridge

directly opposes the seminal groove and the glandular

protuberance is intermediate in size between that of J:

silicula and ]. acutifilosa.

Apart from minor individual variation in shape and

development of the prostate glands, there are no detect-

able discrete differences in male reproductive anatomy

compared to Juga silicula. Some minor differences in-

clude the fact that the ventral fold of the medial lamina

appears less textured as in J. acutifilosa, and the dorsal

posterior flange appears consistently larger.

DISCUSSION

EVALUATION OF MORPHOLOGICAL CHARACTERS: Most

anatomical differences among the three species investi-

gated herein comprise qualitative variations in size and/

or shape (e.g. operculum, hypobranchial gland, oviposi-

tor, radula, and bladder). Although often emphasized in

species-level systematic studies of gastropods, male and

female reproductive anatomy also displays only minor

differences in shape and size of the various glands,

pouches, and gutters. Additional sampling within and be-

tween Juga species is necessary to determine if any of

these comprise discrete rather than continuous character

variation. Surprisingly, the midgut offers a significant

source of variation, with medincatons to size of the cae-

cum, style sac, and proximal intestine, as well as overall

proportions of the gastric chamber and sorting area. This

is very different from eastern North American pleuro-

cerids with species from disparate genera displaying al-

most identical midgut morphologies (Strong, 2005;

Strong, unpublished data). For a summary of ‘these and

other differences, see Table 1

The only published anatomical account of any North

American Juga is that of Prozorova and Raschepkina

(2004) on the female reproductive anatomy of five un-

determined Juga species from the Willamette River sys-

tem in Oregon. That study indicated the size and shape

of the seminal receptacle and spermatophore bursa can

vary, which was confirmed in the present study. How-

ever, Prozorova and Raschepkina reported seve ral addi-

tional findings that could not be confirmed here, includ-

ing a sperm gutter that becomes suddenly shallow (an-

terior one fifth to one sixth), variable le neth of the

opening between the gonoductal groove and momen cav-

ity, and asymmetrical arrangement of albumen and cap-

sule glands. Indeed, the dorso-ventral axis (as defined by

a plane extending between the gonoductal groove and

the opening to the mantle cavity), is not correctly iden-

tified by these authors. Thus, they mistakenly conclude

that the lateral lamina is glandular and the medial lamina

is non-glandular, comprising the spermatophore bursa

and sansivial receptacle. Consequently, the glands of the

oviduct are misinterpreted as a solid block penetrated by

channels within the lateral lamina, rather than as a con-

voluted tube. This misunderstanding does not allow

them to correctly identify the basic symmetry between

the medial and lateral laminae, and the proportional de-

velopment of the capsule and albumen glands along the

antero-posterior axis. Instead, the albumen gland is often

depicted as extending far anteriorly, dorsally overlying

the c capsule g gland. Not only is this i incorrect, it is difficult

to imagine how such an arrangement would function.

ANATOMIC AND SYSTEMATIC AFFINITIES OF [UGA: — No

conchological feature unambiguously distinguish res Juga

from eastern North American pleuroce rid genera; in-

deed, Juga has often been synonymized w ith lamin (as

Goniobasis )—a genus wide spread east of the continental

divide (e.g. Tryon, 1865, 1873; Pilsbry, 1899; Walker,

1918; Henderson: 1935a, b; Goodrich, 1942). However,

increasing evidence not only supports the independence

of Juga, but suggests ties to Asian pleurocerids. In an

analysis of a large segment of the mitochondrial 16S

rRNA gene including representatives of five eastern

North American pleurocerid genera, Juga falls to the

base of the tree rooted on Melanopsis praemorsa (L.)

(Melanopsidae) and Melanoides tuberculata (Miiller,

1774) (Thiaridae) (Holznagel and Lydeard, 2000). In an

analysis of cerithioidean relationships using nearly full

length 16S rRNA sequences (Lydeard et al., 2002), East-

erm North American pleurocerids ( (Elimia, Pleurocera)

are supported as more closely related to Melanopsis than

to a clade uniting Juga with Eastern Asian pleurocerids

(Semisulcospira Boettger, 1886, Hua Chen, 1943).

Available evidence from reproductive anatomy may

also support a link between fuga and Asian pleurocerids.

A recent contribution by Strong (2005) re-described the

anatomy of Pleurocera acuta Rafinesque, 1831 and

Elimia livescens (Menke, 1830) with a re-evaluation of

pallial oviduct homologies. Despite several erroneous ac-

counts (Woodard, 1934; Jones and Branson, 1964; Dazo,

1965), Strong’s (2005) analysis confirmed Eastern North

American pleurocerids described thus far lack a seminal

receptacle—a feature present in all described western

North American ( (Juga) ) and Asian (Hua, Se misulcospira)

species (Itagaki, 1960; Nakano and Nishiwaki, 1989; Pro-

zorova, 1990; Rashchepkina, 2000; Prozorova and Ra-

schepkina, 2001, 2004, 2005). However, as stated above,

the only information available on the anatomy of Juga

concerns female reproductive anatomy. Given the de-

tailed description of three Juga species herein, it is now

possible to better place this unique assemblage within

the emerging anatomic and phylogenetic framework for

limnic cerithioideans.

Page 60

THE NAUTILUS, Vol. 121, No. 2

Table 1. Summary of anatomical differences between three species of Juga.

Juga (Juga) silicula

Juga (Calibasis) acutifilosa

Juga (Oreobasis) newberryi

External Anatomy:

Operculum nucleus size

(as percent of total

length)

Ovipositor pore

Hypobranchial gland

Alimentary System:

Dentition:

Rachidian tooth

Lateral teeth

Marginal teeth

Rachidian basal denticle

Rachidian basal margin

Weakly formed outermost

denticle of lateral teeth

Radular sac

Salivary glands

Sorting area

Crescentic groove

Caecum

Style sac

Proximal intestine at base

of style sac

Reno-Pericardial System:

Bladder excretory tubules

Reproductive System:

Proximal albumen gland

Spermatophore bursa

Sperm gutter

Lateral lamina glandular

ridge

Prostate glands

45%

Shallow

Transversely ridged

3/1/38

2/1/2-3

6/7

Present

Bluntly v-shaped

Present

Short

Thick, tubular

Elongately rectangular

Shallow, broad

Shallow

Large

Large protuberance

Loosely and regularly branched

Moderately large, u-shaped

tube

Large, rounded

Anterior one fourth of oviduct:

extends to seminal

receptacle as shallow groove

within medial lamina

Inside seminal groove; large

protuberance extending into

receptacle aperture

Tightly interlocking

41%

Deep, highly ridged

Thin

2/1/2,

2/1/34

5/6

Lacking, slightly developed

Concave bordering rounded

median projection

Absent

Long

Thin, tubular

Elongately triangular

Shallow, narrow

Deep

Large

Small protuberance

Loosely and regularly

branched

Small, rounded pouch

Moderately large, rounded

Anterior one third of

oviduct; becomes

obsolete before reaching

receptacle

Opposite seminal groove;

low, flat protuberance

extending into receptacle

aperture

Loosely interlocking

34%

Shallow

Warty

2/1/2

34/1/34

4-5/6-7

Lacking

Bluntly v-shaped

Present

Moderately long

Thick, tubular

Broadly triangular

Shallow, broad

Shallow

Small

Large protuberance

Densely and highly branched

Large, u-shaped tube

Narrow, tubular

Anterior one third of oviduct:

extends to receptacle as

shallow groove within medial

lamina

Opposite seminal groove;

moderate protuber rance

extending into receptacle

aperture

Tightly interlocking

Juga, like other pleurocerids, possesses an ovipositor

involved in the deposition of the egg capsules (except the

viviparous Semisulcospira; Itagaki, 1960) (e.g. Jewell,

1931; Woodard, 1934; Magruder, 1935b; Morrison,

1954; Jones and Branson, 1964; Prozorova, 1990; Rash-

chepkina, 2000; Prozorova and Raschepkina, 2001, 2004

2005; Strong, 2005). The ovipositor pore in Juga forme a

broad, shallow triangular shelf with a highly grooved epi-

thelium. This is very different from the deep pore that

expands medially into the foot of Plewrocera and Elimia,

with parallel folds that direct the movement of ova

through the pore (Strong, 2005). Indeed, this feature was

cited by ” ae (1966) as justifying the independence of

Juga.U nlike Juga, the distal ovipositor groove extends to

the foot sole in some but not all of eastern North Ameri-

can pleurocerids (e.g. Van Cleave, 1932; Morrison, 1954;

Strong, 2005): ovipositor morphology of Asian pleurocer-

ids is unknown. In melanopsids, the pore is deep, glan-

dular and complex and the distal 1 groove does not ye

sect the foot sole (Bilgin, 197: 3. Glaubrecht, 1996) (Ta-

ble 2)

Similar to other pleurocerids, the gut of Juga species is

characterized by the presence of tubular salivary glands

that pass through the nerve ring, the absence of a mid-

esophageal eland, and a style sac in restricted communi-

cation with the proximal intestine (Mz agruder, 1935a, b:

Itagaki, 1960; Dazo, 1965; Strong, 2005). Rachidian basal

eae *s are present in at least some Asian pleurocerids

(Ko et al., 2001) and are apparently lacking in many

eastern North American ple surocerids (e. g. Minton et al.,

2004; Sides, 2005). However, they are easily overlooked

in whole mounts and their absence from existing descrip-

tions may be an error; for example, basal denticles are

present in Elimia livescens (Glaubrecht, unpublished

E. E. Strong and T. J. Frest, 2007

Page 61

Table 2. Summary of anatomical differences between Juga and other limnic gastropods classified in the Pleuroceridae and Mel-

anopsidae. Details from Sunderbrink, 1929; Sods, 1936; StarmiihIner and Edlauer, 1957; Itagaki, 1960; StarmiihIner, 1970; Bilgin,

1973; Houbrick, 1958; Nakano and Nishiwaki, 1989; Glaubrecht, 1996; Strong and Glaubrecht, unpubl. data. NA = not applicable.

Elimia livescens

Juga Pleurocera acuta

External Anatomy:

Ovipositor pore

Alimentary System:

Salivary glands

Salivary gland position

Esophageal gland

Digestive gland ducts

Caecum

Reno-Pericardial System:

Bladder

Evagination of bladder

wall

Shallow, simple,

weakly glandular

Tubular

Dace ‘

Pass through

nerve ring

Absent

]

Shallow/Deep

Small, pallial

Present

Deep, simple,

Semisulcospira Melanopsidae

NA Complex, highly

weakly glandular glandular

Tubular Tubular/branched

Tubular

Pass through

nerve ring

Pass through

nerve ring

Pass

through/anterior

to nerve ring

Absent Absent? Present

2 2 |

Shallow Shallow Deep and spiral

Small, pallial ? Small, pallial

Absent ? ?

Nervous System:

Dialyneury, Zygoneury Zygoneury

Zygoneury

Dialyneury? Zygoneury

Accessory ganglion Present Present Absent? Present?

between left pleural

and sub-esophageal

ganglia

Reproductiv e System:

Ovipositor distal groove Present Present/Absent NA : Absent

intersects foot sole

Seminal vesicle Straight Straight Straight Folded

Proximal albumen gland U-shaped Straight 2 ?

Gonoductal groove Convoluted Simple Simple? Simple?

Seminal receptacle Present Absent Present Present

Reproductive strategy Oviparous Oviparous Viviparous Oviparous

data), but have not been described in the literature (e.g.

Baker, 1928; Dazo, 1965). The phylogenetic significance

of these features is unclear as they occur sporadically

among many cerithioidean groups, including batillariids,

planaxids, melanopsids, thiarids (see e.g. Thiele: 1928;

Houbrick, 1987: Glaubrecht, 1996). Melanopsids differ

in eee an esophageal gland, and the salivary glands

may be tubular or branched and variably pass through or

by-pass the nerve ring; the radula is variable and may

present a rachidian ee is similar to that of pleuroce rids,

or may be quadrangular, robust with conical denticles

and with a marked glabella (Sunderbrink, 1929; Thiele,

1928: Bilgin, 1973; Glaubrecht, 1996).

The midgut of Juga species diverges from eastern

North American pleurocerids and other putatively

closely related limnic lineages most noticeably in size and

shape of the glandular pad ‘and configuration of the cres-

centic ridge. In Juga, the glandular pad is rather narrow

with a crescentic ridge hata is often separated from it by

a wide, shallow groove; the proximal end of the crescen-

tic ridge borders a vestibule that receives several ducts of

the digestive gland. Typically, melanopsids, paludomids,

and thiarids have a broadly rounded glandular pad and a

deep, narrow crescentic groove such that the crescentic

ridge closely adheres to the outer edges of the pad. Al-

though the midgut has demonstrated great utility in re-

constructing relationships among seatioiices line ages

(Strong, unpublishe d data), midgut characters of Jiga do

not provide unambiguous evidence of affinity to any one

freshwater family; the small, narrow glandular pad is

rather similar to that of Se misulcospira, "bist the configu-

ration of the crescentic ridge, particularly in J. silic ae

and J. newberryi, is unique among cerithioideans known

thus far. The presence of a single digestive gland duct

vestibule is shared between Juga and melanopsids, but is

also found in a number of cerithioideans; other pleuro-

cerids and paludomids have two digestive gland ducts

(Strong and Glaubrecht, 2002, 2003, 2007, unpublis! ied

data; Strong, 2005).

As in paludomids ( e.g. Strong and Glaubrecht, 2002,

2003) and melanopsids ( (Bilgin, 1973), the kidney of pleu-

rocerids penetrates the palli: ul cavity (Mz agruder, L935b;

oe 1960: Strong, 2005), but the pallial portion of the

bladder is smaller Than that in the former taxa. The

branching pattern of excretory tubules within the blad-

der is essentially identical between Juga and eastern

North American pleurocerids, but Juga is unique in the

outpocketing of the wall separating the main chamber

and the bladder (Strong, 9005) Kidney anatomy is cur-

rently unknown for Asian pleurocerids ‘and me lanopsids

Page 62

THE NAUTILUS, Vol. 121, No. 2

Although published accounts of pleurocerid nervous

systems disagree on the number of nerves produced by

various ganglia (Magruder, 1935b; Itagaki, 1960; Dazo,

1965; Strong, 2005; present study), this is often variable

within species. However, the present account agrees

with that of Strong (2005) that the cerebral ganglia pro-

duce seven nerves, and that there are two prominent

anterior pedal nerves with a variable number of small

accessory nerves (typically four to seven). The thickened

connective between the left pleural and sub-esophageal

ganglia, representing a small accessory ganglion, was

found to give off only a single large nerve in the present

study, bar was found by Strong (2005) to produce one to

three nerves in Elimia and Pleurocera. Str ong (2005) also

found the sub-esophageal ganglion to give off one to

three nerves, whereas three to four nerves were found in

the present study.

The most intriguing aspect of the nervous system is the

presence of the accessory ganglion. It is known only from

eastern North American Sead (Magruder, 1935b:

Strong, 2005) and now Juga (present study) ), and has not

been depicted in most accounts of melanopsid (Bouvier,

1887: Sods, 1936; Starmiithlner and Edlauer, 1957;

StarmiihIner, 1970; Bilgin, 1973; Glaubrecht, 1996) and

Asian pleurocerid Gnesi, 1960) nervous system

anatomy. However, the illustration of the nerve ring of

Melanopsis frustulum Morelet, 1856-57 (St: ee

1970) and that of M. doriae Issel, 1866 (StarmiihIner andl

Edlauer, 1957) clearly show a thickened connective be-

tween the left pleural and sub-esophageal ganglia, indi-

cating that it is most likely present in melanopsids as

well. Paludomids and thiarids have a much more con-

centrated nerve ring with the left pleural and sub-

esophageal ganglia fused or in close contact (e.g. Se-

shaiya, 1934: Glaubrecht, 1996; Strong and G laubrech®

2002. 2003).

Pleurocerids and melanopsids share the same basic

layout of the pallial oviduct to the exclusion of paludo-

mids and thiarids, including the presence of a long open-

ing to the mantle cavity, anda deep sperm gutter open-

ing anteriorly to a spermatophore bursa and posteriorly

to a seminal receptacle; as mentioned above, eastern

North American pleurocerids are unique in lacking the

seminal receptacle (Bilgin, 1973; Nakano and Nishiwaki,

1989: Prozorova, 1990: Glaubrecht, 1996: Rashchepkina,

2000: Prozorova and Raschepkina, 2001, 2004, 2005:

Strong, 2005). Whereas in Juga and Semisulcospira the

sperm gutter becomes obsolete or continues posteriorly

as a shallow groove within the medial lamina (Prozorova

and Aaschepkina, 2004, 2005; herein), in melanopsids

the sperm gutter is continuous along the ventral edge of

the medial lamina and contains the opening to the re-

ceptacle at the posterior end of the oviduct (Bilgin, 1973;

Glaubrecht, 1996). In Elimia and Plewrocera, a deep

sperm gutter 1s present above the opening to the mantle

cavity along its entire length, and closes posteriorly to

form a short, blind spermatophore bursa.

One aspect of cerithioidean reproductive anatomy that

is routinely overlooked is the configuration of the capsule

and albumen glands. Strong and Claus echt (2002, 2003)

have found that the shape of the albumen gland in palu-

domids is quite distinctive, and likely a synapomorphy of

the family. In fuga, the glands and intervening gonoduc-

tal groove of the pallial oviduct are highly conimlued

with a proximal albumen gland that is pouch- -like or u-

shaped (present study), elas eastern North American

pleurocerids possess ¢ clands that form two narrow bands

with smooth opposing surfaces and an essentially linear

proximal albumen gland (Strong, 2005). However, such

potentially informative characters are undescribed for

other pleurocerids and melanopsids.

Male reproductive anatomy of Juga is apparently quite

distinct as no other male pleurocerid (indeed, no other

cerithioidean) has been described with a tightly inter-

locking arrangement of glands (e.g. Woodar d, 1934; Ita-

gaki, 1960; Prozorova, 1990): however, a large fold within

the lateral lamina may be present (Nakano and Nishi-

waki, 1989; Strong, 2005). Among eastern North Ameri-

can pleurocerids, hoch the highly Folded proximal portion

(Woodard, 1934) and distal region of the prostate Strong

(2005) have been inferred as the site of spermatophore

formation. In Juga, there is no differentiated anterior or

posterior region, but intuitively the tightly interlocking

folds of the Tateral and medial laminae must function in

molding the spermatophore.

CONCLUSIONS

While sharing many similarities, numerous features

clearly set Juga apart from eastern North American pleu-

rocerids: ovipositor pore, lateral outer cusps, midgut

glandular pad and crescentic ridge, digestive gland duct

vesubule; evagination of kidney "wall: convoluted gono-

ductal groove, proximal albumen gland, seminal recep-

tacle, interlocking prostate glands. However, as is often

the case for ancient lineages , it is difficult to identify

uniquely shared features that more or less unambigu-

ously demonstrate affinity to any one limnic lineage. Juga

is particularly difficult as many of these features not only

set the genus apart from other pleurocerids, but are un-

dccumente d (ovipositor pore, kidney, pallial oviduct

glands) or apparently autapomorphic among cerithioide-

ans (crescentic ridge, evagination of the kidney wall,

prostate). Nevertheless, this analysis has revealed some

characters that are shared with Asian pleurocerids (mid-

gut glandular pad, seminal receptacle), but some that are

also shared with melanopsids (digestive gland duct).

However, the presence of the seminal receptacle is un-

doubtedly plesiomorphic and uninformative in delineat-

ing rele tionships. Others are so heterogeneously distrib-

uted among limnic lineages, it is difficult to determine if

there may be an unde rlying phylogenetic signal (oviposi-

tor ventral eroove, Ti achidian basal denticles), Unlike mo-

lecular dats. given the available morphological evidence,

there is no overwhelming signal linking Juga to Asian

pleurocerids, nor for that matter linking eastern North

E. E. Strong and T. J. Frest, 2007

Page 63

American pleurocerids to melanopsids (Table 2). While

part of this pattern may be due to the high rate of auta-

pomorphies, it may simply be an artifact of missing data.

Clearly, one of f the more significant impediments in as-

sessing pleurocerid affinities is that comprehensive ana-

tomical accounts of pleurocerids and melanopsids are

rare, leaving many potentially informative characters un-

known. Thus, it is clear that further anatomical studies

within the context of ongoing morphological and molecu-

lar cladistic analyses are necessary to unambiguously re-

solve the question of pleurocerid monophyly and their

affinities to melanopsids.

These results also have implications for the systematics

of Juga, particularly with regards to the y validity of cur-

rent subgeneric subdivisions. Although all three taxa pos-

sess unique features that clearly distinguish them from

one another, ]. silicula and J. newberryi are strikingly

anatomically similar and share many features to the ex-

clusion of J. acutifilosa: 1) shallow, weakly ridged ovi-

positor pore, 2) similar rachidian basal margin, 3) similar

development of weak, outermost denticle of the lateral

teeth, 4) thick, tubular salivary glands, 5) shallow, broad

crescentic groove, 6) shallow caecum, 7) large proximal

intestine protuber rance on base of style sac, 8) u-shaped

proximal albumen gland, and, 9) tightly interlocking

prostate glands (T Table 1). In conclusion, the enatomieal

data do not support separation of Juga sensu stricto and

Oreobasis and Oreobasis is thus here synonymized with

Juga sensu stricto; Calibasis is sufficiently distinct to

merit retention as a valid taxon. The question of the rank

of these taxa is, of course, highly subjective. But given

what is known about Anotouical differentiation among

other pleurocerid genera, it could reasonably be ar cued

that Juga sensu stricto and Calibasis be recognized at the

rank of genus.

ACKNOWLEDGMENTS

We are indebted to Marilyn Schotte (USNM) for inking

the anatomical drawings and for scanning electron mi-

oa We also thank Philippe Bouchet (Muséum na-

tional d'Histoire naturelle, Paris) for critical reading of

the manuscript and consultation on matters of zoological

nomenclature. Arthur Bogan (North Carolina State Mu-

seum of Natural Sciences) and an anonymous reviewer

provided valuable comments. We are also grateful to Ed

Johannes (Deixis Consultants) for Figure 1, for collecting

specimens with TF, and for compiling locality descrip-

tions. Support to Deixis Consultants from the Cantara

Trustee Council is gratefully acknowledged (Grants

C0010010, CO110013).

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THE NAUTILUS 121(2):66-75, 2007

Page 66

Discriminating shells of Gastrocopta pentodon (Say, 182

2) and

G. tappaniana (C. B. Adams, 1842) (Gastropoda: Pulmonata)

with an example from the Delmarva Peninsula, eastern USA

Timothy A. Pearce!

Marvin C. Fields

Carnegie Museum of Natural History

4400 Forbes Avenue

Kayoko Kurita

National Institution for Academic Degrees

and University Evaluation

1-29-1 Gakuen-nishimachi, Kodaira-shi

Pittsburgh, PA 15213 USA Tokyo, 187-8587 JAPAN

ABSTRACT

The North American pupilloid land snails Gastrocopta pent-

odon and G, tappaniana have similar shells that can be difficult

to separate, which raises the question of whether they repre-

sent two species or environmentally influenced variants of one

species. In 1906, Vanatta and Pilsbry presented 53 drawings of

shells of the two species to facilitate separation, but differences

therein were subtle. Discriminant function analysis of measure-

ments from their illustrations gave a discriminant function clas-

sifying 96% correctly, with aiaimnal overlap between groups

shown by factor analysis. The function revealed both forms on

the Delmarva Peninsula, again with minimal factor analysis

overlap. Bimodality of morphology does not reflect sexual di-

morphism in these hermaphrodites.

The two forms maintained their distinct morphologies where

they co-existed, supporting the concept of separate species.

Reports of the forms in habitats of different wetness could

indicate separate species with different moisture preferences or

one species with moisture-influenced morphology. We found

the two forms to show distinct morphologies in medium wet

areas, further supporting the concept of separate species.

Regarding habitat wetness, we confirmed Delmarva Gastro-

copta tappaniana in wetter areas, whereas G, pentodon oc-

curred in a wide range of moistures, but tending to be found in

drier areas. Our surrogate measure of habitat wetness relying

on plant moisture requirements should be useful in future stud-

ies. Geographically, G. tappaniana tended to occur along the

SE Delmarva Atlantic coast while G. pentodon ranged more

wide ly

In an application of the discriminant function, measurements

from an image of the lectotype of Gastrocopta carnegici clas-

sified that species with G. pentodon.

INTRODUCTION

The ability to distinguish one species from another is

central in biology. However, environmental variation in

Corre sponding author pearcet@carnegiemnh org

form can sometimes be mistaken for species-level differ-

ences in taxa including mollusks (Minton and Gunder-

son, 2001).

Historically, making a distinction between the North

American pupilloid land snails Gastrocopta pentodon

(Say, 1822) and Gastrocopta tappaniana (C. B. Adams,

1842) has been difficult (Vanatta and Pilsbry, 1906).

These latter authors discussed and evaluated G. pen-

todon, G. tappaniana, and other similar, previously de-

scribed forms. They also illustrated various modifications

of form, Their paper presented drawings of 53 Gastro-

copta specimens from eastern North America. Although

Vanatta and Pilsbry identified some of their illustrated

specimens as G. pentodon and others as G. tappaniana,

they did not reveal their criteria for specific allocation of

these individual specimens.

Vanatta and Pilsbry (1906) and Pilsbry (1948) de-

scribed distinctions between Gastrocopta pentodon and

G. tappaniana including differences in shell size and

shape, and the number and arrangement of apertural

teeth. Despite Vanatta and Pilsbry’s clarification of the

species differences, confusion persists and some workers

have consolidated the two forms into one single species

while others have kept them as separate species.

Bequaert and Miller (1973: SS-90) lumped the species,

stating that the holotype (actually the lectotype, selected

by Clench, 1965) of G. tappaniana is a typical G. pen-

todon. On the other hand, Hubricht (1976: 107) also

examined the type of G. tappaniana and concluded it was

G. tappaniana and not G. pentodon. Lauriol et al. (2003)

and Nekola (2004) recognized the two forms as separate

species.

Our visual e xamination of Vanatta and Pilsbry’s (1906)

drawings of the 53 specimens in light of their statements

comparing the os species left us uncertain that those

shell characters could reliably separate the two species.

Some of the G. pentodon seemed to have characteristics

of G. de tarlge G vice versa (see in particular their

figs. 17, 32, 45, . Admittedly, Vanatta and Pilsbry at-

tempted ie aha variability in the forms, such that they

might have chosen extreme examples.

T. A. Pearce, M. C. Fields, kK. Kurita, 2007

Page 67

Multivariate statistical procedures are obviously more

accessible now than they were in 1906. Factor analysis

(FA) identifies the axes of mé yor variation in a dataset

and helps show whether specimens cluster into more

than one group in a multidimensional space. Discrimi-

nant function analysis (DFA) can identify which variables

are most useful for separating two pre-defined groups

and can identify coefficients to use with the variables in

a discriminant function (DF) for classifying unknown

specimens. Nekola and Barthel (2002) use da r simlae ap-

proach.

Some researchers have suggested that Gastrocopta

pentodon prefers drier areas and G. tappaniana prefers

moister areas (Sterki, 1906: 134; Pilsbry, 1948: SSS—S90;

Hubricht, 1985: 9: Nekola, 2004). One hypothesis holds

that the forms are separate species as evidenced by their

different moisture preferences. An alternate hypothesis

is that, since larger snails are sometimes associated with

moister conditions (Goodfriend, 1986, and references

therein), the two forms might simply be two ends of an

environmentally influenced continuum of morphology. If

environmental conditions are continuous, then randomly

sampled specimens should show continuous morphological

variation if they represent one environmentally influenced

species or a bimodal distribution if they are two species

with different moisture preferences. F urthermore, examin-

ing w hether the two species maintain their separate mor-

phologies where they co-occur would provide strong evi-

dence whether they are two separate species.

Numerous specimens tentatively identified as Gastro-

copta pentodon were available from a survey of terrestrial

mollusks on the Delmarva Peninsula (Pearce and Italia,

2002). Examining these specimens using morphometric

methods should address whether both forms occur on

Delmarva and, if so, whether the forms are distinct and

whether they differ in their moisture preferences.

In this paper we explore the moisture associations of

the two forms on the Delmarva peninsula to examine

whether specimens identified as G. pentodon occurred in

drier habitats and whether G. tappaniana occurred in

moister habitats. Furthermore, we look for interpretable

differences in geographical distribution of the two forms

on the peninsula. Finally, by examining specimens at

localities where the two forms co-occur, we address

whether the two forms are valid species or ecomorphs of

a single species.

We address three principal questions in this paper:

(1) Are the two forms identified by Vanatta and Pilsbry

(1906) distinguishable via morphometric analysis and, if

so, which fenpines discriminate between them? To an-

swer this question, we examined selected variables from

their 53 illustrations in FA and DFA.

Do the Gastrocopta pentodon and G. tappaniana

from the Delmarva Peninsula fall into two morphometric

groups and, if so, how do the features of those two groups

compare with those of the forms as identified by Vanatta

and Pilsbry (1906)? To answer this question, we applied

the DF from question (1) to 577 Delmarva specimens

and examined results along FA axes.

(3) Does the wetness of the habitats of the Gastrocopta

pentodon and G. tappaniana forms differ as previously

suggested? How are they distributed on the peninsula?

Does co-occurrence evidence addresses whether the two

forms represent separate species or environmentally in-

fluenced morphs of the same species? To answer this

question, we compared occurrences using wetness esti-

mated from plant data and compared morphology of the

forms in sympatry and allopatry.

MATERIALS AND METHODS

Sources of Specimens: We examined illustrations of

the 53 Gastrocopta shells presented by Vanatta and Pils-

bry (1906), which had been drawn to the same scale

oe using a camera lucida. Their specimens were

from geographically widespread areas in eastern North

America. They identified the specimens as 41 G. pen-

todon and 12 G. tappaniana.

We examined 577 unbroken adult Gastrocopta shells

from 130 leaf-litter samples from 123 grid squares on the

Delmarva Peninsula. The number of shells measured per

sample ranged from 1 to 69. The Delmarva Peninsula

shells came from a study of land snails on the peninsula

(Pearce and Italia, 2002) in which leaf litter samples (1-4

liters) were collected from 794.5 « 5 km grid squares,

passed through sieves, and snails retained by screens 0.5

mum or larger mesh were picked and identified. We ex-

cluded juveniles (adults are easily recognized by the

presence of a reflected lip and we I-deve loped apertural

teeth) to avoid introducing variability from shells of dif

ferent ages and excluded fe other clearly different spe-

cies of Gastrocopta found on Delmarva. Although speci-

mens resembling G. pentodon and G. tappaniana were

found in 201 samples from 11 different grid squares on

Delmarva (Figure 1) excluding broken, juvenile, or wet-

preserved specimens (bodies of alcohol-preserved speci-

mens obscured apertural features) reduced the number

of measurable specimens.

Differences between the datasets: the Vanatta and

Pilsbry geographic coverage is about two orders of mag-

nitude larger (throughout « -astern North America for

Vanatta and Pilsbry, just the Delmarva Peninsula for

Delmarva), whereas the Delmarva sample size is an or-

der of magnitude larger (n = 53 for Vanatta and Pilsbry,

n = 577 for Delmarva).

Voucher specimens are deposited at the Delaware

Museum of Natural History.

Selection and Measurement of Variables: \Ve took

measurements from camera lucida drawings of shells,

using 53 drawings by Vanatta and Pilsbry (1906) and 577

of our own drawings of specimens from the Delmarva

Peninsula.

Page 68

THE NAUTILUS, Vol. 121, No. 2

Figure 1. Delmarva Peninsula showing locations where

specimens of Gastrocopta pentodon and G tappaniana were

found

We measured or counted 8 variables from the draw-

ings and derived 13 additional (ratio) variables as com-

binations of the measured variables (Table 1, Figure 2).

We excluded teeth on the parietal wall from the count of

teeth because whereas only one parietal tooth is present

in Gastrocopta tappaniana, either one or two parietal

teeth can be present in G. pentodon (Pilsbry, 1948: 889).

\perture height was measured from the middle of the

parietal callus.

We included derived variables because ratios suc-

cinctly describe shape, which is not described by original

measurements. Because ratios are more intuitive, includ-

ing ratios in this analysis should give a more useful result

for separating these species. Although ratios can theo-

retically produce non-normal distributions, the ranges of

our variables are limited so they can be used in FA and

DFA. We did not check for normality or transform vari-

ables because FA and DFA are robust against non-

normality (Mardia, 1971; Hagiuda and Shigemasu, 1996).

In order to reduce she fcmabe sr of var bles to about

1/5 the number of specimens (Hair et al., 1998), and to

determine the set of variables that have the greatest

chance of separating the species as defined, we used

non-parametric Mann- Whitney U-tests to compare mea-

surements of each variable on the two Gastroc ‘opta forms

as defined (Table 1) and used the variables that showed

significant differences between the two groups in further

analyses.

Statistical Techniques: DFA and FA were per-

formed using SAS version 8.02 (SAS Institute Inc., Cary,

NC). We used DFA to determine the variables most

useful in discriminating between the two forms using

stepwise, backward, and forward methods for variable

selection. The three methods gave similar sets of vari-

ables to use in separating the species, giving us confi-

dence that two recognizable groups exist in the data set.

Then SAS applied the DF to the analyzed specimens to

determine reclassifications and percent correct classifi-

cation. We applied those DF coefficients to measure-

ments of the 577 specimens from the Delmarva Penin-

sula to determine which forms occur there.

We used FA to examine graphically whether two mor-

phological groups of individuals are evident among the

specimens, using the variables determined by the DFA.

We used FA with principal compone nts (esse tially the

same as principal component analysis with rote ition). We

accepted the first three axes, applie sd varimax rotation,

and plotted specimens on the factor axes

Distinctness of the Forms: To further study distinct-

ness of the two forms, in addition to examining FA plots,

we examined whether individual samples from Delmarva

contained mostly one form of Gastrocopta or if forms

tended to be randomly distributed among samples. To

examine whether this non-randomness represents two

species, or one environmentally influenced species, we

examined their morphologies in symp yatry. If the forms

are two morphologically distinct species, then their mor-

phologies should remain distinct and mixed samples,

containing both forms, should exhibit bimodal morphol-

ogy. If the forms are a single species with environmen-

tally de pendent morphology, morphology should not

show bimodality under intermedi: i: environmental con-

ditions; in intermediate conditions, morphology should

be intermediate.

To verify the names being applied to the two forms, we

measured and compared the lectotype of Gastrocopta

tappaniana, To examine visually how it compares with

the other specimens, we plotted the pseudo-factor scores

(calculated from lectotype measurements using results of

FA on the 53 specimens, rather than including it in an

analysis with the other 53 specimens) for the lectotype

where it would appear on the factor plots gene rated for

T. A. Pearce, M. C. Fields, K. Kurita, 2007

Page 69

Table 1. Variables recorded on Gastrocopta pentodon and G. tappaniana. Derived variables relate to shape. Bold p-values indicate

variables showing significant difference (Mann-Whitney U test) between forms as identified by Vanatta and Pilsbry (1906). Measured

variables are in millimeters.

Variable Type Description p-Value between forms

aph/bdh Derived Aperture height/body whorl height 0.6863

aph/h Derived Aperture height/shell height 0.0629

aph Measured Aperture height 0.0003

bdh Measured Body whorl he ight 0.0001

bdh/h Derived Body whorl ht/shell ht 0.0145

bdhAwv Derived Body whorl height/shell width 0.1948

h Measured Shell ht 0.0660

hat Measured Hat height (height above penultimate whorl) 0.2464

hat/h Derived Hat height/shell height 0.0029

hat/spr Derived Hat height/spire (spire = hat height + penultimate 0.0007

whorl height)

pnilth Measured Penultimate whorl height 0.0126

pnlth/bdh Derived Penultimate whorl height/body whorl height 0.6100

pnith/h Derived Penultimate whorl height/shell height 0.2177

pnlth/hat Derived Penultimate whorl he ight/hat 0.0007

pulth/pnltw Derived Penultimate whorl he .ight/penultimate whorl width 0.9915

pnitw Measured Penultimate whorl width 0.0002

pnitw/h Derived Penultimate whorl width/shell height 0.0742

pnitw/w Derived Penultimate whorl width/shell width 0.7823

tthxpar Counted Number of teeth excluding those on the parietal 0.0000

(upper) wall

w Measured Shell width 0.0002

wh Derived Shell width/shell height 0.1313

just the 53 specimens. Second, we performed DFA to

determine how it classified.

Moisture Association of the Forms: To examine the

relationship of the forms with the moisture in their en-

vironment, we calculated a surrogate index of wetness

using plants noted at the collecting sites. Because plant

species differ in their long-term moisture requirements,

estimating éaveninental moisture from plant data

should provide a biologically meaningful measure of

long-term moisture av: ailability.

Wetland facultative indicator status of many North

American plant species is available at USDA-NRCS

(2004) by geographical regions. The plants are assigned

to one of five categories depending on how obligate od

they are to wet lands. For each plant species at a locality,

we assigned a score of 5 for the wettest facultative indi-

cator rane 1 to the driest, and 2, 3, or 4 to the three

intermediate ranks on the website (adding 0.3 for a “+”

and subtracting 0.3 for a “—"). We calculated the mean

facultative indicator rank of all the plants recorded at

each locality and used this mean as a surrogate index of

wetness at the locality.

To test the hypothesis that Gastrocopta pentodon oc-

curs in drier and G. tappaniana in moister areas we di-

vided the wetness score for specimens into three parts,

with the middle part representing 25% of the specimens,

and the other two parts being approximately equally di-

vided. Then we examined the number of individuals in

each of the three wetness areas as a function of their DF

scores.

Geographical Distribution of the Forms: We ex-

amined a map of localities of specimens identified by the

DF to see whether the two forms tended to show inter-

pretable geographic patterns on the peninsula. Of the

130 samples, 106 were only Gastrocopta pentodon or G.

tappaniana; the 16 that were predominantly G. pentodon

were scored as G. pentodon, the 2 that were predomi-

nantly G. tappaniana were scored as G. tappaniana, and

we omitted the 5 samples containing equal numbers of

the two forms. We recognize that geographical location

and wetness might not be independent.

RESULTS

Distinctness of 53 Specimens: DFA of the 53 speci-

mens as defined by Vanatta and Pilsbry (1906) deter-

mined coefficients for four ee to separate the two

forms of Gastrocopta (Table 2). To use these coefficients

to determine an unknown spe cimen, first multiply each

DF coefficient by the corresponding variable value for

that specimen, and then sum all these results. If the

result is negative, then the specimen is more likely G

pentodon, and if greater than zero, the specimen is more

likely G. tappaniana. For example, specimen 1 in Vanatta

and Pilsbry (1906) has the following measurements

(mm)/count: bdh = 0.942. hat/h = 0.267, pnith/hat =

0.645, tthxpar = 4. Multiplying these ae by the DF

coveeut sad adding them yields —8.068, indicating

that the specimen classifies as G. pentodon.

The DF correctly classified 51 (96.2%) of the 53 speci-

Page 70

THE NAUTILUS, Vol. 121, No. 2

tthxpar

Figure 2.

aperture height [measured from base of outer lip to midpoint of

callus connecting the two lip insertions], bdh = body whorl

Variables measured on Gastrocopta spp.: aph =

height, h = shell height, hat = height of shell above penulti-

mate whorl, pnlth = penultimate whorl height, pnltw = pen-

ultimate whorl width, tthxpar = number of apertural teeth

excluding those on the parietal (upper) wall, w = shell width.

Dimensions are measured perpendicular or parallel to axis of

coiling.

mens that had been used to create the function: 39 of the

41 Vanatta and Pilsbry-identified G. pentodon classified

as identified, and all 12 of the 12 G. tappaniana classified

as identified.

In FA, of the 53 specimens, the first three eigenvalues

using the four variables explained 98.1% of the variation.

Two shape variables (hat/h, pnlth/hat) loaded strongly on

factor 1, one size variable (bdh) loaded strongly on factor

2, and o one variable (tthxpar) loaded strongly on factor 3

(Table 3). This result, in which shape loaded strongly on

factor . contrasts with the usual FA pattern in ich

factor | is size and factor 2 is shape. The four variables

were all important for explaining variation in the dataset,

as evidenced by high communality scores (sum of

Table 2. Discriminant function (DF) coefficients for separat-

ing the two forms of Gastrocopta using four variables

Variable Coefficients

Constant =fi071]

bdh 26.82807

hat/h 70

pnilth/hat 33.85039

tthxpat 0.92107

Table 3. Loadings for the four variables on the three factors

from factor analysis (FA) of 41 Gastrocopta pentodon and 12 G.

tappaniana specimens illustrated by Vanatta and Pilsbry

(1906), using the rotated factor method. Bold loadings indicate

the main variables loading on each factor. Percent variance

explained is after varimax rotation.

Factor | Factor 2 Factor 3

pnilth/hat 0.96674 0.00425 0.16676

hat/h -0.96086 —0.08634 —0.17608

bdh 0.0465] 0.98561 0.16166

tthxpar 0.21990 0.17578 0.95955

Variance explained 1.908 1.010 1.006

squares of factor loadings [SS], ranging from 0.961 to

1.000). Figure 3 plots specimens on factors 1 and 2, and

factors 2 and 3, showing that G. tappaniana (solid

squares) occupies a portion of morphospace at the edge

of and somewhat overlapping with G. pentodon (hollow

diamonds). Thus, although the two forms overlap, they

generally occupy separate portions of morphospace.

Shells scoring higher on factor 1 have a relatively

smaller hat to otal height, and a relatively taller penul-

timate whorl relative to hat height. Shells scoring higher

on factor 2 have a larger body whorl height. The third

factor reflects aueaber of teeth, and shells classified as

Gastrocopta tappaniana had 7 non-parietal teeth, in con-

trast to G. pentodon, which had a variable number of

teeth.

Distinetness of 577 Specimens: — Applying the DF

coefficients to the Delmarva specimens classified 475

specimens as Gastrocopta pentodon and 102 as G. tap-

paniana.

In FA of the 577 Delmarva specimens, the first three

eigenvalues using the four variables selected by the DFA

explained 98.0% of the variation. Two sh ape variables

(hat/h, pnlth/hat) loaded strongly on factor 1, one size

variable (bdh) loaded strongly on factor 2, and one vari-

able ( (tthxpi ir) loaded strongly on factor 3 (Table 4). The

four variables were all important for ¢ epbinne ieee

in the dataset, as evidenced by high communality scores

(SS ranging from 0.957 to 1. 000). Figure 4 plots speci-

mens on factors | and 2, and factors 2 and 3, showing

that the two species occupy generally separate portions

of morphospace with minimal ove srlap.

Interpretation of factors 1 and 2 are the same as for

the Vanatta and Pilsbry results. The third factor, largely

reflecting number of teeth, shows variability in both

forms, but shells classified as Gastrocopta tappaniana

tend not to have the minimum number of teeth. The

specimens plot in distinct columns in this analysis be-

cause number of teeth was discreet: the columns appear

more distinct in this analysis than in the analysis of

Vanatta and Pilsbry data because teeth loaded much

more strongly on factor 3 in this Delmarva analysis.

Distinctness in Sympatry and Type Material: The

species composition of samples from Delmarva (contain-

T. A. Pearce, M. C. Fields, kK. Kurita, 2007 Page 7]

37 : 3

* *

2 2

| |

° ° |

° a ° ° a

-_ 9 -_—

g ~ 2" a - oe ean

an a an a

° 2 } a bal Py = ° ‘ 5 Pa S

5 0 S90 ° . Mss 5 f © 990 O% :

uw 2 6 ° WL } o 8

“1 oo O° ° ° ©

; ° | °

: ° 3° % }° - °

22 , 2 ‘ a

37(

-3,-+— = 3. SE) — —_

-3 -2 -1 0 1 2 3 -3 -2 -1 0 1 2 3

Factor 1 (shape) Factor 3 (teeth)

Figure 3. | Factor analysis (FA) plots of 53 specimens on factors | and 2 (left) and factors 2 and 3 (right). Hollow diamonds represent

Gastrocopta pentodon and solid squares are G, pelnaaeie as identified by Vanatta and Pilsbry (1906). The shell image shows, to the

right on factor 1, a relatively smaller hat to total height and a relatively taller penultimate whorl relative to hat height: higher on factor

2 shows a larger body w horl he ight; to the oe on factor 3 shows shells with more apertural teeth. Asterisk (* ) indicates position of

lectotype of G tappaniana and plus sy mbol (+) indicates position of the lectotype of G. carnegici. Numbers beside shell images

correspond to specimen numbers from eae and Pilsbry (1906).

ing 5 or more specimens) was non-random. Samples

tended to be ean primarily of one species or the

other (Figure 5), instead of mostly mixed samples that

would be. expected with random mixing. By the DF, 28 of

39 samples were composed of a single : species. In testing

whether the two forms maintain their distinctness in

sympatry, we examined the FA plot (factors | and 2) of

samples from Delmarva that the DF identified to contain

both forms (mixed samples). Figure 6 shows good sepa-

ration of the two coexisting forms, supporting the idea

that the two forms are separate species. The separation

of the two forms in the mixed-only samples is as good as

the separation in all samples (compare Figures 4 ‘anal 6).

We performed two tests to determine whether one of

the two morphologically discrete forms we identified on

Delmarva corresponds to type material of Gastrocopta

Table 4. Loadings for the four variables on the three factors

from factor analysis (FA) of 577 specimens from the Delmarva

Peninsula, using the rotated factor method. Bold and under-

lined loadings indicate the main variables loading on

each factor. ;

tappaniana (we could not locate type material of G. pen-

todon). First, on the FA plots (Figure 3), the lectotype of

G. tappaniana was located in the se he of the ‘graph

with specimens classified by Vanatta and Pilsbry as G.

tappaniana. Second, the lectotype of G. tappaniana was

classified as G. tappaniana by the DF.

Moisture Association and Geographical Distribu-

tion of Forms on Delmarva: The 22 samples having

5 or more specimens with only Gastroc ‘opta pe mntodon las

classified by the DF) were from areas drier (wetness

factor X = 2.62, sd = 0.46) than the 6 samples having 5 or

mone specimens with only G. tappaniana (wetness X =

3.14, sd = 0.30) (t-test, p = 0.006). Mean wetness factor

foe all 471 specimens classified by the DF as G. pentodon

was 2.59 (sd = 0.44) and that for the 100 G. tappaniana

was 2.95 (sd = 0.44, t-test, p < 0.00005). The histogram

in Figure 7 shows the number of individuals by their DF

score separately for each of three wetness classes. The

vast majority of G. fappaniana (positive DF) occur in

medium or wet areas. On the other hand, G. pentodon

(negative DF) occur in a wide range of moisture, with a

tendency to be in drier areas. In samples from medium

wet areas, bimodality of forms is evident, with a hint of

Factor | Factor 2 Factor 3

—— a siaebes bimodality in the sample s from dry and wet areas.

pnith/hat 0.97099 0.13158 0.03672 A map of localities of specimens identified by DF

hat/h -0.94900 -0.23783 -0.01636 shows Gastrocopta pentodon widespread throughout the

en 022s! 0.97414 pig peninsula but essentially absent from the southeastern

tthxpar ; 0 03081 0.00861 0.99948 coast and barrier islands. In contrast, s: unples containing

Variance explained 1.895 1.023 1.001]

G. tappaniana were most common oe the Atlantic

Page 72

THE NAUTILUS, Vol. 121, No. 2

4 3 -2 -1 0 1 2 3 4

Factor 1 (shape)

Figure 4.

Factor 2 (size)

oO —_

OGD Mo

Rocce om Mm ae

Ot. 20S

aR,

4 -3 -2 -1 0 1 2 3 4

Factor 3 (teeth)

Factor analysis (FA) plots of 577 Gastrocopta specimens from the Delmarva Peninsula plotted on factors 1 and 2 (left)

and factors 2 and 3 (right). The same variables were used as in analysis of Vanatta and Pilsbry data. Hollow diamonds represent

specimens classified as G. pentodon and solid squares those cle issified as G tappaniana.

Coast in southeastern Delmarva and closer to major wa-

ter bodies (Figure §). Interestingly, four Atlantic coast

samples containing 5 or more specimens of only G. tap-

paniana were from four of the wettest areas, as deter-

mined by the plant moisture associations.

DISCUSSION

These results indicate that the two rons of Gastrocopta

can be considered separate spe cies. DFA successfully

discriminated two groups. The 53 specimens in Vani ta

and Pilsbry (1906) could be separated using 4 variables

and the DF classified 96% of specimens as defined. The

two forms were generally distinct by FA de spite, some

morphological overlap. When applie d to the 577 Del-

marva specimens, the DF identified both species on Del-

marva. FA of the Delmarva specimens showed the two

species occupying generally separate portions of mor-

phospace with minimal ove le ap.

~ 20

fe)

O 10

° ° ro) [=] fo} fo) (=) ro) co) =)

. Y oy SS 8 es

oO ro) oO fo) -) fo) [=] f-) fo} ai

= AQ o +t wo oO i ire) Q

Percent of G. tappaniana

Figure 5. Proportion of specimens that were Gastrocopta

£ I I }

ppaniana in the 39 samples containing 5 or more specimens

Even stronger evidence of separate species is that the

forms matntined their distinct morphologies where they

coexist. Gastrocopta pe ntodon and G. tappantana from

the Delmarva Peninsula showed no evidence of conver-

gent morphology in sympatry. Moreover, the forms

maintained their separate morphologies in samples from

5 + -

3 | 2

@ 24 ;

o |

2 | °

N 4 oe)

ro} 20% |

Oo 25 D 2

4 0 | * 50a on

WL | of ° Og a @

6 0s °

4 4 o, F 5 8% sm a

| eee. @

2 | 7

Rogen

4 #-3 -2 -1 0 1 2 3 4

Factor 1 (shape)

Figure 6. Factor analysis (FA) plot of 180 Gastrocopta speci-

mens from the Delmarva Peninsula of the Ll mixed samples

determined to contain both forms.

T. A. Pearce, M. C. Fields, K. Kurita, 2007

jee)

Page

areas of intermediate wetness, further supporting the

idea that the two forms are distinct species.

The dataset from Vanatta and Pilsbry (1906) and ours

from Delmarva showed similar results, strengthening the

conclusion that two species exist. The Delmarva dataset

was more geographically restricted, but had more speci-

mens, while the Vanatta and Pilsbry dataset was geo-

graphically broader with fewer a cimens. Sources ee all

Gastroc opta tappaniana included by Vanatta and Pilsbry

(1906) are from locations more concentrated in NE USA,

from Washington, DC to Maine ( (excepting one G. tap-

paniana from Arizona), whereas their G. pentodon were

more W idespread, from Texas to Iowa and Florida to

Maine. This congruence despite different geographic

sampling suggests that shell morphology does not vary

significantly across geography. Bimodality of morphology

supports the hypothesis of two species on Delmarva and

throughout the eastern USA.

Gastrocopta tappaniana on Delmarva occurred in

samples from moister areas, whereas G. pentodon oc-

curred in samples from a broad range of wetness. Sample

wetness was unimodal and continuous, both for all

samples and for samples containing at least 5 specimens,

such that a difference in habitat by species does not seem

to be an artifact of sample choice. This result confirms

reports of Sterki (1906) and Pilsbry (1948) that G. tap-

paniana occurs in moister areas. However, in contrast to

authors who characterized G. pentodon as being in drier

areas, our results agree with the report of Hubricht

(1985) that G. pentodon has a wider moisture range and

can overlap in habitat wetness with G. tappaniana.

Geographically, G. tappaniana tended to occur on the

SE part of the peninsula, and closer to water bodies,

whereas G. pentodon was more evenly distributed across

the peninsula, including its central ine Moistness seems

to be an important ‘afinenise on the distribution of G.

guia but some other aspect of geography might

also play a role in its distribution, considering that fhe

Atlantic coast samples from the SE part of the peninsula

were the moistest samples.

We examined the type specimen of Gastrocopta tap-

Individuals

-12 to -10 I

paniana (Museum of Comparative Zoology, Harvard,

lectotype MCZ 186171, paratype [not seen] 186172) but

i re unable to locate type material of G. pe ntodon. For

,. tappaniana, Pilsbry (1948: S89) stated “Type locality,

oo coll. Amherst College”, implying that an un-

specified kind of type or types was present at Amherst.

Bequaert and Miller (1973) observed that the holotype

[sic] was transferred from Amherst to MCZ, mistakenly

stating that Pilsbry (1948) referred to it as a holotype.

C lench( 1965) noted that the type locality for G. tappa-

niana is Roscoe, Coshocton Co., Ohio, and not Vermont

as some writers have assumed. He chose a lectotype for

G. tappaniana: “G. tappaniana appears to be a synonym

of G. pentodon (Say); the lectotype (here selected) is

nearest to the figure of pentodon given on pl. 3, fig. 7

1916, Manual oF Conch. (2) 24: 33, and not io fig. 9

which is given as tappaniana. This same plate was re-

published i in Land Mollusca of North America, vol. 2, pt.

2, fig. 477, p. S87, Mono. no. 3, Acad. Nat. Sci. Phila-

delphia, 1948.”

Ve find it peculiar that Clench intended to choose a

lectotype specimen of Gastrocopta tappaniana that was

most like G. pentodon. Interestingly, despite Clench’s

bias in his choice, the lectotype is consistent with G.

tappaniana per both of our tests: it is located among the

G. tappaniana specimens on factor plots, and it Glascihed

as such by the DF derived in this paper.

Vanatta and Pilsbry (1906) did not reveal their objec-

tive criteria for classifyi ing the two forms, so researchers

using their paper must accept their two groups at face

vale. We recognize that DFA, by its nature, will find

differences between nearly any groups. Therefore, to test

whether the groups defined by Vanatta and Pilsbry

(1906) are different from randomly selected eee we

compared the percent correct classification of the DFA

results for the groups as recognized by Vanatta and Pils-

bry, to 10 randomized datasets with 41 specimens in one

group and 12 in the other. In contrast to the 94.3% correct

classification of groups as they defined, the randomized

data sets averaged 70.9% correct classification. The higher

correct classification of the non-randomized groups gives us

Midry

G. tappaniana

med

A wet

0 to2 nal

2to4 L

ato6 ba

8 to 10

10 to 12

Discriminant Function Score

Figure 7.

classes.

Histogram showing the number of individuals by discriminant function (DF) score, separately for each of three wetness

Page 74

THE NAUTILUS, Vol. 121, No. 2

Figure 8.

samples of Gastrocopta pentodon (open circles) and G. tappa-

niana (solid circles). Symbols on the map reflect majority rule (24

samples contained both G. pentodon and G. tappaniana; the 5

samples with equal numbers of the two species are omitted).

Delmarva Peninsula showing locations of 130

confidence that the two groups as defined by Vanatta and

Pilsbry are non-randoin ( (p < 0.000001).

Some disagreement exists in the literature regarding

the correct classification of Gastrocopta carnegici Sterki,

1916. Sterki (1916a) described the new species as G.

minuta, but subsequently changed the name to G, carnegiet

because G. minuta was preoccupied (Sterki, 1916b). Sterki

1916a) did not designate a holotype among the three

specimens, although Pilsbry (1948: S90, 892, fig. 450: 5)

designated a lectotype by referring to the only unbroken

specimen as the type. Turge -on et al. (1998) listed G. car-

negiei as a valid species. Sterki (1916a) had indicated that

G. carnegici is similar to G. tappaniana, but Hubricht

(1985) considered G. carnegie to be a synonym of G. pen-

todon. The DF, when applied to measurements taken from

the lectotype illustration of G. carnegiei presented by

Pilsbry (1948: 892, fig. 480, image | 5), classified the speci-

men as G. pentodon uid the position it occupies in Fig-

ure 3 (indicated by the plus sy seer suggest it is @losete

to G. pentodon. The squat shape of ake lectotype figure

is like G. tappaniana, with its large body whorl, bak it

differs by a smaller shell and only 5 apertural teeth; the

few whorls suggest that it might be an abnormal speci-

men. Although we were unable to locate the lectotype of

G. carnegiei at Carnegie Museum of Natural History, we

located the two paralectotypes, which are broken, as noted

by Pilsbry (1948), so their relevant shape measurements

cannot be discerned. This study suggests that G. carnegiei

falls within the range of variation of G. pentodon.

Although separating the two species visually can be

difficult, some characteristics might be helpful at distin-

guishing between them. Gastrocopta tappaniana tended

to be larger (wider shell, wider penultimate whorl width,

and taller body whorl) and regarding shape, tended to

have a shorter hat (section doov e the penultimate whorl)

relative to both shell height and spire height (hat/h, hat/

spr), and a relatively taller body whorl. Regarding num-

ber of teeth, Vanatta and Pilsbry (1906) stated that all G.

tappaniana have 7 teeth (excluding teeth on the parietal

wall), whereas G. pentodon have 5-9 teeth. However,

according to their eainee the number of teeth for their

G. pentodon ranged from 2-9. In contrast to their sug-

gestion that tooth number might be a useful distinguish-

ing character, our analysis suggested that number of

teeth was not useful for separating species on the Del-

marva Peninsula. A character that could be evaluated in

future studies is whether the lower palatal fold of G.

tappaniana is “usually not so long and entering as in G.

pentodon” (Vanatta and Pilsbry, 1906; Pilsbry, 1945).

Future work could aes molecular data such as

DNA sequences to verify these conclusions. Another av-

enue to address environmental influence on morphology

would be to raise sibling specimens under different en-

vironmental conditions.

ACKNOWLEDGMENTS

The U.S. National Science Foundation (DEB 9972026)

funded part of this project. K. Fewlass Kling and others

helped collect Delmarva samples and A. W. Doolittle

helped with fieldwork. A. S. Italia, L. Brink Beebe, D. L.

Scott, A. Gathers, and others diligently picked snails

from leaf litter samples. We are grateful to Adam J,

Baldinger, MCZ, for loan of the lectotype of Gastrocopta

tappaniana. We are grateful to Amanda E. Zimmerman

for expertise in producing the figures. Reviews by Jeff C,

Nekola and an anonymous reviewer contributed substan-

tially to improving this paper.

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Turgeon, D. D., J. F. Quinn, Jr., 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. Schel-

tema, F. G. Thompson, M. Vecchione, and J. D. Williams.

1998. Common and Scientific Names of Aquatic Inverte-

brates from the United States and Canada. Mollusks, 2nd

edition. American Fisheries Society Special Publication

26, Bethesda, 526 pp.

USDA-NRCS. 2004. The PLANTS Database, Version 3.5

(http://plants.usda.gov). National Plant Data Center,

Baton Rouge, Louisiana 70874-4490, USA. Consulted on

28 Feb. 2007.

Vanatta, E. G. and H. A. Pilsbry. 1906. On Bifidaria pentodon

and its allies. The Nautilus 19: 121-128.

THE NAUTILUS 121(2):76-89, 2007

Page 76

Upper Jurassic Pleurotomariidae (Gastropoda) from

southwestern Madagascar

Steffen Kiel

Earth Sciences

M. G. Harasewych

Department of Invertebrate Zoology

National Museum of Natural History

Smithsonian Institution

P.O. Box 37012

Washington, DC 20013-

[email protected]

Leeds LS2 9JT

7012 USA and

University of Leeds

UNITED KINGDOM

Department of Paleobiology

National Museum of Natural History

Smithsonian Institution

Washington, DC 20013-

[email protected]

7012 USA

ABSTRACT

This paper describes four new species of Upper Jurassic Pleu-

rotomariidae from southwestern Madagascar: Obornella

thompsonorum, Bathrotomaria annejoffeae, Bathrotomaria be-

detteae, and Leptomaria takahashii. In addition, the previously

described Leptomaria texta Delpey, 1948, is reassigned to the

genus Obornella. Comparison of this fauna with that of the

geographically proximal Kutsch region of northwestern India

reveals it to consist of representatives of wide-ranging Tethyan

genera, but also to exhibit strong endemism at the species level.

INTRODUCTION

A substantial number of well preserved Upper Jurassic

pleurotomariid gastropods have recently been uncovered

as a byproduct of commercial mining for ammonites in

southwestern Madagascar. Six specimens, representing

four species, were leadly made available to us for study

by Mr. Chris 1 ‘akahashi. The pleurotomariids and am-

monites were dug by village rs from pit quarrie s near the

town of Zakar ha, in southwe sstern Madagascar.

A survey of the literature on the Mesozoic gastropod

fauna of Madagascar (e.g., De Ipey y, 1948; Collignon,

1949: Collignon, 1959; Kiel, 2006) revealed the majority

of pleurotomariid species known from Madagascar to be

of Cretaceous age, with only a single species, Leptomaria

texta Delpey, 1948, reported from Jurassic strata. Al-

though the sample available to us is of limited size and

stratigraphic range, it expands our insight into the Juras-

: pleurotomariid fauna of Madagascar. Like the well

‘loc ‘umented Jurassic pleurotom: wad fauna of the Kutsch

also spelled Kachc hh region of western India (Jaitley et

al., 2000; Das, 2002; Das et al., 2005), the Madagascar

fauna reveals Tethyan affinities at the generic level, yet

exhibits endemism at the species level. Both faunas

evolved an increasing endemism within the Indo-

Madagascan faunal province that was formed as the

Tethys Ocean widened between Laurasia and eastern

Gondwana, and a seaway started to develop between

East and West Gondwana in the latest Jurassic (Titho-

nian) (Hay et al., 1999; Shome et al., 2004).

The scope of the present study is to review the Oxfor-

dian (Upper Jurassic) pleurotomariid fauna of Madagas-

car, describe five species, four of them new, and to re-

view the relationships and biogeography of these pleu-

rotomariids.

GEOLOGICAL SETTING

Rifting between Africa and Madagascar produced three

large sedimentary basins along the west coast of Mada-

gascar. These are, from North to South, the Ambilobe (or

Diego), Mahajanga (also spelled Majunga), and Moron-

dava basins. Sedimentation in this region commenced in

the Carboniferous with the deposition of the Gondwanan

Karoo sequences and equivalents. The first marine de-

posits resulting from the break-up of the Gondwana su-

percontinent are of Toarcian (late Lower Jurassic) age.

From then on, alternating shallow marine, brackish, and

fluviatile sediments were deposited in these basins. The

pleurotomariids described here are from the Morondava

basin, the southernmost of the three basins. Bio- and

lithostratigraphic work in this basin is difficult because

outcrops are few, index fossils are often not available, and

measurable sections are usually short and difficult to cor-

relate with each other. Consequently, only few litho-

str: tigraphic units have been given formation names so far

(Besairie and Collignon, 1972: I uger et al., 1994; Geiger

and Schweigert, 2006),

The quarries that produced the specimens described

here are located to the west of the town Zakaraha (also

M. G. Harasewych and S. Kiel, 2007

spelled Sakkara), one of them north of the Fiherenana

River, the other to the south of it (Figures 1, 2). The

fossiliferous lavers are thin and consist of pink-yellow

iron-oolithic limestones, and are overlain by grey mud-

stones (K. Bandel, pers. comm., 2007). The southern

locality is very close, if not identical, with the “Amparam-

bato section (VIb)” of Geiger and Schweigert (2006: 99)

which was characterized by them as “a highly fossilifer-

ous iron-oolitic limestone and sandstone bed with thin

mud-stone interlayers.” Accordingly, the oolitic lime-

stones contain an abundant ammonite fauna; the over-

laying mudstones contain nodosariid Foraminifera and

ostracods (Geiger and Schweigert, 2006). The northern

quarry is geographically close to the “Middle-Upper

Oxfordian Ankilimena section (XI)” of Geiger and

Schweigert (2006). This section is characterized by re-

current iron-oolitic limestones, which contain ammo-

nites, rhynchonellids, bivalves, belemnites, echinoderms,

and wood debris (Geiger and Schweigert, 2006: 103).

Although Geiger and Schweigert (2006) did not report

A5et

rs no?

wane

Figures 1,2. Location of collection sites. 1. Detailed map of

two localities in southwestern Madagascar. 2. Location of sites

on a map of land masses during the Late Jurassic.

Page 77

(=)

pleurotomariids or other gastropods from these sections,

the remaining fossil content, their geographic position,

and their lithology agrees well with that observed at the

ammonite quarries visited by K. Bandel (pers. comm.,

2007).

The ammonite fauna of these two localities, especially

the presence of Dhosaites cf. primus Collignon, 1959,

suggests a ‘middle to upper Argovian (early Oxfordian)’

age (Collignon, 1959; H. Keupp, pers. comm., 2004).

The term ‘Argovian’ has been abandoned due to its in-

consistent usage, but largely falls within the range of the

Oxfordian (Zeiss, 2003). The sediments considered as

‘Argovian’ by Besairie and Collignon (1972) correlate

with those mapped as late Callovian-early Oxfordian by

Geiger and Schweigert (2006). Thus, the pleurotomariids

described here are most probably of Oxfordian (Upper

Jurassic) age.

SUPRASPECIFIC CLASSIFICATION

WITHIN PLEUROTOMARIIDAE

The number of genera and subgenera currently recog-

nized within the family Pleurotomariidae (Appendix 1)

has nearly doubled since the family was reviewed in the

Treatise of Invertebrate Zoology (Knight et al., 1960). Of

the 21 genera and subgenera now recognized (Figure 3),

16 are Mesozoic. Of these, five are restricted to the Tri-

assic, three to the Jurassic, and only a single subgenus to

the Cretaceous. At present, Leptomaria and Conoto-

maria are the only genera that are recognized as having

survived from the Mesozoic into the Cenozoic. Of the

seven Cenozoic genera, four survive in the Recent fauna.

According to the literature surveyed, there is no chrono-

logical overlap between the Mesozoic genera and the

Cenozoic genera.

As noted by a number of researchers (e.g., Hickman,

1976; 1094: Szab6, 1980: 49; Das, 2002: 99) fossil pleu-

rotomariids are difficult to classify objectively, since the

criteria upon which fossil pleurotomariid genera are de-

fined differ conspicuously from those applied to Ceno-

zoic genera. The monophyly and phylogenetic relation-

ships of the surviving Cenozoic genera have been con-

firmed using molecular data from living representatives

(e.g., Harasewych et al, 1997; Harasewych, 2002). By

contrast, the relationships of Mesozoic genera and the

species assigned to them are far less certain, as generic

classification tends to be based on relatively few con-

spicuous morphological features (Table 1) especially

those that are most easily derived from poorly preserved

specimens and external or internal molds, while other

characters are unconstrained and may vary widely. Szabé

(1980: 49) commented that “almost all genera can be

identified with certainty” on the basis of the shape of the

whorl section and the surface of the whorls, as well as the

position and width of the selenizone. Conti and Szabd

(1987: 43) raised a question as to the significance of the

presence or absence of an umbilicus in pleurotomariid

Page 78

THE NAUTILUS, Vol. 121, No. 2

MESOZOIC

TRIASSIC JURASSIC |CRETACEOUS

io)

Figure 3. Geological ranges of the

{¢ ssil rect ord

classification. The presence or absence of an umbilicus is

sufficient to distinguish the most basal dichotomy among

living Pleurotomariidae, yet this feature remains uncon-

strained and may vary widely within most Mesozoic gen-

era and even within some “species” as they are currently

diagnosed. It is therefore not surprising that fossil species

are frequently reassigned from one genus to another

e.g., Pyrgotrochus to Laevitomaria Conti and Szabo,

LOST

2005

16: Perotrochus to Leptomaria, see Das et al.,

33]

served specimens become available

especially as more numerous and better pre-

ausd0bIlO

genera and subgenera within the family Pleurotomariidae,

CENOZOIC

QUad0I|4

QU800}SI9|q

GENERA OF

PLEUROTOMARIIDAE

Mamoeatomaria Begg and Grant-Mackie, 2003

Tahua Begg and Grant-Mackie, 2003

Murihikua Begg and Grant-Mackie, 2003

Pleurotomaria Defrance, 1826

Ornatospira Pan, 1982

Stuorella Kittl, 1891

Talantodiscus P. Fischer, 1885

Pyrgotrochus P. Fischer, 1885

Anodotomaria Szabo, 1980

Bathrotomaria Cox, 1956

Cyclostomaria Szabo, 1980

Laevitomaria Conti and Szabo, 1987

Obornella Cox, 1959

Conotomaria Cox, 1959

Leptomaria E. Eudes-Deslongchamps, 1864

Indomaria Das, 2002

Chelotia Bayle in P. Fischer, 1885

Entemnotrochus P. Fischer, 1885

Perotrochus P. Fischer, 1885

Bayerotrochus Harasewych, 2002

Mikadotrochus Lindholm, 1927

arranged by first occurrence in the

SYSTEMATICS

Family Pleurotomariidae Swainson, 1S40

Genus Obornella Cox, 1959

Obornella Cox, 1959: 238.

Type Species: Pleurotomaria plicopunctata J. é

Eudes-Deslongchamps, 1849 (By original designe ney

Bajocian (Middle Jurassic) of France.

Diagnosis: The genus Obornella is characterized by a

shell that is low turbiniform to sublenticular, with a nar-

Harasewych and S. Kiel, 200

—

spray pur MOTeYS yuasqe XOAUO. [LoyAr-prut

sp109 yeadg ‘peotg 10 YRoAy A[BUO.QS sugqnqy yuosqy yuosqy yuasqy QAOqY — popls-P promos snyoo.jo1ahivg

sp.l09 MOTeYS quasqe [LOYA\-plUt

jeuds osopon ‘proig, 10 yRaAy XOAUOD) Suyuqy quasqy yuosqy juosqy MOLIG — Papls-p fRolloy — smpoo.gopvyiyy

asopou

10 out MO]EYS quasqr [LoTp\-plul

spi09 peurdg “proig IO YROA\ XOAUOD) suyqqnqy quasqy quasqy yuosqy Mopaq ysuf— papts-F jeowog SM IOMOMIT

spioo pur doop yuasqe [Loya-prut

sprampy pemdg = ‘MoumN 10. YRoA\ rl ysnya opU yuasqy yuosqy QA0qY — popls-F [ROOD SNYIOAJOULWAPUTT

sptoo daap quasqe [HOYA\-prutt

jetids asopoN =“ MOLIBN LO. YROAY Wi ysty a opU yuosqy yuasqy QAOGY — papls-f [Rolulon pyopayy

aqe]jeours MOTEYS yuasqe x apr 0}

sptoo prudg = ‘Mourn 10. YRay A[BUO.S Suyquqy quasqy juasqy uosqy [LOY A-plUL PY Popts-F [Rowwo9g pupwojday

doop Opi 0} [toyar-prut

spi0o yeudg = ‘more Usa wll sup] yuosqy juosqy yuosqy aAoqe 10 FY papts-c peowwod DLIDWOJOUDT

URUTULOp

avysoo MOTPYS juosqr. XOAUOK

‘oPRTPOOURD, = SMOLIBNY LO ROA, A[BUOLYS SUNY Asa. Juosa.tg quesqy = Atoyditod uwon papts-p [row DIJOULOGO

yuasqe [LoyA\-prut

sp0o peardg MOLE IO YROA\ Wa possordury opi yuasqy yuasqy MOOG — Popls-F jeomtoy DUDUONAIVD'T]

spear peatids asopou MO.LIRU Atoydiiod

‘avysoo [RIXy MO.LIRN \uaso.g Rl ysnyy 0 Juasqy UIs yuasqy aaoqe ysuf papts-F [Rolo DIJQLON]S

spurq asopou asopou MO.LIVU juosoid [LOYPA\-piut

‘spa09 ped prog \uasotg ra ysnpy 10 yuasqy yuasotg SOpON MOLI — popls-p [Romo smyoospod.thg

}USUTULOp

peixe

‘oqyyyjoourg, MO.LIEN quasqy Wp Apron sunynqy op quasqy yuasqy [LOYA\-PIU PY papis-p — [ROLWOD AO] v.ndsoppuUui<a

XOAULOD Aroyditod quosqe

aqye]jaourD MOLI quasqy AJBUOYS MOPO yuasqy — ulasag yuasqy [LOY M-plUt FY popts-p peoluwory DUYDL.

quasqe

juiosqe spooupoas

ayRIfoour|” MOLIEN 10 ROA d poaoo.ry é quosqy SOpON — Lopphoys Mojog. —poapis-G aywVpVly, — PLEDUO|DAOLUD IY

sp09 perdg MO.LIEN quosqy Fi Ssunnqy d quasqy SLI JRIXY — oppHoYys Mojog. —popts-G ompR Duy yoy

Squt

[RIXe ROM yuasoid LULIOF LUG

‘spc09 ped prog d XOAUOT) Suny SARA quosqy quasqy Atoydurod ovoqy — popts-G MO] DLUDUOPUT

qu [ere XOAUOD oyPpHoyRuro LUO] LG.

‘spaoo ped MOLIVN yuosqy A[BUO.S AP[ROAN opi juasqy qyuasqy Atoyditod oaoqy [RAO MO] PLUDUOISOPIND

sapou pur MOT[PYS

spray ead ‘prosg }UISI.1g rll povoo.sy UOSa1g Uosoig SOpON — Loppuoys Mopoq. — popts-G perdi SNOSIPOPUD]D

ayR|fooura juasqr juosqe

to yetids MOLIEN LO ROAA XOAUOT) sugynqy — Auasoig juosqy JUosqy — daopphoys Suopy — popts-G opm pLeptuojo.nyiyg

sproiyy pur juasqr juosqe

spt0o jeudg proig LO YROAK XOAUOT) sugynqy — Auesotg quosqy JUasqy — Loppuoys Moog, Popts-G ope DIIDULOJOPOUuy

sopou pur MOT[RYS juosqe. judosqr juosqe

spaoo jrardg ‘pRrotg, 10 Yost XOAUOD) SuMnqy — Uosetq. — /Uasog SOPON — Lopfroys Mv\opoq. — Popls-G ope DIIDULOJOAND]

aangdynos WS an[nq asrg ong snoyiquiy, Atoyditod — oangdypnos OUOZIUOJOG oyyoud oud SOND

aOPJANG prtoypditoc suope topyuoys LOU MA

SOPpON jeixy

“ULOTy oulfop O7 OATVLOPLT ot ul posu SLOPOVICYO our puv IVPHAVULOJOITL] | ATHY om UIP TAN poziudoodo. AP}UO.Ltno vLouosqus pure VIOUOF) MNQe

Page 80

THE NAUTILUS, Vol. 121, No. 2

rowly open umbilicus and a strongly convex base. Surface

sculpture consists of closely spaced cna costellae

(usually dominant) and spiral threads. The periphery is

commonly crenate. The selenizone is narrow, smooth,

often projecting onto the upper whorl face near the pe-

riphery. The labral slit is short.

Remarks: Obornella is known from strata of Toarcian

(Lower Jurassic) to Oxfordian (Upper Jurassic) age (183

Ma to 156 Ma). Greatest diversity has been documented

from Europe (see Griindel, 2003; Hiigele, 2003), and this

genus has also been reported from northeastern [ran

(Majidifard, 2003) and the Kutch region of western India

(Jaitley et al., 2000; Das et al., 2005).

Obornella texta (Delpey, 1948)

(Figure 4, reproduced from Delpey, 1948: pl. 2, fig. 1)

Leptomaria texta Delpey, 1948: 9, pl. 2, fig. 1

Original Description (Translated): “(Height: 25

mm; diameter 31.5 mm; number of whorls: 5). The strip

[selenizone] is anterior, wide relative to the last turn, and

develops/changes normally until the multicarinate stage,

becoming a little convex. The sculpture is latticed. An

anaes obscured by a lamina pierces the convex base.

It is similar to Pleurotomaria eudora @ Orbigny, 1850,

but the selenizone of this Oxfordian species is concave

between two carinae, which is noticeably different from

the Malagasy form. Argovian of Ankirijy (coll. Houreq).”

Remarks: Unfortunately, Delpey did not specify were

her type material was ‘deposited. Inquiries at the

Muséum national d'Histoire naturelle in Paris as well as

the Université de Paris revealed that the specimens are

not deposited in their collections. The description is

minimal, and the illustration of the single, partial speci-

men (Figure 4) is poor, showing a specimen in which the

selenizone runs along the shell periphery, with the suture

Figure 4.

Obornella texta (Delpey, 1948). Reproduction of

original illustration (Delpey, 1948: pl. 2, fig. 1, as Leptomaria

lexta

adpressed along the lower edge of the selenizone. These

features preclude the inclusion of this species in Lep-

tomaria E. Eudes-Deslongchamps, 1864, which is char-

acterized by convex whorls, with the selenizone at mid-

whorl. It is more likely that this taxon is referable to the

genus Obornella, in which the selenizone is situated

closer to the pe riphery. Delpey’s illustration resembles

the partial specimen (Jaitly et al. 2000: 39, pl. 2, figs. SA,

B) from the contemporary Dhosa Oolite Member ar the

Kutsch region of India that was identified as Obornella

aff, granulata ( (J. Sowerby, 1818). Delpey’s taxon is here

ae to Obornella, but its generic affinities remain

obscure until the type, which is the only known speci-

men, is located.

Obornella thompsonorum new species.

(Figures 5-12)

Description: Shell Mgures 5-12) small (holotype

maximum diameter 42.2 mm, minimum diameter 34.6

mm, height 26.4 mm) low, turbiniform, consisting of ap-

proximately 7 7 teleoconch whorls. Base moderately con-

vex, with narrow umbilicus (Figure 7, u). Spire angle

103-L06°. Spire slightly convex in profile. Suture ad-

pressed, joining previous whorl at or just below periph-

eral bulge (Figure 6, pb). Protoconch and first 3-5 te-

le eseuch whorls eroded, only final 2—4 teleoconch whorls

well preserved. Weak shoulder present on whorls 3-5,

becoming convex, rounded in final two whorls. Axial

sculpture of oblique radial costae most pronounced on

whorls 3-5 (40-50 per whorl), weaker on subsequent

whorls (SO-108 per whorl), producing cancellate gran-

ules at intersections with spiral cords, especially at pe-

a as bulge and on either side of selenizone (Figure

10, sz). Number of strong, simple spiral cords 7-9 be-

mite ss and selenizone, 0-3 on selenizone, 2—4 be-

tween selenizone and peripheral bulge, 21-22 along

base, between peripheral bulge and broad parietal callus

(Figure 7, pe). Selenizone (Figure 10, sz) narrow, con-

vex, with 0-3 spiral cords, and numerous strong to weak

lunulae, situated just above peripheral bulge. Aperture

ovate, roughly perpendicular to coiling axis. Outer lip

smooth, portion below slit offset from portion above slit

by 39°. Slit narrow (~2.5 mm), extending posteriorly 62°

from end of suture. Lip thickest in Solana llar and basal

region, forming broad parietal callus that partly overlaps

the umbilicus.

Type Locality: Zakaraha, near Toliara (also spelled

Tulear), southwestern Mad: iwascar. 6-7 m below surface

on plateau cut by river.

Type Material: Holotype, USNM 534480; paratype 1

USNM 534481: paratype 2 USNM 534482, all from the

type locality.

Age: Oxfordian (Upper Jurassic).

Etymology: This new species honors Jon and Beverly

Thompson for their many contributions and years of ser-

vice to The Bailey-Mi atthews Shell Museum in Sanibel,

Florida.

M. G. Harasewych and S. Kiel, 2007

Page $1

Figures 5-11.

Remarks: The assignment of this new species in the

genus Obornella is provisional. The type species of

Obornelia. Pleurotomaria plicopunctata J. A. Eudes-

Deslongchamps, 1549. from the Bajocian ( (Middle Juras-

sic) of France and England, is near one end of a mor-

phological spectrum (see Hiigele, 2003: fig. 10) that is

distinguished by a low, conical spire and conspicuous

axial fluting along the shell periphery and base. The

other end of this “morphological spectrum is character-

ized by shells with a higher, stepped spire, a rounder

Obornella thompsonorum new species. 5. Apical, 6. apertural, and 7. basal views of the holotype, USNM 534480.

8. Apical and 9. dorsal views of paratype 1, USNM 534481. 10. Details of sculpture between suture and periphery on last three dorsal

whorls of paratype 1. 11. Apical view of paratype 2, USNM 534482. Abbreviations: p, periphery; pb, peripheral bulge: pe, parietal

callus: s, suture; sl, posterior limit of slit; sz, selenizone; u, umbilicus.

aperture, and spiral cords along the base, features remi-

niscent of the genus Pliocene. Obornella thompso-

norum more closely resembles O. trapeza (Hudleston,

1895: pl. 40, figs. 5a, b) and the “elevated variety” of O.

granulata (Sowerby, 1818) illustrated by Hudleston

(1895: pl. 40, figs. la, b) as Pleurotomaria granulata var,

ceelata Deslongchamps, 1848, but differs in having a less

pronounced, more rounded peripheral bulge, a spire that

is stepped along intermediate whorls, “and strongly

beaded sculpture valong s both sides of the selenizone. Two

Page 82

THE NAUTILUS, Vol. 121, No, 2

European species of Obornella had been reported from

the Kutch fauna (Jaitly et al., 2000; Das, et al., 2005).

Obornella wuerttembergensis (Sieberer, 1908) from

Lower Jurassic [upper Bathonian] strata, is much flatter,

lacks a parietal callus, and has a far broader umbilicus

than O. thompsonorum. Specimens of Obornella granu-

lata (Sowerby, 1818) from the Dhosa oolite member of

the Chari Formation [Oxfordian} of Kutsch (Jaitly et al.,

2000: pl. 1, figs. 5-7) resemble O. thompsonorum in

overall proportions, but have more convex whorls be-

tween suture and periphery, far weaker axial sculpture, a

broader umbilicus, and lack the distinctive parietal cal-

lus. Obornella thompsonorum may be distinguished trom

Obornella texta Sate 1948), the only previously de-

scribed pleurotomariid from the Jurassic of Madagascar,

by the position of the selenizone above rather than. along

the periphery, and by having more prevalent sculpture,

especially adjacent to the salenizone.

The shell microstructures found in Obornella thomp-

sonorum (Figure 12) are similar to those of pleuroto-

mariids from the Carboniferous (Batten, 1972), Triassic

(Bandel, 1991), Jurassic (Boggild, 1930), Cretaceous

(Kiel, 2006), ant Recent (Harasewych, 2002), indicating

that shell microstructure is a very conservative character

in this group.

Genus Bathrotomaria Cox, 1956

Type Species: Trochus reticulatus J. Sowerby, 1821.

(By original designation). Kimmeridgian (Upper Juras-

sic) of England.

Diagnosis: Bathrotomaria can be distinguished by its

usually large (to 130 mm), trochiform shell with a spire

that may he elevated to depressed. The umbilicus may be

broad to entirely absent. The whorl profile is usually

angulate and non-tuberculate, with a broad ramp and a

second carina or angulation, just overlapped on the spire.

SEM image of a fracture surface of the shell of

Obornella thompsonorum new species at the aperture just be-

Figure 12.

low the slit, showing the transition from the simple prismatic

outer layer to the nacreous inner layer. The arrows indicate

areas with recrystalized shell material. Abbreviations: Nae, na-

cre; spr, simple prismatic crystals

The selenizone is situated below the ramp angle. Surface

sculpture of spiral cords and threads, commonly cancel-

late at intersection with collabral threads. The selenizone

is moderately broad, the labral slit short.

Remarks: The oldest member of this genus is Bathro-

tomaria paipotensis Griindel, 2001, from the Sinemurian

(Lower Jurassic) of northern Chile. The selenizone is

narrow for the genus and the spiral sculpture is faint

compared to ether species. During the Middle and Up-

per Jurassic the genus was diverse and widely distributed

from Peru (Cox, 1956) to Europe, India, and the Afro-

Arabian East coast (Cox, 1960, 1965; Howarth and Nor-

ris, L995; Das et al., 2005), and possibly also New Zea-

land (Gardner and Campbell, 1997), although the New

Zealand record was not figured and needs confirmation.

A number of species were reported from the Lower Cre-

taceous of the Tethyan realm, including the largest spe-

cies with 130 mm diameter (Das, 2002: "Kolltaann, 1982,

2002). From the Cenomanian (lower Upper Cretaceous)

Kiel and Bandel (2004) reported four species of Bathro-

tomaria trom an intertidal rocky shore setting in Ger-

many, the highest diversity at any Cieiacdous locality.

Further Upper Cretaceous records are few: the last

record is from the lower Maastrichtian of France (Koll-

mann and Odin, 2001).

Bathrotomaria annejoffeae new species

(Figures 13-16)

Description: Shell (Figures 13-16) small for genus

(holotype maximum diameter 47.7 mm, minimum diam-

eter 43.5 mm, height 50.2 mm), with a tall, conical,

strongly gradate spire, consisting of 6+ teleoconch

suihole Base weakly but evenly convex, bo very narrow

umbilicus (Figure 15, uw). Spire angle 7 °. Spire very

slightly convex in profile. Suture w eakly ae join-

ing previous whorl at or above periphet ral bulge (Figure

14, pb). Protoconch and approximately first 2 ilesconch

whorls missing. Subsequent early whorls with straight to

weakly convex ramp between suture and ramp ‘angle

(Figure 14, ra) that becomes more inflated, convex on

body whorl. Shell surface with broad, uneven, undulating

rugae most evident near the suture. Axial sculpture of

evenly spaced, weak axial costae (about 120 on body

whorl) that produce weakly cancellate sculpture at inter-

sections with spiral cords. Spiral sculpture (Figure 16)

dominant, of 10-12 narrow, nearly abutting spiral cords

between suture and ramp angle (comprised of single,

smooth broad cord), 0-2 spiral cords between ramp

angle and selenizone, 6-7 along selenizone, 3-4 between

selenizone and peripheral bulge. Base with 36-38 spiral

cords that are twice as a as intervening spaces. Se-

lenizone (Figure 16, sz) broad, weakly convex, nearly

abutting ramp angle, me slightly more than half the

distance between ramp angle and peripheral bulge. Ap-

erture elongate, roughly pentagone al ieee axis near an per-

pendicular to the coiling axis. Outer Tip broken. Slit

broad (~3.5 mm), exte nding posteriorly 117° from the

end of the suture. Lip thick along columellar region, with

M. G. Harasewych and S. Kiel, 2007

Page 83

Figures 13-16. Bathrotomaria annejoffeae new species. 13. Apical, 14. apertural, and 15. basal views of the holotype, USNM

534483. 16. Details of sculpture between suture and periphery on last three dorsal whorls of the holotype. Abbreviations: p,

periphery: pb, peripheral bulge: pe, parietal callus; ra, ramp angle; s, suture; sl, posterior limit of slit; sz, selenizone; u, umbilicus.

narrow, weakly reflected parietal fold that forms a nar-

row parietal callus that partially occludes the umbilicus.

Type Locality: Zakaraha, near Toliara, southwesterm

Madagascar, 6-7 m below surface on plateau cut by river.

Type Material: Holotype, USNM 534483, from the

type locality.

Age: Oxfordian (Upper Jurassic).

Etymology: This new species honors Anne Joffe in

recognition of her many contributions and long service to

the field of malacology, particularly to the American Ma-

lacological Union (now American Malacological Society)

and most recently to The Bailey-Matthews Shell Mu-

seum in Sanibel, Florida.

Remarks: Bathrotomaria was among the most wide-

spread and diverse of the pleurotomariid genera

throughout the Middle and Upper Jurassic and Creta-

ceous. This is particularly true of the Jurassic fauna of the

Page 84

THE NAUTILUS, Vol. 121, No. 2

Kutch region, from which ten species have been re-

ported ( (Maithani, 1967; Jaitly et al., 2000; Das et al.,

2005), of these, five are of Oxfordian age. From its con-

temporary congeners within the Indo- Madagascan prov-

ince, Bathrotomaria annejoffeae is readily distinguished

from B. tewarii (Maithani, 1967), B. buddhai Das et al.,

2005, B. prasantai Das et al., 2005, and B. dhosaensis

Das et al., 2005 in having a proportionally taller, more

gradate spire and a narrower umbilicus. In shell profile,

B. annejoffeae more closely resembles specimens of B.

reticulata (Sowerby, 1821) (Das et al., 2005: figs. 4, A-G)

and B. siebereri (Jaitly et al., 2000:pl. 3, figs. 2-3) both

from the Bathonian of Kutsch, and the ae B.

millepunctata ( (Eudes- ree tauele: 1849) (Jaitly et

al., 2000: pl. 3, fig. 4), but lacks granular whorl angula-

tions of B. sie Die and B. reticulate: | (see Das et al.,

2005:334). Das et al. (2005: 334) also noted that the

Kutch specimens of B. reticulata lack an umbilicus

(present in B. annejoffeae), whereas the suture of B. sie-

bereri is deeply canaliculated, unlike that of B. annejof-

feae. Bathrotomaria annejoffeae can also be recognized

on the basis of the columellar portion of the aperture

being long, straight, and nearly co-axial.

Bathrotomaria bedetteae new species.

(Figures 17-20)

Description: Shell (Figures 17-20) small for genus

(holotype maximum diameter 65.0 mm, minimum diam-

eter 58.8 mm, height 47.5 mm), with a short, broad,

weakly gradate spire, consisting of 5+ teleoconch whorls.

Base weakly but evenly convex, lacking an umbilicus

(Figure 19). Spire angle 92°. Spire w veakly convex in pro-

file. Suture weakly adpressed, joining previous whorl at

or above peripher ral bulge (Figure 1S, pb). Protoconch

and approximately first 3 icleoeonch whorls missing.

Subsequent early whorls gradate, with straight ramp he

tween suture and ramp angle (Figure 1S, ra) that be-

comes more inflated, convex with increasing whorl num-

ber. Axial sculpture of broad, low, closely spaced, axial

costae (about 56 on body whorl) that produce a coarsely

cancellate sculpture at intersections of spiral cords, in-

cluding weakly beaded ramp angle and peripheral bulge.

Spiral sculpture codominant, of 3-4 low, broad, ards

i tween suture and ramp angle, 0-1 spiral cords be-

tween ramp angle and selenizone, 0-1 between seleni-

zone and peripheral bulge. Base with 17-19 finer spiral

cords that are 24 times as broad as intervening spaces.

Selenizone (Figure 20, sz) broad, weakly convex, nearly

abutting peripheral bulge, spanning slightly more than

half the eae between ramp angle and peripheral

bulge. Surface without spiral cords, sculpture limited to

strong lunulae. Aperture elongate, weakly pentagonal,

long axis deflected from the coiling axis by 103°. Outer

lip damaged, slit morphology not known. Inner lip thick-

est along columellar region.

Type Locality: Zakaraha, near Toliara, southwestern

Madagascar, 6-7 m below surface on plateau cut by river

Type Material:

type locality.

Holotype, USNM 534454, from the

Age: Oxtordian (Upper Jurassic).

Etymology: This new species honors the late Barbara

A. Bedette, whose 52 years of service to molluscan pa-

leontology at the National Museum of Natural History

has benefited a multitude of researchers.

Remarks: Bathrotomaria bedetteae may be readily

distinguished from B. annejoffeae, with which it co-

occurs, on the basis of its lower, broader shell, with a

weaker, more rounded ramp angle, by its coarser and

more prominent cancellate sculpture that extends onto

the ramp angle and peripheral bulge, and by the absence

of an ainbilieus, Bathrotomaria bedetteae is similar in

profile to B. tewarii, B. prasantai, and B. dhosaensis, all

from contemporary strata in Kutch, but differs in having

a more gradate spire and a pronounced peripheral band,

and in lacking an umbilicus.

Genus Leptomaria E. Eudes-Deslongchamps, 1864

Type Species: Plewrotomaria amoena J. A. Eudes-

Deslongchamps, 1849. (By original designation). Bajo-

cian (Middle Jurassic) of France.

Diagnosis: Species of Leptomaria can be recognized

on the basis of their large, turbiniform shells with low to

moderately high spires ne weakly to strongly rounded

whorls. The naibilieus may range from broz a to entirely

absent. The whorl profile is rounded, lacking an angulate

shoulder. The selenizone is situated at mid-w haul Sur-

face sculpture consists primarily of narrow spiral threads

with finer axial threads forming weakly cancellate sculp-

ture in some species,

Remarks: = Leptomaria has been reported from strata

ranging in age from Bajocian (Middle Jurassic, Knight et

al., 1960) to Selandian (Paleocene, Kollmann and “Peel,

1983). Hickman (1976) suggested that even some

Eocene species may be included. Not surprisingly, the

genus had a cosmopolitan distribution, with diverse fau-

nas ranging from England (Cox, 1960) to New Zealand

(Hudson, 2003). Leptomaria has been previously re-

ported from Cretaceous deposits of NW Madagascar

(Delpey, 1948; Collignon, 1949; Kiel, 2006). It is repre-

sented in the Jurassic fauna of Kutch (Jaitly et al., 2000;

Das et al., 2005) but is not as diverse as the genera

Bathrotomaria or Plewrotomaria.

Leptomaria takahashii new species

(Figures 2]—24)

Description: Shell (Figures 21-24) moderately large

for the genus shelotype maximum diame ter $2.4 mim,

minimum diameter 67.23 mm, height 57.3 mm) low, tur-

biniform, consisting of approminalely 6 teleoconch

whorls. Base broadly and eu convex, with very nar-

row umbilicus (Figure 23, Spire angle LO9®. Spire

strongly convex in profile. ie (Figure 24, s) abutting,

M. G. Harasewych and S. Kiel, 2007

Page 85

2cm

Figures 17-20. Bathrotomaria bedetteae new species. 17. Apical, 18. apertural, and 19. basal views of the holotype, USNM

534484. 20. Details of sculpture between suture and periphery on last three dorsal whorls of the holotype. Abbreviations: p,

periphery: pb, peripheral bulge: ra, ramp angle; s, suture; sl, posterior limit of slit; sz, selenizone.

joining previous whorl just below selenizone (Figure 24,

sz) in early whorls, at or below periphery (Figure 29, 24,

p) in later whorls. Protoconch and part of first teleoconch

whorl eroded. ae teleoconch whorls (whorls 2—4)

evenly rounded, later whorls (whorls 5-6) becoming

more gradate, but evenly rounded, lacking a angular

shoulder. Axial sculpture of numerous fine srowth striae

that produce a strongly reticulate pattern most evident

between adjacent cords in region between suture and

selenizone of early whorls, and broader low, axial costae

that form a weakly cancellate pattern at intersections

with the spiral cords. Spiral sculpture dominant, with 10

strong, closely spaced cords between suture and seleni-

zone, 7-9 cords between selenizone and periphery, and

29-32 cords along base. Spiral cords may become

broader and less disGnct with increasing whorl number.

Selenizone (Figure 24, sz) narrow, convex, situated just

above periphery, with a single, median spiral cord

present in early whorls, abpenk in later whorls, and nu-

merous strong to weak lunulae throughout its length.

Aperture ev enly ovate, long axis forming an angle of 103°

with pe axis. Outer lip smooth, portion below slit

offset from portion above slit by 30°. Slit narrow (~3.3

mim), extending posteriorly 51° from end of suture. Lip

Page 86

THE NAUTILUS, Vol. 121, No. 2

2cm

Figures 21-24. Leptomaria takahashii new species. 21. Apical, 22. apertural, and 23. basal views of the holotype, USNM 534485.

24. Details of sculpture between suture and periphery on last three dorsal whorls of the holotype. Abbreviations: p, periphery; pe,

parietal callus; s, suture; sl, posterior limit of slit; sz, selenizone; u, umbilicus.

thickest in columellar and basal region, forming narrow

parietal callus that partly overlaps the umbilicus.

Type Locality: Zakaraha, near Toliara, southwestern

Madagascar, 6—7 m below surface on plateau cut by river.

Type Material: Holotype, USNM 534485, from the

type locality.

Age: Oxfordian (Upper Jurassic).

Etymology: This new species honors Mr. Chris Ta-

kahashi of Honolulu, Hawaii, in recognition of his many

contributions to the study of living and fossil mollusks.

Remarks: = Leptomaria daityai Das et al., 2005 (Cal-

lovian to Oxfordian of Kutch, India) occurs in coeval de-

posits, and most closely resembles L. takahashii in terms

of size and general profile, but L. takahashii has a shorter

spire, more rounded whorls, and has a very narrow rather

than a wide umbilicus. Although the genus Leptomaria is

well represented in the Jurassic fauna of the Indo-

Madagascan region, most published records are either

from Bathonian and Callovian strata of Kutch, or from

Cretaceous deposits of Madagascar. Older taxa, includ-

ing L. fraga (J. A. Eudes-Deslongchamps, 1S49), L. sim-

plex Jaitly et al., 2000, L. pseudoumbilicata Jaitly et al.,

M. G. Harasewych and S. Kiel, 2007

Page 87

2000 (transferred to Bathrotomaria by Das et al., 2005:

340) are easily distinguished as they are smaller, and have

a tall, conical whorl ‘profile. Although the Bathonian L.

asurai Das et al., 2005, is of comparable size, it differs in

also having a tall conical profile and lacks an umbilicus.

DISCUSSION

The species most similar to those described herein occur

in contemporaneous fossil deposits of northwestern In-

dia. This is not surprising considering that the Madagas-

car species lived near the southern tip of a long and

narrow embayment between East and West Gondwana,

while northwestern Indian species lived at the northern

tip of this | ment, where it opened to the Tethys

Ocean (Figure 2). Although Recent pleurotomariids are

iran to a substrates along the outermost conti-

nental shelf and upper continental slope, Mesozoic pleu-

rotomariids inhabited sublittoral depths along the conti-

nental shelf. Studies of Upper Jurassic phytogeography

have indicated that southern Madagascar was part of a

warm temperate biome, while northern Madagascar and

northwestern India were part of a warmer, subtropical

biome (Rees et al., 2000: fig. 7C). B

stable isotope compositions of the ammonite Per risphinc-

tes (Dichotomoceras), Lécuyer and Bucher (2006: 7, fig.

3) reported seawater temperatures ranging from 21.5°C

to 24.1°C in the Morondava Basin of southwestern

Madagascar, during the Oxfordian stage of the Upper

Jurassic.

Extensive studies of Jurassic aaa faunas

within this Indo-Madagascan Province (e.g., Cox, 1965;

Jaitly et al., 2000; Das, 2005) have documented the pres-

ence of the genera Bathrotomaria, Leptomaria,

Obornella, Plewrotomaria, Anodomaria, and Pyrgotro-

chus. the first three by far the most diverse. As additional

specimens from the Jurassic of Madagascar become

available, it is likely that the generic composition and

diversity of this fauna will mirror that of the Kutch region

of northwestern India.

ACKNOWLEDGMENTS

We are grateful to Chris Takahashi for generously mak-

ing the specimens available for study, anil a llowing the

type material to be deposited in the collections of the

National Museum of Natural History, Smithsonian Insti-

tution. We thank K. Bandel, J. Hartmann, and W.

Weitschat, (all Hamburg) and H. Keupp (Berlin), for

their help regarding the focalities and their age; and T. A.

White for providing the Jurassic paleo-map. A. Oleinik

and an anonymous reviewer are acknowledged for their

reviews.

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Appendix 1. Supraspecific taxa included in the family

Pleurotomariidae, listed in the order in which they were

proposed. Their type species are provided, as are known

stratigraphic ranges, and geographical distributions.

Pleurotomaria Defrance, 1826 — Type species, Trochus angli-

cus J. Sowerby, 1818. Middle Triassic (Anisian; range ex-

tended by Begg and Grant-Mackie, 2003: 259) to Lower

Cretaceous (Aptiz in). C Josmopolitan.

Leptomaria E. Fudes- De slongch: amps, LS64 — Type ators

Pleurotomaria amoena |. A. Eudes-Deslongchamps, 1549.

Middle Jurassic (Bajocian) to Paleocene (Danian). Cosmo-

politan,

Talantodiscus Vischer, 1885 — Type species, Pleurotomaria

mirabilis Deslongchamps, 1848. Middle Triassic

(Kaihikuan ; range extended by Begg and Grant-Mackie,

2003: 259) to Middle Jurassic (Bi york an). E surope and New

Zealand. Knight et al. (1960; 1214) included this genus in

the family Porcelliidae Broili, 1924. Subsequent authors

(e.g., Szabo, 1980: fig. 3; Begg and Grant-Mackie, 2003:

299) included this genus within Pleurotomariidae.

Pyrgotrochus P. F isolien. 1885 — Type species, Pleurotomaria

bitorquata |. A. Eudes-Deslongchamps, 1S49. Lower

M. G. Harasewych and S. Kiel, 2007

Jurassic to Upper Cretaceous (Cenomanian). Cosmopoli-

tan.

Perotrochus P. Fischer, 1885 — Type species, Plewrotomaria

quoyana Fischer and Bernardi, 1856. Oligocene to Re-

cent. Cosmopolitan.

Chelotia Bayle in P. Fischer, 1885 — Type species, Pleuroto-

maria concava Deshayes, 1832. Eocene. Europe.

Entemnotrochus P. Fischer, 1885 — Type species, Plewroto-

maria adansoniana Crosse and Fischer, 1861. Eocene to

Recent. ree

Stuorella Kittl, 1S91 — Type species, Trochus subconcavus

Miinster, 1541. Middle” Triassic (Ladinian) to Upper Cre-

taceous (Campanian; range extended by Kiel and Bandel,

2000). Europe.

Mikadotrochus Lindholm, 1927 — Type species, Plewrotomaria

beyrichii Hilgendort, 1877. Western Pacific, Pliocene to

Recent.

Bathrotomaria Cox, 1956 — Type species, Trochus reticulatus J.

Sowerby, 1821, by original designation. Lower Jurassic to

Upper Cretaceous (Senonian). Cosmopolitan.

Conotomaria Cox, 1959 — Type species, Pleurotomaria mail-

leana VOrbigny, 1843. Middle Jurassic (Bajocian) to Pa-

leocene (Danian; range extended by Pacaud, 2004). Cos-

mopolitan.

Obornella Cox, 1959 — Type species, Plewrotomaria plicopunc-

tata J. A. Eudes-Deslongchamps, 1549. Lower Jurassic

(Toarcian) to Upper Jurassic (Oxfordian) Cosmopolitan.

Anodotomaria Szab6, 1980 [described as a subgenus of Pleu-

rotomaria| — Type species Pleurotomaria scacchi Gem-

mellaro, 1874. Lower Jurassic (Pliensbachian) to Middle

Page 89

Jurassic (Callovian; range extended by Jaitly et al., 2000:

36). Europe to noehanestert India.

Cyclostomaria Szab6, 1980 — Type species, Plewrotomaria

suessi Hérnes, 1853. Lower Jurassic (Pliensbachian) to

Middle Jurassic. Europe.

Ornatospira Pan, 1982 — Type species Ornatospira mirifira

Pan, 1982. Triassic, China.

Laevitomaria Conti and Szabo, 1987 — Type species, Pyrgotro-

chus? problematicus Szab6, 1980. Lower Jurassic (Pliens-

bachian) to Middle Jurassic (Bajocian) Europe. Question-

ably included in Pleurotomariidae by the authors.

Indomaria Das, 2002 [described as a subgenus of Pleuroto-

maria] — Type species, Pleurotomaria intdontonta) umien-

sis Das, 2002. Lower Cretaceous (Berriasian). Northwest-

ern India.

Bayerotrochus Harasewych, 2002 — Type species, Pleuroto-

maria midas Bayer, 1964. Miocene to Recent. Cosmopoli-

tan.

Murihikua Begg and Grant-Mackie, 2003 — Type species,

Murihikua tuhawaiki Begg and Grant-Mackie, 2003,

Middle Triassic (Aniasian) to Upper Triassic (Norian).

New Zealand.

Tahua Begg and Grant-Mackie, 2003 — Type species, Talia

waipiro Begg and Grant-Mackie, 2003 Middle Triassic

(Aniasian) to Late Triassic (Norian). New Zealand.

Mamoeatomaria Begg and Grant-Mackie, 2006 (new name

for Mamoea Begg and Grant-Mackie, 2003) — Type

species, Mamoea wairakiensis Begg and Grant-Mackie,

2003. Middle Triassic (Aniasian to Ladinian). New Zea-

land.

THE NAUTILUS 121(2):90-94, 2007

Page 90

Sassia melpangi, a new ranellid species (Gastropoda: Tonnoidea)

from the Central Pacific

Alan G. Beu

GNS Science

P.O. Box 30368

Lower Hutt 5040

M. G. Harasewych

Department of Invertebrate Zoology

National Museum of Natural History

Smithsonian Institution

P.O. Box 37012

Washington, DC 20013-

[email protected]

7012 USA a. [email protected]

NEW ZEALAND

ABSTRACT

Sassia melpangi is described from bathyal depths off Oahu,

Hawaii. This new species is most similar to S. nassariformis

(Sowerby, 1902) from comparable depths off southeastern Af-

rica, and to a lesser extent to S. remensa (Iredale, 1936) from

the western Pacific. Sassia melpangi may be distinguished from

all other Sassia on the basis of its broad, evenly rounded whorls,

absence of a distinct shoulder on the varices, numerous axial

ribs and spiral cords that produce an evenly reticulate surface

sculpture, a broadly ovate aperture with distinctive inductura

and strongly pigmented pattern along the edge of the outer lip.

INTRODUCTION

Sassia, the oldest of the ranellid genera, is represented in

the Upper Cretaceous deposits of the United States, Eu-

rope, and northern Africa. This genus became cosmo-

politan by the Eocene and has Been considered to be a

basal group that gave rise to all other Ranellidae (Beu,

1998a: 800). In the Recent fauna, Sassia appears to have

dispersed via a Tethyan distribution route, ranging from

South Africa [Sassia nassariformis (G. B. Sowerby I,

1902)| and Japan [Sassia semitorta (Kuroda and Habe in

Habe, 1961)| to the South Atlantic [Sassia philomelae

(Watson, 1SS0)|. Greatest diversity occurs in the Indo-

Pacific, with most species inhabiting outer shelf to upper

slope de pths (100 to 600 m). The Indo-West Pacific Sas-

sia were revised by Beu (1998b: 137), who distinguished

is “Sassia sp. nov.?” a distinctive specimen from Raeva-

vae, French Polynesia. He mentioned examining photo-

or iphs of additional specimens from Guam and Hawaii,

but deferred from naming it formally until more speci-

mens became available.

Through the kindness of Mr. Chris Takahashi, five

additional specimens, all taken in traps off Oahu, Hawaii,

were made available for study. Based in this new mate-

rial, the taxon Sassia melpangi is described as a new

species

Abbreviations and acronyms used in the text are: dd,

dead-collected shell: MNHN, Muséum national

(Histoire naturelle, Paris; USNM, National Museum of

Natural History, Smithsonian Institution, Washington,

DG:

SYSTEMATICS

Superfamily Tonnoidea Suter, 1913

Family Ranellidae Gray, 1854

Subfamily Cymatiinae Tredale, 1913

Genus Sassia Bellardi, 1873

Type Species: § Triton apenninicum Sassi, 1527, from

the Miocene and Pliocene of Europe (by subsequent

designation, Cossmann, 1903: 93). For an extensive syn-

onymy, see Beu, 199Sb: 139.

Sassia melpangi new species

(Figures 1-13)

Sassia sp. nov.? Beu, 1998: 141-142, fig. 43, 1.

Diagnosis: Sassia with up to 8 broad, evenly rounded

Ww Horie: without distinct shoulder on varices; sculpture of

numerous axial ribs and spiral cords that produce an

evenly reticulate surface sculpture; a broadly ovate ap-

erture forming a distinctive inductura with ventrally re-

flected edges and strongly pigmented pattern along the

edge of the outer lip.

Description: Shell (Figures 1, 3-6, 11-13) large for

genus (reaching 51.7 mm; Table 1), fusiform, with tall,

conical spire, large aperture, oval inductura with re-

flected edge, and short, open, axially oriented siphonal

canal. Protoconch (Figures 12-13), tall- conical, increas-

ing in diameter from 185 jum to 1.67 mm in 2.75 evenly

rounded, glossy whorls. First half whorl smooth, followed

by onset of shi arp, narrow axial cords (26-29 per whorl)

and 34 narrow, weaker, spiral cords. Transition to te-

leoconch abrupt, marked by change in color from amber

M. G. Harasewych and A. G. Beu, 2007 Page 9]

Figures 1-6. Sassia me [pangi new species 1. Holotype USNM 1099759, apertural, right lateral, and dorsal views. 2. SEMs of the

inner and outer surfaces of the operculum of the holotype 3. Apertural view of paratype |. 4. \pertural view of paratype 2. 5.

] eft lateral views of Parat pe 3. USNM 1099760, showing adherent poly haete tubes (wt). 6. Ape rtural view of Paratype

1 cm (applies to all shells.) Abbreviation: wt, worm tube

Page 92

THE NAUTILUS, Vol. 121, No. 2

Table 1. Measurements of the holotype and four paratypes of

Sassia melpangi. Linear measurements in mm.

HT PTI PT2 PT3 PYr4

Shell length 41.1 37.6 51.1 5L.7 50.3

Aperture length 161 149 198 20.5 19.3

Siphonal canal length 7.5 7A 9.8 112 9.5

No. whorls, protoconch 3.3 3.3 3.2 3.2 3.2

No. whorls, teleoconch 6.5 6.5 13 7.0 7.2

No. cords on penultimate

whorl D 4 5 5 5

No. cords on last whorl 8 i) 9 8 i)

No. cords on siphonal

canal 6 4 5 6 7

No. axial ribs on last

whorl 24 20 29 26 PA

No. axial ribs between

varices 1S 15 2] 17 13

Spire angle 50.0° 48.5° 448° 45.8° 43.0°

to tan, loss of surface gloss, and coarsening of axial and

spiral sculpture. Teleoconch of up to S convex, evenly

rounded whorls. Suture adpressed. Axial sculpture of i

29 weakly opisthocline to weakly prosocline ribs per

whorl, as broad as interspaces, forming reticulate sculp-

ture of hemisphe srical beads at intersections with strong

spiral cords (3 on first whorl, 4-5 on penultimate worl.

8-9 on last whorl, 4-7 on siphonal canal). Cords com-

prised of 3-5 broad fused threads with 3-9 finer threads

between adjacent cords. Varices broad, strongly raised,

begin after about 0.5 5 whorl and recur every 230-240°

thereafter. Plane of inductura tangential to previous

whorl, thus, varices form an angle of up to 10° with axial

rib, intersecting at base of siphonal canal. Aperture large

(0.38-0.40 shell length), broad (0.5-0.6 of aperture

length), oval, major axis deflected from shell coiling axis

by 20-23°. Outer lip reflected, forming rim of ‘inca,

thickened, with 7 strong teeth that do not extend beyond

the varix. Most adapical tooth largest, opposed to strong

parietal tooth. Flaring edge of outer lip with rectangles of

dark reddish brown pigment along its outer margin that

are aligned with spiral cords. A broad, oval inductura

with reflected edges extends over the parietal area. Pa-

rietal region with single, strong tooth that does not ex-

tend into the aperture and one or two weak folds that

overlay spiral cords of previous whorl. Columella with

multiple weak folds of varying lengths and angles, some

e nding before reac ching the ingucean a, others chose origi-

nating on the inductura. Columellar fold at junction “of

siphonal canal most pronounced. Siphonal canal about

half as long as aperture, axial, dorsally deflected, nar-

Eowy open, with proximal half covered by inductura.

3ase color cream to light tan, with axial bands of darker

red lish brown, 3 bands between adjacent varices on

early whorls, increasing to 6 bands between varices. Va-

rices pigmente -d with reddish brown, darker on dorsal,

lighter on ventral region, interrupted by slightly to much

lig ghte r bands along spi al cords. Interior of she ll nacre-

ous white, base color visible in thinner areas. Perios-

tracum (Figure 11), thin, brown, finely lamellose, hir-

sute, with hairs aligned along spiral threads and growth

lines. Periostracum best preserved along suture in most

specimens. Operculum (Figure 2) large (> 0.8 aperture

length), ovate, thin, corneous, with terminal nucleus.

Radula (Figures 7-9) short (0.33 aperture length) con-

sisting of 45-51 rows of teeth (7 per row). Rachidian

teeth broad, with wide, curved basal plate, with strong

central cusp flanked by 5-8 short, conical denticles. Lat-

eral teeth broad, with stout bases, 6-8 denticles along

ventral edge. Marginal teeth scythe-like, with eaieath

edges. Jaws (Figure 10) paired, narrow dorsally, ex-

panded ventro-laterally, with fringed edges.

Type Locality: Oahu, off the north shore district of

Haleiwa, muddy sand, in red shrimp traps set at 300-

390 m.

Type Material: Holotype, USNM_ 1099759; Paratype

3, USNM 1099760, Paratypes 1, 2, 4, Takahashi collec-

tion. All from the type locality.

Additional Material Examined: AUSTRAL ISLANDS:

MNHN, Raevavae, 23°50.54 S, 147°42.73° W, in 400 m.

BENTHAUS: stn DW 1884, Iles are 570-620 m (1

dd): sth DW 1885, as last, 700—SO0 m (1 dd); stn DW

1897, ouest de Rapa, 480-700 m (2 dd); stn DW 1899, as

last, 342-600 m (1 dd); stn DW 1903, Banc NE Rapa,

400-600 m (1 dd); stn DW 1923, Recif Nielsen, 360-840

m (1 dd): sth DW 1924, as last, 340-800 m (1 dd): stn

DW 1925, as last, 560-790 m (1 dd): sth DW 1929, Banc

Président Thiers, 350-370 m (1 dd); stn DW 1932, as

last, 500-600 m (1 dd): stn DW 1933, as last, 500-859 m

(2 dd); stn DW 1940, N de Rae vavae, LOO—460 m (3 dd);

stn DW 1943, as last, 950 m (2 dd): sth DW 1945, Banc

Lotus, 120-500 m (2 dd); stn DW 1951, as 206-450

m (1 dd): stn DW 1957, Tubuai, 55S—1000 m (2 dd): stn

DW 1961, as last, 470-800 m (3 dd): stn DW 1973, Banc

Arago, 300-350 m (2 dd); stn DW 1974, as last, 450-615

m (2 dd): stn DW 1992, Rurutu: Mont de Lotus, 442-444

m (1 lv): stns 1997-2001, Rurutu, 200—LOOO m (2 dd); stn

DW 1998, cote N de Rurutu, 250-302 m (1 dd?); stn

DW 1999, as last, 270-500 m (4 dd); stn DW 2000, cote

N de Rurutu, 270-480 m (1 dd); stn DW 2001, port de

Rurutu, 200-550 m (1 Iv?); stn DW 2006, cote E de

Rurutu, 35-450 m (1 dd); stn CAS 2008, cote E de Ru-

rutu, 280-300 m (2 dd); sth DW 2018, Rimatara, 770—

771 m (1 dd); stn DW 2021, Rimatara, 1200-1226 m

(1 dd).

Distribution: Sassia melpangi is broadly distributed

throughout the central West Pacific, from Guam to Ha-

waii and Raevavae in the Austral Islands, but does not

occur in the Marquesas. [t inhabits upper bathyal depths,

with live specimens collected between 200 and 550 m.

Etymology: This species is named in honor of Mr.

Melvin Pang, of Oahu, Hawaii, who collected the type

series.

Comparative Remarks: ~The new species Sassia mel-

pangi is readily distinguished from the western Pacific S.

M. G. Harasewych and A. G. Beu, 2007 Page 93

Figures 7-13. Sassid ™meé lpangi new species

of radula near distal end. lateral teeth spread to reveal rac hidian teeth. 9. Right lateral view or radular ribbon. Scale bar

s 7-9). 10. Jaw of holotype, with detail of edge. 11. Periostracum of paratype 1, at midpoint of final vari

13. Lateral views of protoconch of paratype 1. Scale bar | mm (applies to Figures 12, 13

7-9. Radula of the holotype. 7. Dorsal view of radula near mid-length. 8. Dorsal vic

LOO jxn

12. Apical

applies to Figur

Page 94

THE NAUTILUS, Vol. 121, No. 2

remensa and 8S, semitorta in having: more evenly rounded

whorls that lack a pronounced shoulder the presence of

more numerous, evenly spaced axial ribs that form a

reticulate surface sculpture; a more evenly ovate aper-

ture surrounded by an inductura with ventrally reflected

edges; as well as a strong pigmentation pattern along the

outer lip. Sassia melpangi most closely resembles S. nas-

sariformis from southeastern Africa, with which it shares

the rounded whorl profile and even cancellate sculpture.

Sassia nassariformis can be differentiated by its shoul-

dered varices, smaller aperture, weaker pigmentation

along the outer lip, and by its thicker inductura, which is

more triangular than ovate, and which is not reflected

along it edges.

Sassia melpangi has a broad range throughout the

tropical Pacific. While the type lodalitay is off the Island of

Oahu, a number of specimens are known from French

Polynesia, and photographs of two specimens from

Guam, Marianas Islands have been examined. This spe-

cies inhabits upper bathyal depths (300-400 m). The

position of worm tubes (Figure 5, wt) on the ventral

surface of living specimen suggests that Sassia melpangi

is epifaunal atid inhabits hard “Sabsirates.

ACKNOWLEDGMENTS

We are grateful to Chris Takahashi, for bringing this

material to our attention, and for donating the holotype

and one agg to the National Museum of Natural

History. Richard Salisbury kindly made available

photogr ae of two specimens dredged off Guam, Mari-

anas Islands, and Professor Alison Kay provided photo-

graphs of additional specimens dredged in Hawaii.

LITERATURE CITED

Beu, A. G. 199Sa. Superfamily Tonnoidea. In: Beesley, P. L.,

G. J. B., Ross, and A. Wells (eds.) Mollusca: The Southern

Synthesis. Fauna of Australia 5. CSIRO Publishing, pp.

792-803.

Beu, A. G. 1998b. Indo-West Pacific Ranellidae, Bursidae and

Personiidae (Mollusca: € Gastropod: 1). A monograph of the

New Caledonian fauna and revisions of re lated taxa. Mem-

oires du muséum national dhistoire naturelle 178: 255 pp.

Cossmann, M. 1903. Essais de Paléoconchologie comparée.

Vol. 5. M. Cossmann, Paris, 215 pp.

Habe, T. 1961. Coloured Hlustrations of the Shells of Japan vol.

2. Hoikusha Publishing Co., Osaka, ix + 182 pp, appendix

42 pp.

Iredale, T. 1936. Australian molluscan notes. No.2. Records of

the Australian Museum 19: 267-340.

Sowerby, G. B. HI, 1902. Mollusca of South Africa. Marine

Investigations in South Africa 2: 93-100.

Watson, R. B. 1881. Mollusca of the HMS Challenger Expe-

dition, part 7. Families Pyramidellidae, Naticidae,

Cassidae, Tritonidae. Journal of the Linnean Society of

London 15: 245-274.

THE NAUTILUS 121(2):95-98, 2007

Page 95

A new species of Microcancilla (Gastropoda: Cancellariidae)

from the continental slope off northeastern Brazil

José Carlos N. de Barros

Departamento de Pesca e Aquicultura

Richard E. Petit

S06 St. Charles Road

Universidade Federal Rural de Pernambuco North Myrtle Beach, SC 29592 USA

Avenida Dom Manuel de Medeiros, S/N,

Dois Irmaos

52171-030, Recife, BRAZIL

[email protected]

ABSTRACT

Microcancilla jonasi new species is described from deep waters

off northeastern Brazil. The genus Microcancilla Dall, 1924,

had not previously been recorded in Brazilian waters. Speci-

mens were collected from sediment dredged as part of the

REVIZEE program (Assessment of the Sustainable Potential of

Living Resources in the Exclusive Economic Zone) during

prospecting work on the continental slope off the state of Per-

nambuco at depths of 425 to 690 meters. The present study

reveals that these specimens present affinities to the species

Microcancilla microscopica (Dall, 1889), which differs from the

new species, among other features, by having strong spiral

sculpture between sigmoid axial ribs. In addition, a lectotype

for Cancellaria microscopica Dall, 1889, is designated herein.

Additional keywords: Cancellaria, Neogastropoda, Pernam-

buco.

INTRODUCTION

In his work on the gastropods collected in the West In-

dies (1879-80) by the U.S. Coast Survey Steamer BLAKE,

Dall (1889a) described Cancellaria microscopica Dall,

1889, based entirely on empty shells. Dall later (1889b:

106) placed that species with a ‘toa in the genus Ad-

mete ie er in Moller, 1842. Still later, Dall erected the

cancellariid genus Microcancilla Dall, 1924. When intro-

ducing this genus, Dall did not give a genus description,

and listed only the single species Admete [sic] micro-

scopica Dall, 1889. Until now no congeners have been

reported.

Most species of Admete are found in shallow waters of

polar regions but a few occur in deepe r water in tem-

perate zones (Harasewych and Petit, 1986; Knudsen,

1964). However, all of the taxa described as Admete have

not been studied in detail and it is probable that some are

not properly placed in this genus (Bouchet and Warén,

1985).

A recent study of deep-water Cancellaridae from the

New Caledonia area (Bouchet and Petit, in preparation)

shows that the central Pacific fauna contains species

clearly attributable to Microcancilla and others that are

morphol ogically similar to Admete aethiopica Thiele,

1925, font Somalia [illustrated in Verhecken (1997: 312,

fig. 52)|. Intermediate between these seemingly dispar-

ate morphologies and having various features in common

with them are a series of species. Verhecken (1997),

figuring the Somalia specimen, placed it in the genus

Adee with a query but offered no other possible plac e-

ment.

It is not contended here that Microcancilla jonasi new

species, “Admete” aethiopica Thiele, and Microcancilla

microscopica (Dall) are truly congeneric species. How-

ever, Microcancilla is considered at the moment the best

available placement within the existing genera of the

Cancellarioidea and such placement is provisional. Until

the small deep-water species of Cancellariidae are better

understood and the validity and limits of other available

genus group taxa are established, we do not wish to in-

troduce a new genus-group taxon. This problem with the

available genera for small cancellariids was succinctly

pointed out by Maxwell (1992: 167). Microc ancilla jonasi

new species is considered endemic to the continental

slope off the state of Pemambuco, Brazil.

MATERIALS AND METHODS

All specimens examined were obtained in 1999-2000,

during oceanographic prospecting work on the continen-

tal slope off the state of Pernambuco, Brazil. No live

specimens were collected. Shells were mounted on

specimen stubs and exmained and photographed under a

Jeol JSM 6360 Scanning Electron Microscope, at the

“Instituto Tecnolégico de Pernambuco (ITEP)”. Shells

were measured using a stereo microscope with eyepiece

micrometer. The type material was deposited at the

Academy Natural of Sciences, Philadelphia, USA

(ANSP): Museu Nacional, Rio de Janeiro, Brazil

(MNRJ); Museu de Zoologia da Universidade de Sao

Paulo, Brazil (MZUSP):; Museu Oceanogratico do Rio

Grande, Rio Grande, Brazil (MORG).

Page 96

THE NAUTILUS, Vol. 121, No. 2

SYSTEMATICS

Family Cancellariidae Forbes and Hanley, 1851

Genus Microcancilla Dall, 1924

Type Species: © Admete microscopica (Dall, 1889) [=

Cancellaria microscopica Dall, 1889a] by original desig-

nation. Recent, Caribbean.

Microcancilla jonasi new species

(Figures 1-8)

Description: — Shell conical, small, fragile, translucent,

whitish, short spire. Protoconch smooth, globose, pau-

cispiral with 1.5 whorls, terminating with the emergence

Figures 1-8.

Scanning electron micrographs of Microcancilla jonasi new species (all deposited in MOR¢

of the axial sculpture of the teleoconch (Figures 7-8).

Teleoconch with 2.5 gee convex whorls. Suture well-

marked, with a small, flattened subsutural region orna-

mented by the adapical portion of the axial “ibs. On the

margin of this region, there is a strongly nodular shoul-

der, nodule 4S coinciding with axial ribs below, resembling

a small crown. Below the shoulder, there is a second

weaker spiral cord, with nodules somewhat smaller than

those of the crown on margin of a small, concave, spiral

depression (Figure 6). Body whorl developed, very wide,

with about 69% of the total size of the shell, sculptured

with 15 to 18 rounded, regularly spaced ribs, which dis-

appear toward the base. Interspaces ornamented by ob-

scure threads. Base imperforate, strongly conical, with

+): L. Holotype, apertural

iew, leneth fmm; 2. Paratype, apertural view, length: 3.28 mm; 3. Paratype, apertural view, length = 3.12 mm; 4. Paratype

ipertural view, length = 2.55 mm; 5-8. Paratype, length: 3.60 mm Apertural view. 6. a view, showing strongly nodular

shoulder and weak spiral cords (Figure 5). 7-8. Protoconch (shell in Figure 5). Scale bars: Figure 6 = 500 jm; Figure 7 = 200 jzm;

Figure 8 = LOOwm

J.C. N. de Barros and R. E. Petit, 2007

Page 97

convex contour, ommamented by 3 to 5 weak spiral cords

that emerge from the interior of aperture. Aperture el-

liptic. Outer lip fragile and smooth inside. Inner lip

cad y reflected, wide parietal region, without callus,

median portion of the columella with reflected appear-

ance, thickened upon the umbilical wall with no columel-

lar folds. Final anterior portion of the columella inclined

to the left of the shell axis.

Type Material: —© Holotype, MORG 50.716, length = 4

mm; 4 paratypes, MORG 50.718 (Figures 2-8); 3

paratypes, MZUSP $1755; 3 Paratypes ANSP 413549, 1

paratype, MNRJ 1OS39, Pernambuco, Brazil, D-4,

08°42.1" S, 34°44.1' W, 425 m, muddy bottom, 25 Mar.

2000; 2 paratypes, MNRJ 10835; 3 paratypes, MZSP

$1756; 1 paratype MORG 50.717 (Pernambuco, Brazil,

D-11, 08°46.5' S, 34°44.5' W, 690 m, muddy bottom, 18

Sep. 2000).

Type Locality:

Pernambuco, northeastern Brazil, D-4, 08°42.1' S,

34°44." W, 425 m, muddy bottom, 25 Mar. 2000.

Etymology: The species is named after Mr. Rainer

Jonas, scientific director of the Gesellschaft fiir Biotech-

nologische Forschung (GBF), Germany, for his assis-

tance in obtaining literature and his constant support

during the identification work of gastropods from the

contnenals slope of Brazil.

Geographic Distribution: — Endemic to the Conti-

nental Slope of the State of Pernambuco, northeastern

Brazil.

Habitat: © Muddy substrate, 425 to 690 m.

Discussion: — Microcancilla jonasi new species studied

here is distinguished from Microcancilla mic roscopica

and ?Admete aethiopica based on the following charac-

teristics: (1) sculpture of the body whorl: M. micro-

scopica has a strong spiral sculpture, with subequal

rather coarse threads, forming a reticulum with the axial

spirals. 2A. aethiopica presents a wide body whorl, with a

strong axial sculpture and smooth spiral bands se parated

by narrow grooves disappearing near the base, M. jonasi

has strong axial ribs, sinuous and rounded, and an ob-

scure spiral ornamentation with no reticulation; (2) Spi-

ral ornamentation: M. microscopica has threads that al-

ternate between strong and weak, crossed by raised

growth threads, ?A. aethiopica has strong axial ribs

crossed by weaker spiral threads, M. jonasi only presents

raised axial ribs: (3) Shoulder: on M. microscopica, the

shoulder is obsolete and the subsutural platform be-

comes rounded on the body whorl, ?A. aethiopica pre-

sents an angular shoulder, strongly nodular, and a nar-

row, flat subsutural platform, M. jonasi has a small, flat

subsutural platform, bordered by a nodular shoulder

with a lower depression and followed by a second sub-

shoulder cord that is more weakly nodular: (4) Umbili-

cus: M. microscopica has a distinct, moderate umbilicus

with no bounding carina or siphonal fasciole, 2A. aethi-

opica and M. jonasi do not possess an umbilicus; (5

Continental Slope from the State of

Aperture: M. microscopica has an aperture that is

rounded behind and strongly angular in front, ?A. aethi-

opica has an oval aperture, slightly square-cut adapically,

M. jonasi has an elliptic aperture, weakly constricted be-

hind: (6) Inner lip: M. microscopica has a moderately

callous inner lip with one extremely faint fold about the

middle, PA. aethiopica has an inner lip with two very

weak folds near the halfway height, M. jonasi has a re-

flected inner lip, wide parietal region, with no folds: (7

Columella: In M. microscopica and PA. aethiopica, i

columella is straight, parallel to shell axis; in M. jonasi, it

is inclined to the left of the shell axis.

Only the type species has been allocated in Microcan-

cilla since the genus was ae The specimen fig-

ured by Dall (1902: pl. 29, fig. 4; 1903: pl. 75, fig. 4 [same

Figures 9-11.

Scanning electron micrographs of Microcan-

cilla microscopica (Di ull, 1889 lectotype, USNM 82977. 9.

Apertural view. 10. Lateral view. 11. Protoconch. Scale bars

Figures 9, 10 = 1 mm; Figure 11 = 200 pm

Page 98

THE NAUTILUS, Vol. 121, No. 2

drawing]) and illustrated herein (Figures 9-11) is more

rounded at the anterior than in most specimens in the

syntype series and also has a heavier columellar callus.

Kaicher (1978: Card 1940) Pee al illustrated

this same specimen from the USNM type collection

(USNM 62977 |sic; error for Som as Admete micro-

scopica (Dall), Admete being the genus used by Dall in

1889b and 1903. She incorrectly felenied to this speci-

men as holotype but Dall did not designate a type speci-

men. The species was originally deserbe d from two lo-

calities and there are numerous specimens in the original

lots. Under Article 74.5 of the current Code (Interna-

tional Commission on Zoological Nomenclature, 1999)

Kaicher’s statement does not “qualify as a lectotype des-

ignation. In order to rectify this, USNM 82977, off

Yucatan, 366 m (originally 200 fathoms), is here desig-

nated lectotype of Cancellaria microscopica Dall,

1889. The type locality thus becomes restricted to

Campeche Bank, off Yucatan, Mexico.

ACKNOWLEDGMENTS

We are grateful to Mr, Enilson Cabral of the Research

and Management Center of Fishing Resources of the

Northeastern Coast — CEPENE/IBAMA for the collec-

tion of the material analyzed in the present work. This

study Was par tially supported by the Assistance to Sci-

ence and Technology Foundation of Pernambuco

(FACEPE/CNPq).

Dr. M. G. Harasewych, National Museum of Natural

History, Smithsonian Institution, W ashington, DC,

kindly furnished the SEM of Microcancilla microscopica

(Dall) and reviewed the manuscript.

LITERATURE CITED

Bouchet, P. and Warén, A. 1985. Revision of the Northeast

Atl ee bathyal and abyssal Neogastropoda excluding Tur-

ridae (Mollusca, Gastropoda). Bollettino Mal: acologico,

suppl. 1: 121-296.

Dall, W. H. 1889a. Reports on the results of dredgings, under

the supervision of Alexander Agassiz, in the Gulf of

Mexico (1877-78) and in the Caribbean Sea (1879-80), by

the U. S. Coast Survey Steamer “Blake”, Lieut.- Com-

mander ©. D. Sigsbee, U.S. N., and Commander J. R.

3artlett, U.S. N., commanding, XXIX. Bulletin of the Mu-

seum of Comparative Zoology 1S: 1-492, pls. LO-40.

Dall, W. H. 18S9b. A preliminary catalogue of the shell-bearing

marine mollusks and brachiopods of the southeastern

coast of the United States, with illustrations of many of the

species United States National Museum Bulletin 37:

1-221, pls. 1-74. [Reprinted 1903 with additional plates:

1-232, pls. 1-95.].

Dall, W. H. 1902. Illustrations and descriptions of new, unfig-

ured, or imperfectly known shells, chiefly in the U. S.

National Museum. Proceedings of the United States Na-

tional Museum 24(1264); 499-566, pls. 27-40.

Dall, W. H. 1903. A preliminary catalogue of the shell-bearing

marine mollusks and brachiopods of the southedertn

coast of the United States, with illustrations of many of the

species. United States National Museum Bulletin 37:

—232, pls. 1-95.

Dall, W. H. 1924. Notes on molluscan nomenclature, Proceed-

ings of the Biological Society of Washington 37: S790.

Harasewych, M. G. and Petit, R.E. 1986. Notes on the mor-

phology of Admete seein (Gastropoda: Cancellariidae).

The Nautilus 100(3); 85-91.

International een on Zoological Nomenclature, 1999.

International Code of Zoological Nomenclature. Fourth

Edition. International Trust for Zoological Nomenclature,

London, xxix + 306 pp.

Kaicher, S$. D. 1978. Pack #19, Cancellariidae.

of world-wide shells. Cards 1559-1964.

Knudsen, J. 1964. Scaphopoda and Gastropoda from depths

exceeding 6000 meters. Galathea Report 7: 125-136,

Maxwell, P.A. 1992. Eocene Mollusca from the vicinity of Mc-

Culloch’s Bridge, Waihao River, South Canterbury, New

Zealand: Pale oecology and systematics. New Zealand Geo-

logical Survey Pe leontological Bulletin 65: 1-280.

Petit, R.E. and Harasewych, M.G. 2005. Catalogue of the su-

perfamily Cancellarioidea Forbes and Hanley, 1851 (Gas-

tropoda: Prosobranchia) - 2"4 edition. Zootaxa 1102:

1-161.

Thiele, J. 1925. Gastropoda der Deutschen Tiefsee-E —

I. Teil. Deutsche Tiefsee-Expedition 1595-1899, 17(2

35-382, Pls. 13-46.

Verhecken, A. 1997. Mollusca Gastropoda: Arafura Sea Can-

cellariidae collected during the Karubar Cruise. Mémoires

du Muséum National d'Histoire Naturelle 172: 295-323.

Card catalogue

THE NAUTILUS 121(2):99-103, 2007

Page 99

A new species of Gerdiella (Gastropoda: Cancellariidae) from

the South Atlantic Ocean off Brazil with discussion of an

undescribed species

Silvio Felipe B. de Lima

José Carlos N. de Barros

Departamento de Pesca e Aquicultura

Universidade Federal Rural

de Pernambuco

Avenida Dom Manuel de Medeiros, S/N,

Dois Irmaos

52171-030, Recife, BRAZIL

[email protected]

re. [email protected]

Richard E. Petit

806 St. Charles Road

North Myrtle Beach,

SC 29582-2846 USA

ABSTRACT

Two rare species of Cancellariidae were identified during the

study of material from oceanographic dredge hauls indlepalken

in 2000 by the fishing vessel NATUREZA in ‘deep waters off the

state of Pernambuco, Brazil. The species belong to the genus

Gerdiella Olsson and Bayer, 1972. Gerdiella alvesi new species

is similar to Gerdiella c ‘ingulata Olsson and Bayer, 1972, as both

have strong, nodular ornamentation that is coarsely cancellated

and a heavily thickened, lirated outer lip. A second species,

Gerdiella sp.. is identified based on the protoconch, cancellated

ornamentation and the presence of two columellar folds, dis-

tinguished from the species described herein by its ornamen-

tation and the absence of a subsutural keel.

Additional keywords: Mericella, Neogastropoda, bathyal, Per-

nambuco

INTRODUCTION

The family Cancellariidae Forbes and Hanley, 1851,

represented by a large number of fossil and recent gas-

tropods da bated among diverse marine regions

throughout the world. The group inhabits subtidal to

bathy al sandy and muddy bottoms of tropical and tem-

perate regions, with the greatest diversity found along

the eastern Pacific coast of the Americas and the ceniel

Indo-Pacific area (Harasewych and Petit, 1982). In the

western Atlantic Ocean, the number of known species is

still relatively small especially with regard to the Brazilian

coast (see for instance Harasewych et al. 1992).

The genus Gerdiella Olsson and Bayer, 1972, was in-

troduced to include three species described by these two

authors from bathyal depths of the Florida ‘Straits and

south of Jamaica. These species are: Gerdiella gerda

from the Straits of Florida, 648-622 m: G. santa from the

Straits of Florida, 64S—622 m: and G. cingulata from S of

Jamaica, 549-530 m. Another specimen of G. cingulata,

collected in 1961 by R/V OrEGON, sta. 3552, 130 miles

ESE of New Orleans, Louisiana, 29°07’ N, 88°05’ W,

trawled in 732 m, is now catalogued as USNM 811462.

No additional species of Gerdiella have been discovered

until now.

The genus Mericella Thiele, 1929, was introduced by

Thiele to accommodate the bathyal Mericella jucunda

(Thiele, 1925) from off Tanzania. He originally placed

the species in Cancellaria (Merica). Morice. bozzetti

Petit and Harasewych, 1993, was described from off So-

malia. Petit and Harasewych at the same time placed

Cancellaria (Merica) paschalis Thiele, 1925, in the genus

Mericella. Mericella paschalis was described from a bro-

ken fragment, but recently collected material from off of

Mozambique allowed Verhecken and Bozzetti (2006: 15-

16) to confirm the allocation of the species in Mericella.

In a recent paper, Verhecken and Bozzetti (2006)

ae Gerdiella in the synonymy of Mericella Thiele,

1929. As observed by those two authors, Mericella was

ed by Olsson and Bayer in the original description

of Gerdiella. Verhecken and Bozzetti (2006: 17) stated

that the two genera are “very much alike conchologically,

the main differences being ‘the relative spire height and

the suture form.” They also considered relative aperture

heights, observing that, as Gerdiella has a shorter aper-

ture, the ratio in this latter genus agrees “with Petit and

Harasewych (1993: 223) Go comer a value of 50.5 a

diagnostic feature for Mericella.” Verhecken and

Bozzetti did not point out that Petit and Harasewych

used additional characters to differentiate these genera.

Verhecken and Bozzetti also stated that “there are no

important differences in shell characteristics that would

justify a separation between Mericella and Gerdiella.”

Although shown on their table, the text does not mention

the fact that Gerdiella species have axial ribs on the pro-

toconch. However, in an earlier work Verhecken (ohos:

Page 100

THE NAUTILUS, Vol. 121, No. 2

513) stated that “protoconch characters are not consid-

ered of diagnostic importance at generic level by this

writer.” We disagree with that approach, especially when

protoconch characters allow for the distinction of west-

ern Atlantic taxa from those from the Indian Ocean. We

consider Gerdiella to be a valid genus with species

known at present only from the w vestern and southern

Atlantic Ocean.

The geographical grouping of Mericella, with all

known species being from off eastern Africa, and Gerdi-

ella, with all known species being from the western and

southern Atlantic, is obvious.

Verhecken and Bozzetti (2006: 17) mentioned that the

eastern Pacific Cancellaria corbicula Dall, 1908, was

placed in Gerdiella by Kaicher (1978: card 1952). We do

not agree with that placement as the species has a

euneatl protoconch and an aperture height greater than

one-half shell height. Its generic placement remains in

doubt.

Verhecken (2002: 512) studied three juvenile and frag-

mented shells collected from the Continental Slope at

Pernambuco, northeastern Brazil, during probes of the

CHALLENGER EXPEDITION in 1873. Those specimens

were considered by him to be conspecific and the pos-

sibility that they represent a new species of Gerdiella was

mentioned. We agree with Verhecken that more and

better specimens are needed for identification to be con-

firmed.

In this paper we describe a fourth species of Gerdiella

from the Western Atlantic, collected from the Continen-

tal Slope off Pernambuco, Brazil. This is the first definite

record of the genus for the South Atlantic. The soft parts

and radula are as yet unknown, but the conchological

characters are sufficient to justify the naming of a new

species.

MATERIALS AND METHODS

All specimens examined were obtained by the research

vessel NATUREZA along the Continental Slope off Per-

nambuco during oceanic prospecting work for the Re-

search and Management Center of Fishing Resources of

the Northeastern Coast—CEPENE/IBAMA. At the Ma-

lacology Laboratory of the Universidade Federal Rural

de Pernambuco, the specimens were sorted under a ste-

reomicroscope, cleaned in a diluted detergent solution,

rinsed in distilled water, and air-dried. Shells were mea-

sured using a stereomicroscope with eyepie ce microme-

ter and photographed with a Nikon COOLPIX 885 digi-

tal camera. Scanning electron micrographs were made

using a Jeol JSM 6360 Scanning Electron Microscope at

the Electron Microscope Laboratory of the “Instituto

Tecnolégico de Pernambuco (ITEP)”

Abbreviations used: ANSP, Academy of Natural Sci-

ences Philadelphia: LMUFRPE, Laboratério de Mala-

cologia da Universidade Fedral Rural de Pernambuco,

Brazil; MNRJ, Museu Nacional, Rio de Janeiro, Brazil:

MORG, Museu Oceanografico do Rio Grande, Rio

Grande do Sul, Brazil; MZUSP, Museu de Zoologia da

Universidade de Sao Paulo, Brazil.

SYSTEMATICS

Family Cancellariidae Forbes and Hanley, 1851

Genus Gerdiella Olsson and Bayer, 1972

Type Species: Gerdiella gerda Olsson and Bayer,

1972 by original designation. Recent, Caribbean.

Gerdiella alvesi new species

(Figures 1—5)

Description: Shell fusiform, stout, white, strongly or-

namented axially and spirally, entirely marked by growth

lines between spiral cords. Protoconch globose, cap-

shaped, with 1.5 whorls, ormamented by numerous mi-

croscopic spiral threads and weak axial ribs on final por-

tion. Transition to teleoconch marked by strong raised

axial rib. Nucleus small when compared with rest tof pro-

toconch, which is inflated. Teleoconch with 5.5 whorls.

Spire narrow, smaller than body whorl. Whorls rounded,

strongly omamented, with reticulated sculpture. Spiral

cords intersect the equally strong axial ribs, forming

strong nodules with a pustulose aspect, which progres-

sively increase in size toward body whorl. First whorl

with 20-22 axial ribs and 6-7 spiral cords, more often 6,

Second whorl with 20 axial ribs and 7 spiral cords, third

whorl with 22 axial ribs and 7 spiral cords, fourth whorl

with 26 axial ribs and 7 spiral cords, fifth whorl with

32-33 axial ribs and 4 spiral cords. Body whorl with 32

axial ribs and 4 upper spiral cords. Suture strongly con-

stricted, bordered by a strong, nodular, subsutiel spiral

cord. Base imperforate, strongly conical and gently con-

vex, ornamented by 15 nodular spiral cords, 5 of which

form siphonal fasciole. Aperture elliptical, fusiform, nar-

row at terminations. Peristome shiny, very thick and

strongly expanded. Outer lip thick, with a broad posterior

sinus, withl2 denticles, most anterior denticle more

elongated. Parietal region strongly reflected, with inter-

nal nodules. Columella gently concave, with two pro-

nounced, rounded folds, the adapical fold slightly larger

Siphonal canal short and narrow in distal extremity.

Type Material: © Holotype, MZUSP 75932 [Length 22

mm, Width 8.3 mm]; juvenile shells: 1 paratype, ANSP

413550; 3 paratypes, MORG 50.658; 2 paratypes, MNR]

LOTIS; 4 paratypes, MZUSP 78933. All from type local-

1S Noy. 2000.

Type Locality: Northeastern Brazil, off the State of

Pernambuco, 08°46.5' S, 34°44.5' muddy bottom,

690 im.

Geographical Distribution: Continental slope off

Pernambuco, 08°46.5' S, 34°44.5' W, northeastern Brazil.

S. F. B. de Lima, J. C. N. de Barros and R. W. Petit, 2007 Page 101

Figures 1-5.

Gerdiella alvesi new species, holotype MZSP 78932, length

22 mm. 1. Apertural view; 2. Detail of aperture; 3.

Lateral view showing profile of outer lip. 4. Protoconch. 5. View of ornamentation of second and third teleoconch whorls. Scale bars

Figures 2, 3,5 = 500 pm; Figure 4 =

200 zm

Etymology: Named in honor of Dr. Marcos Souto

Alves of the Biology Department, in the field of Zoology

at the Universidade Federal Rural de Pernambuco

UFRPE) for having sent the first author under an in-

ternship at the Malacology Laboratory of the UFRPE.

Remarks: The protoconch of the holotype is not well

illustrated as it is damaged. Nonetheless, we were able to

characterize the protoconch of juvenile specimens of the

new species (Figures 6-1]

Gerdiella sp

Figures 15-19

Material Examined: One damaged specimen

MZUSP 78934. length 18.3 mm, off the state of Pernam-

buco, northeastern Brazil, 08°46.5' S, 34°44.5' W,

muddy bottom, 690 m, 18 Noy. 2000

Geographical Distribution: The Continental Slope

off Pernambuco, northeastern Brazil.

Remarks: The single specimen of Gerdiella sp. may

represent a new species. However, we prefer not to

name it as the shell is damaged and eroded. The proto-

conch of this species has the same characteristics, and

the same number of whorls, as Gerdiella alvesi. This

specimen has two equal and very narrow columella

folds, slightly different from those of G. alvesi. The first

second and third post-nuclear whorls are rounded and

ornamented by finely cancellated axial ribs and spiral

cords, forming small nodules at their intersections, simi-

lar to those of Gerdiella gerda and Gerdiella santa. The

axial ribs are thicker than the spiral cords, (9 on the body

whorl), as opposed to 6 for G. alvesi. The subsutural cord

is weak on the first and second whorls of the teleoconcl

Page 102

THE NAUTILUS, Vol. 121, No. 2

Figures 6-14. Juveniles of Gerdiella alvesi new species. 6-8. Protoconch under SEM, MORG 50.688; 9. SEM of protoconch of

paratype MORG 50.688 showing microscopic spiral threads. 10-11. SEM of paratypes MORG 50.688. 12-13. Paratype, MZUSP

78933. 14. SEM of paratype MORG 50.688 showing growth lines. Scale bars: Figures 6—S = 200 wm; Figure 9 = 50 jm; Figures

10-13 = 500 jm; Figure 14 = 50 pm.

The subsutural region is flat, encompassing the first and

second spiral cords, which do not form a shoulder. There

are seven spiral cords on the first and second whorls, and

10 on the third. There are 27, 30, and 36 axial ribs on the

first, second and third post-nuclear whorls, respectively.

In relation to G. alvesi, Gerdiella sp. has the same num-

ber of spiral elements on the first and second post-

nuclear whorls. This number is higher, however, on the

third whorl. The number of axial ribs on the first three

whorls of the teleoconch of G. alvesi is less than that of

Gerdiella sp.

DISCUSSION

The conchological characters of Gerdiella sp. cannot be

completely and conclusively compared to any of its con-

geners until better material is collected for study at the

species level. The analysis presented above distinguishes

this species from the unnamed shell figured by Ver-

hecken (2002: figs. 9, 10) by the greater number of axial

and spiral ribs and threads on the first and second whorl.

Gerdiella alvesi stands out from its congeners by the

strong, uniform axial and spiral ornamentation, which

forms large. rounded nodules that are quite pronounced

pecially on the body whorl. The heavily thickened

outer lip is very similar to that of Gerdiella gerda, prin-

cipally on the sigmoid contour, and is lirated in the same

way as Gerdiella cingulata, but not as coarsely as de-

scribed by Olsson and Bayer (1972: 879). Two columellar

folds are present on all the species. In both G. cingulata

and G. alvesi the adapical fold is larger and there are no

tubercles between the folds in the latter of the two spe-

cies. In G, alvesi, there is no projection of the parietal

callus on the outer lip, which is present in G. gerda and

G. santa. Among the Gerdiella, the spire of G. alvesi has

the least number of whorls and lacks varices.

ACKNOWLEDGMENTS

We are deeply grateful to: Enilson Cabral of the Re-

search and Management Center of Fishing Resources of

the Northeastern Coast—CEPENE/IBAMA—for his

personal efforts in the collection of sediment from the

Continental Slope off northeastern Brazil and for the

donation of all this conchological material to the Mala-

cology Laboratory of the Universidade Federal Rural de

Pernambuco, Brazil; Fernanda Emanuele, Instituto Tec-

noldgico de Pernambuco (ITEP) for taking the electron

scanning micrographs; Mr. Richard Boike for an early

translation of the text into English. Dr. M. G. Harase-

S. F. B. de Lima, J. C. N. de Barros and R. W. Petit, 2007

Page 103

Figures 15-19. Gerdiella sp.. MZSP 78934, length = 18.3 mm. 15. Abapertural view. 16. Columellar folds under SEM. 17. View

of ornamentation and subsutural flattening of the second and third teleoconch whorls. 18-19. Protoconch, Scale bars: Figures 16,

18, 19 = 200 pm; Figure 17 = 500 pm.

wych, National Museum of Natural History, Smithsonian

Institution, Washington, DC, kindly reviewed the manu-

script.

LITERATURE CITED

Harasewych, M. G. and R. E. Petit. 1982. Notes on the mor-

phology of Cancellaria reti: ulata (Gastropoda: Cancellari-

idae). The Nautilus 96: 104-113

Harasewych M. G.. R. E. Petit, and A. Verhecken. 1992. Two

new species of Cancellariidae (Gastropoda: Neogas-

tropoda) from Brazil. The Nautilus 106: 43-49

Kaicher. $. D. 1978. Pack #19, Cancellariidae. Card catalogue of

world-wide shells. Cards 1859-1964

Olsson, A. A. and F. M. Bayer. 1972. Gerdiella, a new genus of

deep-water cancellariids. Bulletin of Marine Science 22

8$75-S80

Petit, R. E. and M. G Harasewych 1993. A new Mericella

(Mollusca: Gastropoda: Cancellariidae) from northeastern

Africa. Proceedings of the Biological Society of Washing-

ton, LOG: 221-994

Thiele, J. 1925. Gastropoda der Deutschen Tiefsee-Expedition

II. Teil. Deutsche Tiefsee-Expedition 1598-1899, 17(2):

35-382, pls. 13-46. [Dual pagination; also numbered

1-348, pls. 1-34. |

Thiele, J. 1929-35. Handbuch der systematischen Weich-

tierkunde. Gustave Fischer, Jena. 2 vols. [1(1), 1-376

(1929): 1(2), i-vi, 377-778 (1931); 2(3), 779-1022 (1934)

9(4), i-vi, 1023-1154 (1935)].

Verhecken, A. 2002. Atlantic bathyal Cancellariidae (Neogas-

tropoda: Cancellarioidea): Additional data and description

of a new species. Journal of Conchology 37: 505-514

Verhecken, A. and L. Bozzetti. 2006. New data on East-African

Mericella species, and description of a new species of

Scalptia (Neogastropoda Cancellarioidea: Cancellariidae

Gloria Maris 45: 14-25

THE NAUTILUS 121(2):104, 2007

Page 104

Book Review

Land and Freshwater Molluscs of Brazil

Luiz Ricardo L. Simone. 2006. Land and Freshwater Mol-

luses of Brazil. Museu de Zoologia, Universidade de Sao

Paulo, Sao Paulo, Brazil. 390 pp., including over 1100 text-

figures. ISBN 859066700-6. 8.5 by 11 in. Hardback; in En-

ali sh. $125 from US booksellers: 92.5 to 94 € in Europe.

This large-scale work is a timely and sorely needed rem-

edy for one of the most stark deficiencies in the global

inventory of molluscan biodiversity. The author has re-

cruited no less than 70 identified collaborators and used

the resources of 27 prominent institutional collections to

formulate a presentation of 1074 native and 33 intro-

duced species-level molluscan taxa inhabiting the land

and fresh waters of Brazil and/or nearby areas. “The com-

position of this native non-marine fauna (about 700 ter-

restrial and over 370 aquatic, over 950 gastropods, and

116 pelecypods) invites comparison with other areas such

as the USA, where freshwater clams are notably more

speciose, and non-marine snails occur in roughly com-

parable diversity.

The work is organized thus: a one-page Introduction, four

pages of legends and acknowledgements, a Table of Contents,

which is a systematic outline of taxa treated, 278 pages of

consistently formatted illustrations and companion text blocks

in telegr. aphic style, a bibliography of 2696 titles, and an index

of all taxa from phylum to species-level, the later presented

with trivial name first.

The format is simple and efficient. The Introduction indi-

cates the method of presentation of taxa, each consecutively

numbered in a conventional phylogenetic sequence (genera

non- menage family unit, but species alphe ibetical

within genus) The reiterative eye- -catching marginal icons

(up to four) in each text block are a ad: geographic dis-

tribution (blue globe); literature citations (re d printe sd page):

source of companion figure, each of which is like-numbered

(green eyeball); a non-critical synonymy (equal sign on or-

ange); and “N.B.” for random notations when appende cl. The

English diction is deficient on this page, but the author's

intent is generally comprehensible.

One defining feature of the work is enunciated in the In-

troduction: Simone characterizes the figures as “normally

based on type specimens.” Close perusal confirms this as the

case, with name-bearing types from virtually all of the cited

(27) institutional collections being depicted in dedicated pho-

tographs. For the exceptions, it is apparent that paratypes,

voucher specimens, iconotypes, and other levels of authen-

icity were assiduously pursued and exploite od: for the slug

groups this proved generally impossible. The photogr: iphs are

generally of high fide lity, with appropriate magnification to

acilitate identification. About three dozen photogr: iphic vi-

gnettes, mostly of living snails, appropriately placed at various

oints add a dime sion of vitality.

Bibliographic citations are arr anged in near flawless alpha-

vetical-chronological sequence and numbered consecutively.

One or more such numbers appear in each text block, and an

attempt is made to code them with one to six lower case code

letters indicating if the work contained a description, figure,

etc. Regrettably there are many omissions of the important

“On code. which indicates “original,” as in description. The

concermed reader must alphi ibetically search the bibliography

using author and date expressed after the binomen at the

heading of such entries. Perhaps unfortunately, the works of

: ae and Pilsbry are particularly prone to this oversight.

The fidelity of the citations appears to be excellent, although

Bahiensis miliola (no. 591) ) appears to date from @’ Orbigny,

1837, rather than 1835 as stated (Pilsbry, 1901: 32: Sherborn

and Griffin, 1934). A minor and easily remediable biblio-

graphic nuisance is the lack of identification of the G. B.

Sowerbys by generation,

In the course of the work several generic reassignments

(clearly marked as “n. comb.”) are install d and nomina nuda

revealed. Species no. 1071, Byssanodonta riograndensis

(Ihering and Morretes, 1949), is thus designated, but a pho-

eee of an ANSP specimen accompanies this entry. Even

though the Code eins 1999: Article 13.1.1), since 1930, has

not recognized a binomen and figure indication in the ab-

sence of a written description as basis for an available name—

in this instance attributable to Simone—an explanation for his

treatment of this apparent taxon would be welcome by the

reader. Does Simone think this is a valid, unnamed taxon?

The specimen figure on p. 309 captioned Lamellaxis

clavinulus (Potiez and Michaud, 1838) ) appears not to be that

species but L. micrus (@Orbigny, 1835), which is treated as

Allopeas micra {sic] on p. 14. On p. 312, Europe is given as

the origin of the non-native Bradybaena similaris (Férussac,

1821). It is more li kely from eastern Asia.

There are technical problems with the typesetting such as

wholesale deletion of dozens of single letters, particularly

noteworthy on page 23, and the occasional misspelling or

improper diacritical mark can be a minor distraction.

The foregoing minor critique notwithstanding, Simone’s

book is monumental. It is a prodigious work in both scope and

the quality of the research. It is certain that all serious work-

ers will find it indispensable in the understanding of the ex-

tensive and complex Neotropical malacofauna. On another

plane, its application to the analysis of other major world

faunas will impel us to a better appreci ation of the systemat-

ics, evolution, zoogeography, and macroecology of nonmarine

Mollusca on a global scale.

LITERATURE CITED

International Commission on Zoological Nomenclature (ICZN). 1999.

Intemational Code of Zoological Nomenclature. Fourth edition

International Trust for Zoological Nomenclature, London, xxix +

306 pp

Pilsbry, H. A. 1901-02. Manual of Conchology (second series). 14.

Oriental bulimoid Helicinidae. Academy of Natural Sciences,

Philadelphia. [iv] + 302 + xcix pp, 62 pls

Sherborn, C. D. and F. J. Griffin, 1934. On dates of f pub lication of the

natural history portions of Alcide d'Orbigny’s “Voyage dans

Annals and Magazine of Natural His-

tory series 13 1L0(13): 130-134.

Harry G. Lee

1132 Ortega Forest Drive

32210 USA

r Ame rique me sridioné ile

Jacksonville, FI

shells@hglee com

| MBL WHO! Libr

uni)

5 WH

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Volume 121, Number 3

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Volume 121, Number 3

October 10, 2007

CONTENTS ISSN 0028-1344

Gary W. Schmelz The Epitoniidae (Gastropoda: Ptenoglossa) from the lower Alum Bluff

Roger W. Portell Group (lower to middle Miocene) of Florida, with descriptions of nine

NEW SPSClES 5 ke alar aed SAGE OR NewS Se Mend Wc @ Aldean aiararin 105

Francisco M. Heralde III Molecular phylogeny of some Indo-Pacific genera in the subfamily Turrinae

Maren Watkins H. Adams and A. Adams, 1853 (1838) (Gastropoda: Neogastropoda) ... . . 13]

John-Paul Ownby

Pradip K. Bandyopadhyay

Ameurfina D. Santos

Gisela P. Concepcion

Baldomero M. Olivera

Claudia Muniain Reproductive biology of the nudibranch Doris fontainei @Orbigny, 1835

Carlos S. Gallardo (Gastropoda: Opisthobranchia) from the Magellanic Region ........... 139

Pablo E. Penchaszadeh

Juliana M. Harding Two modern records of the southern oyster drill Stramonita haemastoma

M. G. Harasewych floridana in Chesapeake Bay, USA 2... 2. ee 146

Paolo Mariottini Brachycythara beatriceae, a new species from the Alboran Sea and the

eastern Atlantic Ocean (Gastropoda: Neogastropoda: Conidae) ......... 159

Sponsored in part by the State of Florida, Department

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THE NAUTILUS 121(3):105-130, 2007

Page 105

The Epitoniidae (Gastropoda: Ptenoglossa) from the lower Alum

Bluff Group (lower to middle Miocene) of Florida, with

descriptions of nine new species

Gary W. Schmelz!

5575 Dogwood Way

Naples, FL 34116 USA

P.O. Box 117800

Gainesville, FI

Roger W. Portell

Florida Museum of Natural History

University of Florida

32611-7800 USA

[email protected]

ABSTRACT

A comprehensive study of the family Epitoniid: ve that exists in

the lower to middle Miocene portion of the Alum Bluff Group

of Florida (USA) was conducted. A total of 14 species was

examined. Of these, 12 are considered valid members of the

family Epitoniidae. They include three previously described

species, Amaea gardnerae, Cirsotrema cirritum, and Epitonium

virginiae, plus nine new species. Seven of the new species were

collected from the Chipola Formation and two from the Shoal

River Formation. One new species is assigned to the genus

Cirsotrema, two are assigned to the genus Opalia, and six are

assigned to the genus Epitonium.

Cirsotrema previously reported from the Chipola Formation

as Cirsotrema dalli, a Pleistocene to Recent species, has been

described as a new species. Epitonium alaquaense reported

from the upper Miocene Choctawhatchee Formation (upper

Alum Bluff Group) and questionably placed in the Shoal River

Formation fauna by Gardner (1947) is no longer recognized as

a Shoal River Formation species and Gegania acutissima has

been placed with the Architectonica-like members of the family

Mathildidae

Additional Keywords: Miocene, Chipola Formation, Shoal

River Formation, Cirsotrema, Opalia, Epitonium, Chipola

River, Tenmile Creek, Farley Creek

INTRODUCTION

The family Epitoniidae has an extensive history with rep-

resentatives dating back to the early Mesozoic. According

to Clench and Tumer (1950), the group appears to have

reached its peak of diversity during the Eocene and Mi-

ocene epochs. In Florida (USA), members of the family

are well represented in early Miocene deposits and, to a

lesser extent, in middle Miocene deposits of the Alum

Bluff Group.

The Alum Bluff Group consists of five named strati-

graphic units (Figure 1). From oldest to youngest these

units are the Chipola Formation, Oak Grove Sond. Shoal

' Author for correspondence

River, Choctawhatchee, and Jackson Bluff formations

(Huddlestun, 1984). All of the Alum Bluff strata occur in

outcrops in the western portion of the Florida panhandle

(Figure 2). The lower Miocene Chipola Formation was

depositec | approximately 1S mya (Jones et al., 1993) and

outcrops along Tenmile, Farley, and Fourmile creeks,

and the (¢ Chipola, Yellow, Choctawhatchee, and Apalachi-

cola rivers. To date, most Chipola Formation specimens

have been collected from Tenmile, Farley, and Fourmile

creeks, and the Chipola River. Collections of fossil speci-

mens from the Oak Grove Sand along the Yellow River

and the Chipola Formation along the Apalachicola River

are limited, given that these locations typically can only

be accessed during very low water levels. According to

Vokes (1989), Tenmile Creek, F arley Creek, and C hipola

River complex alone encompasses over 7.5 miles (~12

kim) of Chipola Formation exposures. The middle Mi-

ocene Shoal River Formation was de posite od approxi-

mately 12 mya (Jones et al., 1993). All known Shoal River

Formation outcrops are west of the Chipola Formation

exposures with most of the collecting areas situated along

the Shoal River in Walton County. Overviews of the ge-

ology, stratigraphy, and paleontology of the Chipola aad

Shoal River aa can be found, respectively, in

Vokes (1989) and Portell et al. (2006).

Gardner (1947), as part of her monographic treatment

f the ae ee in fauna of the Alum Bluff Group, dis-

cussed three species belonging to the family Epitoniidae.

These species were Epitonium (Spiniscala) virginiae

(Maury, 1910), Epitonium (Clathrus ) ) alaquae nse (Mans-

field, 1935), and Gegania acutissima (Dall, 1892). How-

ever, only two of these taxa are herein considered to

belong to the family Epitoniidae. Epitonium virginiae

was collect d from a Chipola Formation site along the

east bank of the Aps ilachicola River. It was describe ad by

Maury (1910) from a single, extremely small (3.7 mm

maximum height x 1.5 mm maximum width) specimen

that was part of the Cornell University Collection (now

deposited at the Paleontological Research Institution).

Epitonium alaquaense, recorded by Mansfield (1935)

Page 106

THE NAUTILUS, Vol. 121, No.

1c)

Cooke (1945)

DUPLIN MARL

(Ecphora and

Cancellaria zones)

Huddlestun (1984)

JACKSON BLUFF

(Ecphora and

Cancellaria zones)

Vokes (1989)

JACKSON BLUFF

(Ecphora and

Cancellaria zones)

Scott

(2001)

Carter et al. (2003)

JACKSON BLUFF

(Cancellaria zone)

This Paper

JACKSON BLUFF

(Ecphora and

Cancellaria zones)

PLIOCENE

CHOCTAWHATCHEE

RED BAY FM

Arca rubisiniana

zone

(Arca zone)

CHOCTAWHATCHEE

RED BAY FM —M

(Arca zone) (Arca zone)

Upper

(Yoldia zone)

Yoldia waltonensis

Middle

(Main shell bed)

MIOCENE

SHOAL RIVER FM

Glycymeris

waltonensis

zone

WHITES CREEK

MEMBER

YELLOW RIVER FM

(Yoldia zone)

SHOAL RIVER FM

YELLOW RIVER FM

CHOCTAWHATCHEE FM.

(Yoldia zone) SHOAL RIVER FM

(Includes

“Cardium beds”

of Gardner (1926),

Yoldia and Glycymeris

zones, and

Whites Creek Member)

ALUM BLUFF GROUP

SHOAL RIVER FM

ALUM BLUFF GROUP (UNDIFFERENTIATED)

WHITES CREEK BEDS

Cardium taphrium

zone

OAK GROVE SAND

CHIPOLA FM CHIPOLA FM

Figure 1

from the Arca zone of the Choctawhatchee Formation, is

late Miocene (Huddlestun, 1984). When Gardner (1947)

collected a shell fragment similar to Mansfield’s shell at

a Shoal River site she included it as part of the lower

Alum Bluff Group as defined by Cooke (1945). The col-

lective evidence now suggests that E. alaquaense should

not be listed as part of the fauna of the lower Alum Bluff

Group (see Figure 1). The reasons for this are twofold.

First, an extensive examination of the Chipola and Shoal

River formation fossils in the Florida Museum of Natural

History (including Tulane University and Florida Geo-

logical Survey enlle sctions), Paleontologic: il] Research In-

stitution, Smithsonian’s National Museum of Natural

History, Museum of Natural History, The

Academy of Natural Sciences, Alabama Ge ologic: ul Sur-

vey, and private collections during, this study did not

American

uncover a single specimen that could be referred to as E

(1947:

from *

no7

577) stated that she

shell fragment ‘a horizon. slightly

than the Based

upon this remark and that no additional specimens simi-

lar to E

mentioned collections, it seems fair to assume that Gard-

alaquae nse. Second, Gardner

collected her

higher typical Shoal River formation.”

alaquaense were ever found in the above-

ner s specimen be longe d to strati tyounger than the Shoal

River Formation; most probably to the upper Miocene

Choctawhatchee Formation

Dall (1892) described Tuba acutissima and placed it in

the family Mathildidae

Tuba

Gardner (1947) replaced the ge-

nu with Gegania and tentatively assigned it to the

OAK GROVE FM. /,

CHIPOLA FM

Alum Bluff Group stratigraphic nomenclatural history and correlation (in part).

OAK GROVE SAND OAK GROVE SAND

CHIPOLA FM CHIPOLA FM

Modified from Portell et al. (2006),

family Epitoniidae because she felt that its morphologi-

cal characteristics more close ly coincided with the sub-

order Ptenoglossa. Gardner's generic assignment of Ge-

gania was based upon the shell similarity to Gegania

pinquis Jeffreys, 1884, a species collecte d during the

Porcupine Expedition off Cape Mondego, Portugal.

However, the genus Gegania has since been assigned to

Architectonica-like members of the family Mathildidae

(Vaught, 1989).

In addition to the three species of Epitoniidae listed

for the Alum Bluff Group, Gardner (1947: 575) reported

shell fragments belonging to “at least a dozen”

Gardner stated that most of the specimens were so im-

species.

pe rfectly preserve od that only subge meric determinations

could be made. Four of the unidentified « :pitoniids came

from the Aldrich Collection (housed at Johns Hopkins

University) and the remaining species were from Gard-

ner’s collection, Eleven of the fragments were obtained

from Shoal River Formation localities and two were col-

lected from Chipola Formation sites. Gardner placed

eleven of the fragments in the genus Epitonium, one in

the genus Scalina, and one in the genus Gegania. Six

were placed in the subgenus Hyaloscala, two in Spinis-

cala, three in Cinctiscala, and one in Nodiscala.

[t is apparent from Gardner's (1947) discussion on the

Epitoniidae of the Alum Bluff Group that future work

remained to be done on the family. ae her publica-

tion, three additional species of EF pitoniid ie have been

reported from the Chipola Formation. These are Cir-

G. W. Schmelz and R. W. Portell, 2007

Page 107

Jackson

Alum Bluff

Group

Figure 2... Map of Florida showing counties with Alum Bluff Group sediments found at or near the surface.

sotrema dalli Rehder, 1945; Scalina gardnerae (Olsson,

1967): and Cirsotrema cirritum Duerr, 2004. Cirsotrema

dalli is an extant species reported to have undergone

little morphological change since the early Miocene (Ols-

son, 1967). Scalina gardnerae was described by Olsson

(1967) from a single large specimen collected from Mc-

Clellan’s Farm in Calhoun County, Florida. It is a fairly

common species, and since its discovery, it has been

found at numerous Chipola Formation sites by the au-

thors and other investigators. Cirsotrema cirritum is a

much less common species collected at a few sites along

Tenmile and Farley creeks and the Chipola River (Du-

err, 2004).

Scattered among the material in museums and private

collections are a large number of epitoniids from the

Chipola and Shoal River formations of Florida that have

been amassed over the last fifty years. Since many of the

recently collected specimens are in excellent shape, we

have undertaken the task of identifying and describing

the new species and providing better descriptions and

updated taxonomic placements of existing ones. As will

be evidenced in this paper, many of these epitoniids have

shell structures that are similar to those of extant and

fossil species from Florida, the Caribbean, westem Eu-

rope. and the eastern Pacific.

According to Clench and Tumer (1951), DuShane

1979). Kilburn (1985), Nakayama (2003), Robertson

(1983a; 1983b; 1993), Weil et al. (1999), and others, cur-

rent classification of the Epitoniidae is based upon shell

characteristics. For this study, morphological features

such as shell size (height and width), number and shape

of the nuclear whorls, number and shape of teleoconch

whorls, number and placement of the costae on the body

whorl, presence or absence of varices, type of sculpturing

on the body whorls, shape of the aperture, shape and

thickness of the outer lip, depth of the suture, presence

of punctae, and the spire angle were used to help differ-

entiate species. Spire angles were measured from pho-

tographs. A vertical line was drawn through the axis of

the shell and a protractor was used to determine the

angle between the margins of the body whorls on both

sides of the shell.

Additionally, in order to ascertain whether or not the

Chipola Formation species of Cirsotrema is the same as

the Recent Cirsotrema dalli, we closely examined the

varices on both forms. As noted by Weil et al. (1999) and

others, the presence or absence of varices can be a key

diagnostic feature for some species of Epitonium. We

believe that the structural configuration and the number

of varices that appear within a specific generic group,

such as in certain Epitonium, will change over geologic

time and that this morphological difference between the

Recent and fossil forms is significant enough to justify

naming the fossil form as a separate species. In part, this

Page 108

THE NAUTILUS, Vol. 121, No. 3

decision to use the changes observed in varix count as a

diagnostic feature at the species level is based upon the

precedence of using the number, shape, and configura-

tion of varices as a method of identifying different : spe-

cies and genera in the family Muricidae. This technique

has been used broadly by muricid gastropod investigators

(Herbert, 2005, and references therein).

Besides examining the varices on Cirsotrema dalli, we

also abraded the external portion of an uncatalogued Re-

cent shell in order to compare its underlying sculpture

with that of its fossil counterpart.

Even though many of the epitoniids examined are in

excellent shape, the task of classifying them and placing

them into their appropriate generic and subgeneric

groupings was challenging. This is, in part, because there

is very little natural history information on Recent spe-

cies that provide insight into how these animals grow and

how their growth is ee by environmental condi-

tions. Subsequently, we have relied heavily upon the

combined works of numerous past investigators to help

us with this decision making process. These researchers

included, but were not limited to: Brunet (1995), de

Boury (1909), Clench and Turner (1950; 1951; 1952),

DuShane (1979: 1988), Gardner (1947), Kilburn (1985),

laa (2003), Robertson (1983a; 1983b; 1993), Weil

et al. (1999), and Woodring (1959).

At the generic and subgeneric levels of classification

many investigators have y widely div ergent opinions about

in which group a specific epitoniid should be placed.

According to Clench and Turner (1950), de Boury's work

on E pitoniidae lk left behind a long list of generic and

subgeneric names with only the types avail: ble for diag-

nostic analysis. This list includes seven generic and 19

subgeneric names (Weil et al., 1999). It was Clench and

Turner's (1950) opinion that de Boury became confused

about the overlapping characteristics of the Epitoniidae

and rather than trying to place them into existing catego-

ries, he established new genera and subgenera for them.

To date, this confusion with ove lapping characteristics

appears to have continued with the list of generic and

subgeneric extant Epitoniid ae alone being expanded to

34: and; 35 names, respective ly (Weil et al., 1999). For our

classification purposes we have decided to adhere, as

closely as possible, to the more conservative phylogenetic

scheme followed by Clench and Turner (1950) rather

than the more rece nntly expanded version used by Weil et

al. (1999) and Nakayama (2004).

With fossils, placement of certain epitoniids into ap-

propriate generic and subgeneric groups has been ham-

pe re va by iC rosion ( { taphonomic degrad: ition). This process

removes surface sculpturing present on living specimens

and exposes subsurface characteristics that are remark-

ably different. In this paper, an example of the impact

erosion has on the external features of a shell is ilhas-

trated with the new species Epitonium conwaiae.

We have tried to survey all the pertinent descriptions

and illustrations of both fossil and Recent « »pitoniids. For

taxonomic Comparison purposes the most significant lit-

erature came from publications dealing with the descrip-

tions of Recent and fossil species collected from the

United States, Central and South America, the northwest

Atlantic, European continent, and the eastern Pacific.

Institutional abbreviations used herein are: USNM:

National Museum of Natural History, Smithsonian Insti-

tution, Washington, DC; UF: Florida Museum of Natu-

ral History (FLMNH), University of Florida, Gainesville;

TU: Tulane University, (formerly housed in New Or-

leans, Louisiana and now housed at the FLMNH):; PRI:

Paleontological Research Institution, Ithaca, New York;

ANSP: T The Academy of Natural Sciences, Philadelphia,

Pennsylvania; and BMSM: The Bailey-Matthews Shell

Museum, Sanibel Island, Florida. Because of privacy

rights of landowners, specific locality data for specimens

described below are not given. However, specific locality

information is available to qualified researchers upon

written request to [email protected].

SYSTEMATICS

Superfamily Janthinoidea Lamarck, 1812

Family Epitoniidae Berry, 1910

Genus Amaea H. and A. Adams, 1853

Type Species: by subsequent designation, Scalaria

magnifica Sowerby, 1544.

Subgenus Scalina Conrad, 1865

Amaea gardnerae (Olsson, 1967)

(Figures 3-9)

Description: Shell large, turriculate; protoconch miss-

ing; nine or ten teleoconch whorls. Spire angle 19.5°,

W hors convex, relatively thin, with cancellate sculpture.

Cancellate pattern created by four spiral cords crossed

by smaller, evenly spaced axial costae: square spaces

within cancellate sculpture possess textured pattern cre-

ated by fine overlapping axial and spiral threads. Suture

deeply impressed. B Basal cord well-defined, surface

slightly elevated and sculptured with thin spiral and axial

hreails: Sculpture below basal disc lacks elevated spiral

cords. Umbilicus absent.

Holotype: USNM 645180, maximum height 47.5 mm,

maximum width 14.4 mm.

Type Locality: McClellan Farm, Calhoun County,

Florida.

Other Material Examined: UF 117045, | shell, Ten-

mile Creek 04 (CA020) (= Tulane University locality TU

51), Clarksville Quadrangle USGS 7.5’ Series (1945),

Calhoun County, Florida, Chipola Formation; UF

117087, 1 shell, locality and formation same as pre-

ceding; UF 91459, 16 shells, Tenmile Creek 03 (CAQOI7)

= Tulane University locality TU 546), Clarksville Quad-

rangle USGS 7.5’ Series (1945), Calhoun County,

Florida, Chipola Formation.

Distribution: Amaca gardnerae is a fairly common

Chipola Formation species. [t is abundant at several col-

G. W. Schmelz and R. W. Portell, 2007

Page 109

Figures 3-9. Amaea gardnerae (Olsson, 1967). 3-4. Apertural and abapertural views of holotype USNM 645150 originally named

Scalina gardnerae Olsson, 1967. Since its original description the specimen has degraded; maximum height 47.5 mm, maximum width

14.4 mm. 5-8. Apertural, lateral, abapertural, and basal views of UF 117045 shown for comparison to holotype (USNM 645150) and

other Chipola and Shoal River formation epitoniid species; maximum height 17.55 mm, maximum width 5.4 mm. 9. Magnified view

of sculpture of teleoconch of UF 117045. Scale bar = 0.6 mm

lection sites along Tenmile Creek but is less abundant

along Farley Creek and the Chipola River. Thus far,

there are no reports of it having been collected along the

Yellow or the Apalachicola rivers

Etymology: Named for Julia A. Gardner, a pioneer

researcher on Chipola Formation mollusks

Discussion: Gardner (1945) reported an epitoniid

fragment from the Chipola Formation which she as-

signed to the genus Scalina. Olsson (1967) later de-

scribed this species as Scalina gardnerae. A more thor-

ough analysis of additional specimens of S. gardnerae

now indicates that it should be placed in the genus

Amaea. DuShane (1988) noted that members of the ge-

nus Amadea are larger than any k nown Scalina and have a

less distinct basal cord. In addition, DuShane observed

that in the genus Amaea, the sculpture above and below

the basal cord is different. Unfortunately, the poor con-

dition of the type specimen described by Olsson makes it

difficult to tell what the sculpturing was like in the vi-

cinity of the basal cord Subsequent specnhnens, however

have revealed that the sculpturing above and below the

basal cord is different in S. gardnerae (Figures 5, 8)

Clench and Tumer (1950), Weil et al. (1999), and

Nakayama (2003) have all placed epitoniids with the shell

sculpture described by Olsson (1967) for Scalina gard-

nerae into the genus Amaea. We are in complete agree-

ment with this placement and have assigned Olsson’s

epitoniid to that genus

Weil et al. (1999) has identified eight subgeneric forms

of the genus Amaea. These subgenera are distinguished

from one another by the types ol sculpture that appeal

above and below the basal cord. Nakayama (2003) in hi:

review of northwest Pacific epitoniids retained six of th«

subgenera listed by Weil et al. (1999). Among thi

subgenera listed for the genus Amaea by Nakay

2003), the present authors have assigned the Chipola

’ ]

Formation species to the subgenus Scalina. According to

Page 110

THE NAUTILUS, Vol. 121, No. 3

Nakayama (2003) and Weil et al. (1999) members of this

subgenus possess convex body whorls with a cancellate

sculpture of spiral cords and axial ribs.

Representatives of the genus Amaea have been re-

ported from a number of other fossil locations. These

include Scala (Opalia) reticulata Martin, 1904, from the

Miocene Calvert Formation of Maryland, Amaea

(Scalina) ferminiana (Dall, 1908) from the Pliocene Es-

meraldas beds of Ecuador, Scala (Acrilla) wiegandi

(Bose, 1910) from Mexico and the Miocene Chagres

Formation of Panama (formerly Canal Zone), Scalina

pseudolerogi (Maury, 1925) from the Pliocene of Trin-

idad, Epitonium (Ferminoscala) manabianum (Pilsbry

and Olsson, 1941), and Epitonium (Ferminoscala) ele uth-

erium (Pilsbry and Olsson, 1941) from the Pliocene

fauna of western Ecuador, Scalina boylae (Olsson, 1967)

from the Pliocene Tamiami Formation of Florida, and

Scalina kendacensis Jung, 1971, from the Miocene Ken-

deace Formation of Carriacou. Amaea ferminiana, origi-

nally described from the Recent of Baja California, is an

offshore species. It ranges from Mexico south to Peru

(Weil et al., 1999). DuShane (1988) considered E.

eleutherium and S. wiegandi to be synonymous with A.

ferminiana and suggested that S. pseudolerogi as well as

some other fossil species of Amaea may be synonymous

with A. ferminiana. Comparison of A. gardnerae with A

ferminiana clearly illustrates that the two species are et

synonymous. Amacea fe rminiana possesses six to nine spi-

ral cords of uniform thickness on the body whorls, while

A. gardnerae possesses four broad primary cords with

fae cords in between. The costae on A. gardnerae are

also less prominent then they are on A. ferminiana. Com-

parison of Amaea mitchelli (Dall, 1896), a Recent west-

ern Atlantic species, to A. gardnerae was also made.

Amaea mitchelli has 6-7 primary spiral cords on the body

whorls (four of which are closely spaced below the whorl

mid-line and two to three that are evenly spaced above)

as compared to A. gardnerae which has four evenly

spaced primary spiral cords.

Genus Cirsotrema Morch, 1852

Type Species:

monotypy.

Scalaria varicosa Lamarck, 1822, by

Cirsotrema chipolanum new species

(Figures 10-14)

Description: Shell large, turriculate, protoconch miss-

ing; seven to eight teleoconch whorls. Spire angle 27°.

Whorls slightly convex, strongly shouldered and joined.

Primary unde rlying sculpture present on te sleoconch

whorls consists of toy elevated vertical ribs. Suture

deep, covered by exte “mal eal Seventeen to 23

sinuous, inclined, feather-like, broad costae on body

whorls. Edges of foliated costae occasionally touch the

preceding ones giving the shell surface a pitted appear-

ance. In other instances foliated costae are slightly sepa-

rated from one another. When feather-like costae sepa-

rate from each other, five slightly elevated spiral cords

seen On body whorls. Spiral cords, and Spaces between

them, possess numerous distinct spiral threads. Thin ver-

tical threads intersect spiral threads, creating faint can-

cellate pattern. Varices formed from accretion of foliated

costae; varices poorly developed and only slightly el-

evated. Apical end of costae with pointed nodes. Surface

of costae pitted with obliquely arranged small holes.

Three or four obliquely arranged, slightly elevated, nar-

row cords traverse each costa. Oblique cords on costae

less pitted than remainder of surface area. Base of each

costa stem-like, attached to a broad, crenulated, basal

cord. Basal disk composite, created by a large outer basal

cord with crenulated edges; a middle circle of narrow

linear pits; and an inner columellar cord with a crenu-

lated edge. Aperture subcircular. Columella short and

arched. Lip margin thickened, pitted in unworn speci-

mens. Less anode d specimens possess a slightly elevated

node on the posterior outer margin of the ‘lip.

Holotype: UF 117088, maximum height 32.9 mm,

maximum width 15.0 mm.

Type Locality: Tenmile Creek 03 (CA017) (= Tulane

University locality TU 546), Clarksville Quadrangle

USGS 7.5’ Series (1945), Calhoun County, Florida,

Chipola Formation,

Paratypes: UF 117089, | shell, locality and formation

same as holotype; UF 91490, 5 shells, locality and for-

mation same as ee UF 84575, 7 shells, Tenmile

Creek 01 (CA002) (= Tulane University locality TU 830),

Clarksville eee USGS 7.5’ Series (1945), Cal-

houn County, Florida, Chipola For mation; UF 95161, 1

shell, Tenmile Creek 04 (CA020) (= Tulane University

locality TU 951), Clarksville Quadrangle USGS 7.5’ Se-

ries (1945), Calhoun County, Florida, Chipola For ma-

tion: UF 85383, 1 shell, Chipola 09 (CAOIS) (= Tulane

University locality TU 547), Clarksville Quadrangle

USGS 7.5’ Series (1945), C Cahous County, Florida,

Chipola Formation; UF 13966, 1 shell, Chipola 03

(CA005), Chipola River (exact collection site unknown),

Calhoun County, Florida, Chipola eye UF

94650, 1 shell, Tenmile Creek 02 (CA003) (= Tulane

University locality TU 70), Altha West Omdniile

USGS 7.5’ Series (1982), Calhoun County, Florida,

Chipola ares UF 84444, 1 shell, Farley on 03

(CA009) (= Tulane Unive rsity locality TU 825), Clarks-

ville See USGS 7.5’ Series (1945), Gaon

County, Florida, C bape “eae om UF 91356, 1 shell,

Chipola O07 (CAOI5) (= Tulane University locality TU

554), Clarksville Guadrangle USGS 7.5’ Series (1945),

Calhoun County, Florida, Chipola Formation; USNM

534489, 1 shell, Tenmile Creek 01 (CA002) (= Tulane

University locality TU $30), Clarksville Quadrangle

USGS 7.5' Series (1945), Calhoun County, Florida,

Chipola Formation; USNM 534490, 2 fragments, locality

sail formation same as pre ceding; U 5 M 534491, 2 frag-

ments, Tenmile Creek 05 (CA021) (= Tulane Unive rsity

locality TU 998), Clarksville Quadr ee USGS 7.5! Se-

ries (1945), ( ‘lhieniyi County, F lorida, G hipola Forma-

tion: USNM 534492. 1 shell, F arley Creek 03 (CA009)

G. W. Schmelz and R. W. Portell, 2007

Figures 10-19.

Cirosotrema chipolanum new species and Cirsotrema dalli Rehder, 1945. 10-13. Cirsotrema chipolanum a

tural, lateral abapertural and basal views of holotype UF 117088; maximum height 32.9 mm

pel

maximum width 15.0 mm. 14.

Magnified view of s¢ ulpture of teleoconch of holotype UF 117088. Scale bar = 2.85 mm. 15-18. Cirsotrema dalli Rehder, 1945

apertural, lateral. abapertural, and basal views of UF 238698; maximum height 32.8 mm and maximum width 13.8 mm. 19. Magnified

view of sculpture of teleoconch of UF 238698. Scale bar = 5.0 mm. UF 238698 live collected off Egmont Key, Pinellas Count

Florida at about 52 m depth

Page 112

THE NAUTILUS, Vol. 121, No. 3

= Tulane Univ ersity locality TU 825), Clarksville Quad-

eee USGS 7.5’ Series (1945), Calhoun County,

Florida, Chipola Formation; USNM 534493, 10 shells,

Tenmile Creek 03 (CAO17) (= Tulane University locality

TU 546 and USGS 2212 “one mile west of Bailey's

Ferry”), Clarksville Quadrangle USGS 7.5’ Series (1945),

Calhousi County, Florida, Chipola ep teen USNM

534494, 8 shells, same locality and formation as preced-

ing,

Distribution: Cirsotrema chipolanum is a fairly com-

mon species. It is locally abundant at several Chipola

Formation collecting sites along the Chipola River and

Tenmile and Farley creeks. The fact that it has not been

reported from the Oak Grove Sand along the Yellow

River or from Chipola Formation sites along the

Apalachicola River may simply reflect the degree of dif-

ficulty collecting these localities at the appropriate peri-

ods of low water level.

Etymology: Named for the Chipola River.

Discussion: The genus Cirsotrema has an extensive

evolutionary history. Sohl (1964) established the genus

Striaticostatum to represent a Cretaceous form of Cir-

sotrema that lacked the faint spiral striations on the body

whorl. By the Eocene the genus Cirsotrema was well

established with numerous species being described from

different localities (Dockery, 1980; Harris and Palmer,

1946; Palmer, 1937). C urrently, among extant forms, two

species of Cirsotrema exist in ‘the western Atlantic (Weil

et al., 1999). These are Cirsotrema dalli Rehder, 1945

and Cirsotrema pilsbryi McGinty, 1940.

Two species of Cirsotrema have also been reported

from lower Miocene Chipola Formation. Olsson (1967)

identified a specimen collected along the west bank of

the Chipola River south of Tenmile Creek as C. dalli and

more recently Duerr (2004) described Cirsotrema cirri-

tum from material collected at several different Chipola

Formation sites. Olsson (1967) stated that after close

examination of the Chipola Formation Cirsotrema he

found practically no differences between it and the ex-

tant species currently living in the western Atlantic. Since

Olsson’s (1967) analy sis of the ¢ Chipola Cirsotrema, over

50 additional specimens have been collected from a va-

riety of Chipola locations. When most of these specimens

were closely scrutinized it became apparent to the

present investigators that the Chipola Formation Cir

sotrema studie d by Olsson (1967) was not C. dalli.

As noted by Clench and Turner (1950), the sculptur-

ing on the body whorls of representé oe. of the genus

Cirsotrema, is extremely complicated. It was suggested

by Clench and Turner, and observed in the present Pure

that two layers of sculpturing are prese nt on the body

whorls of Cirsotrema dalli. There is an outermost one

that consists of foliated costae that may or may not join

each other along their convoluted e dee »s and asec ‘ond: uy

layer of aeinare cd costae and spiral cords. Because of this

dual type of sculpture Clench and Tumer warned against

identifying different species of Cirsotrema from beach

worn specimens.

Often with fossil specimens it is difficult to find shells

that have not been eroded. However, a sufficient number

of well-preserved Chipola Formation Cirsotrema were

discovered which allowed a more detailed comparison

between the fossil form of this genus and its Recent

counterpart. These detailed studies lead to the following

observations:

First, when the underlying sculpture of the Chipola

Formation Cirsotrema was exposed no difference was

discovered between it and the underlying sculpture ex-

hibited by C. dalli.

Second, although varices are present on both the

Chipola Formation Cirsotrema and its Recent counter-

part, the varices on the Chipola Formation species are

clearly not as well developed as those of Recent C. dalli.

In fact, the varices on C. chipolanum are at times so

poorly formed that it is difficult to identify them as va-

rices. In C. dalli the costae are fused together to forma

pronounced, elevated ridge (Figures 15-19) that, in well-

preserved specimens, ee a slightly crenulated margin.

Conversely, with C. C. chipolanum the varices consist of

little more than one or two slightly raised costae (see

Figures 10-13). In addition, even in instances where two

costae are fused together to form a varix, the fusion is

often not complete and a distinct line of demarcation can

be seen.

Third, when a comparison of the number of varices in

relation to the height of the shell was undertaken with

well-preserved specimens of C. dalli and C. chipolanum,

and a least squares regression analysis was done on the

two species, the qeaulis showed a significant correlation

between the height of the shell sel the number of va-

rices with C. dalli (0.922), and a low correlation between

shell height and the number of varices with C. chipo-

lanum (0. 499). Table 1 below provides the statistical re-

sults of this study and a comparison of the number, size

range, average size, varix range, and average number of

varices of the specimens | in the analysis.

As a result of the 1 regression analy sis tw o other differ-

ences between the varices of the two species was also

noted. Although the varices appeared at random intervals

on the body + of both species, in C. chipolanum the

first varix did not appear until after the third teleoconch

whorl, while in C. dalli varices would appear just after

the protoconch. In addition, when the number of varices

Table 1. Results of least squares regression analysis compar-

ing shell he sight with the number of varices found in C. dalli and

C. chipolanum.

C. dalli C. chipolanum

No. of specimens examined 23 16

Correlation coefficient 0.922 0.499

Range of shell height (mm) 5.8-37.5 5.0-44.2

Mean height (mm) 15:31 17.2]

Range of varices 2-27 0-10

Mean no. of varices §.82 4.75

G. W. Schmelz and R. W. Portell, 2007

Page 113

was compared between the different species it was discov-

ered that C. dalli, on average, had significantly more varices

than C. chipolanum [X = 8.82 vs. 4.75].

Table 2 below summarizes the structural differences

in shell morphology between C. dalli and C. chipolanum.

In addition to comparing the differences between the

shape and number of varices between species, the table

also compares the number of teleoconch whorls, the

number and arrangement of costae, the spire angles, the

arrangement of the costae, and the placement of varices

on the body whorls.

Another point worth noting is that C. dalli and C.

chipolanum came from very different environments. Cir-

sotrema dalli lives in cooler continental shelf waters at

depths of 37 to 227 m (Clench and Turner, 1950) while

C. chipolanum thrived in a shallow, warm, tropical, reef

habitat. Although not a great deal is known about how

the environment and food supply impact the growth of

wentletraps [Robertson (1953a) and Weil et fal (1999)],

there has been sufficient studies done on the family Mu-

ricidae (Vokes, 1973) that will allow us to speculate as to

the reasons for the number and structural differences

observed between the varices of C. dalli and C. chipo-

lanum. According to Vokes (1973) varix (axial growth

ridge) development in muricids occurs during resting

stages in shell formation. These stoppages may come

about as the result of food shortages or perhaps as the

result of environmental changes. The weaker develop-

ment and lower number of varices in C. chipolanium may

indicate that this species lived in a rather stable environ-

ment where there were a large eee of prey species

for it to consume. Conversely, dalli has more and

better developed varices ou: stoppages) which may

be due to lack of prey or perhaps increased predation

pressures.

In all probability, C. chipolanum is the ancestral form

of C. dalli. It is the opinion of the investigators that C.

chipolanum probably retreated from the coastal waters

ste the deeper offshore waters during intervening ice

ages where it gradually evolved into the extant, and mor-

phologically ciendlae: C. dalli.

Clench and Turner (1950) did not assign a subgeneric

name to this genus even though de Boury (1909) did.

Table 2. A comparison of the structural differences in shell

morphology between C. dalli and C. chipolanum.

dalli C. chipolanum

No. of specimens examined 23 16

Teleoconch whorls 9-10 7-8

Spire angle 26.5° 27.0°

Costae arrangement touching sometimes

separated

Mean no. of costae 20.6 19.]

Range in costae number 18-23 17-2]

Range in varix number 2-27 0-10

Mean varix number 8.82 4.75

ard

start on 3

body whorl

start on 1**

body whorl

Varix placement

Clench and Turner's reason for not using subgenera is

not stated, but in all probability the authors did not think

that structural differences between members of this ge-

nus were sufficient to warrant their use.

Since Clench and Turner's publication, Weil et al.

(1999) and Nakayama (2003) have assigned a number of

subgenera to the genus Cirsotrema. It is interesting to

note that between these authors there is not complete

agreement as to which subgenera are valid, For example,

ene oon aoa the pees Boreosc sea to de-

Hie: like the W He W eil et al. ( 1999) 1 raised fe sub-

genus Boreoscala to the generic level. In addition, Weil

et al. questions the use of Elegantiscala as a subgenus of

Cirsotrema while Nakayama retained this subgenus and

used it for many of the species of Cirsotrema he de-

scribed from the northwest Pacific.

The present authors have followed the more conser-

vative gee used by Clench and Turner (1950), Ab-

bott (1974), Duerr (2004), and Landau et al. (2006) and

have not nome a subgenus to the new Chipola For-

mation species.

Cirsotrema togatum (Hertlein and Strong, 1951) i

morphologically sionilay to C. chipolanum. Duerr (5004)

considers C. togatum to be a westem cognate of C. dalli.

Cirsotrema togatum has also been reported from the

Pliocene Esmeraldas beds of northwestern Ecuador by

Pitt (1981) and DuShane (1988). Cirsotrema chipolanum

differs from C. togatum in that it has fewer and less

distinct spiral cords on the body whorls and the varices

are much less developed.

Cirsotrema woodringi Olsson, 1967, reported from the

Miocene Gatun Formation of Panama and the Pliocene

Tamiami Formation at Sunnyland, Florida, is similar to

C. chipolanum. The main difference between C. woo-

dringi and C. chipolanum is that C. woodringi lacks the

inclined spiral sculpture that is present on the flat outer

surface of the costae in C. chipolanum.

Cirsotrema cirritum Duerr, 2004

(Figures 20-28)

Description: Shell medium to large. Turriculate.

Much of protoconch missing. Last protoconch whorl

transitions from a smooth surface into wavy axial lamellae

that gradually enlarge into thickened foliated costae.

Spire angle 97° Eight strongly shouldered teleoconch

whorls present. Suture deep. Thirteen crenulated, re-

curved, axial costae present on last body whorl. Apical

ends of axial costae sharp. Costae made up of multiple

wavy lamella that possesses a fine irregular diamond-

shape ‘d pattern that is inclined adaperturally. Costae

separated by wide intercostal spaces. Intercostal spaces

possess five ‘rounded spiral cords. Cords and intervening

spaces have a cancellate sculpture created by overlapping

vertical and horizontal threads. Anterior reflected pro-

jections of costae on last body whorl form a basal ridge

with a crenulate outer margin. Costae on basal ridge not

fused. Varices absent. Columella short and arched. Ap-

erture subcircular. No umbilicus.

Page 114 THE NAUTILUS, Vol. 121, No. 3

Figures 20-28. Cirsofrema cirritum Duerr, 2004. 20-23. Apertural, lateral, abapertural, and basal views of holotype UF 110972;

Jor

mum height 28.5 mm, maximum width 11.8 mm, 24. Magnified view of sculpture of teleoconch of UF 110972. Scale bar = 2.75

5-28. Apertural, lateral, abapertural, and basal views of paratype UF 67746; maximum height 29.6 mm, maximum width 13.4

Note: Arrows point to apparent varices on paratype UF 67746 thus based upon description by Duerr (2004, p-. 154-155)

a key diagnostic feature of no varices, this paratype was mistakenly identified. In ow opinion UF 67746 is representative

/,; ] Fa

Upolanun Tre species

G. W. Schmelz and R. W. Portell, 2007

Page 115

Holotype: UF 110972, maximum height 28.5 mm,

maximum width 11.8 mm.

Type Locality: —Tenmile Creek 01 (CA002) (= Tulane

University locality TU 830), Clarksville os eae

USGS 7.5’ Series (1945), Calhoun County, Floves,

Chipola Formation. Note: Duerr (2004) erroneously

listed the GPS coordinates of 29° 30.05’ N, 85° 11.00’ W

for this locality and provided no datum. These coordi-

nates are approximately 17 km south of Saint Vincent

Island, Florida in the Gulf of Mexico. Additionally, Du-

err (2004) listed the type locality as in the SE1/4 of Sec.

7. It should have read SEI/4 of Sec. 12.

Other Material Examined: UF 112019, paratype,

Farley Creek 03 (CA009) (= Tulane University locality

TU 825), Clarksville Quadrangle USGS 7.5’ Series

(1945), Calhoun County, Florida, Chipola Formation;

UF 67746, paratype, locality and formation same as pre-

ceding; BMSM 15301, paratype, Tulane University a

cality TU 458, Chipola pariew USNM 534499,

shells, Tenmile Creek 03 (CA017) (= Tulane Univ ersity

locality TU 546), C lanksville Deere USGS 7.5’ Se-

ries (1945), Calhoun County, Florida, Chipola oe

tion.

Etymology: The name cirritum is derived from the

Latin cirrus meaning “filamentous” which refers to the

fringed costae of this species.

Discussion: One of the key diagnostic features of C.

cirritum is its lack of varices. However, when comparing

the paratypes of C. cirritum with the holotype, one

paratype, identified as C. cirritum, was discovered to

possess varices that were similar to those observed in C.

chipolanum (see Figures 25-28). This discovery gener-

ated some confusion and resulted in an exhaustive inves-

tigation to determine if only one species of Cirsotrema

(C. cirritum) existed in the Chipola Formation or if there

were two distinct species and a mistake had been made

with the selection of one of the paratypes of C. cirritim.

The conclusion drawn from this analysis was that there

are two different species of Cirsotrema in the Chipola

Formation and that the C. cirritum paratype (UF 67746)

was mistakenly selected.

When examining the different species of the Chipola

Formation Cirsotrema it is easy to see how this mistake

occurred. The varices on C. chipolanum sometimes can

be easily overlooked without careful examination under a

microscope. A summary of differences between C. cir-

ritum and C. chipolanum is found in Table 3

Table 3. A comparison of the structural differences in shell

morphology between C. cirritum and C. chipolanum.

C. cirritum C. chipolanum

Mean varix number 0 4.75

Costae number 13 17-21

Foliated costae widely separated closely packed

Basal ridge costae not fused costae fused

Cirsotrema cirritum is a fairly rare species that has

been obtained from only three fossil localities: one each

on the Chipola River (TU locality 458), Tenmile Creek

(TU locality 951), and Farley Creek (TU locality $25

As Duerr (2004) noted, C. cirritim is a fairly a e

tive species that bears some resemblance to several types

of Recent Indo-Pacific Epitoniidae. Among the compa-

rable Recent species, Duerr (2004) reported were Cir

sotrema plexis Dall, 1925, Cirsotrema fimbriatulum

(Masahito et al., 1971), Cirsotrema rugosum (Kuroda

and Ito, 1961), and Cirsotrema excelsum Garcia, 2003.

Among fossil species C. cirritum is most similar to the

Miocene species Cirsotrema undulatum (Jung, 1965)

from the Paraguana Peninsula, Venezuela. Cirsotrema

undulatum is a medium-size shell with six to eight post-

nuclear whorls that possess thin, widely sp: aced axials

(Jung, 1965). Cirsotrema cirritum differs from C. undu-

latum by having fewer axial costae on the body whorls (13

versus 17-21) rer five spiral cords on each ‘body whorl

versus four.

The presence of two species of Cirsotrema in the

tropical Chipola Formation environment is consistent

with what currently exists in Florida offshore waters to-

day. According to Clench and Tumer (1950), both C

dalli and C. pilsbr yi can currently be found in deep water

off the Florida coast.

Genus Opalia H. and A. Adams, 1853

Type Species: by subsequent designation, Scalaria

australis Lamarck, 1822.

Subgenus Nodiscala de Boury, 1889

Opalia politesae new species

(Figures 29-33)

Description: Shell small, slender; 2.5 smooth proto-

conch whorls, 8 convex teleoconch whorls. Spire angle

20°. Twelve to 13 rounded axial costae on last teleoconch

whorl. Shell surface covered with fine horizontal and

vertical threads giving surface a pitted appearance. Pitted

surface absent on distal surface of axial costae. Costae

terminate at the suture, creating a crenulated ridge. Su-

tures moderately impressed. No basal ridge present. No

varices. Ape rture oblique, subcircular, ioe d by

thick rounded lip. Inner portion of lip encircled by a thin,

unsculptured rim. Outer portion possesses fine ‘vertical

threads that radiate outwards towards periphery. No um-

bilicus. Columella short and arched.

Holotype: UF 114913,

maximum width 1.7 mm.

Type locality: Tenmile Creek 03 (CA017) (= Tulane

University locality TU 546), Clarksville Quadrangle

USGS 7.5’ Series (1945), Calhoun County, Florida,

Chipola Formation.

Paratypes: USNM 534495, USNM 534496 and UF

113897, 1 shell each, protoconchs missing, locality and

formation for each specimen the same as holotype:

USNM 534497, 1 shell, protoconch missing, Tenmile

maximum height 5.5 mm,

Page 116 THE NAUTILUS, Vol. 121, No. 3

Figures 29-38. Opalia politesac new species and Opalia mica new species. 29-32. Opalia politesac; apertural, lateral, abapertural,

and basal views of holotype UF 114913; maximum height 5.5 mm, maximum width 1.7 mm, 33. Magnified view of sculpture of

teleoconch of UF 114913. Scale bar = 0.41 mm, 34-37. Opalia mica; apertural, lateral, abapertural, and basal views ol holotype UF

66077; maximum height 4.0 mm, maximum width 1.5 mm, 38. Magnified view of sculpture of teleoconch of UF 66077. Seale bar

() {

boy

G. W. Schmelz and R. W. Portell, 2007

Page 117

Creek 01 (CA002) (= Tulane University locality TU 830)

Clarksville Quadrangle USGS 7.5! Series (1945), |

ee County, Florida, C Jhipola Bonaaten: UF 67499, :

shells, C dhipol: LOL (MecClelland’s Farm) (CA001) (= Tu

lane Univ ersity locality TU 457), Clarksville Quadrangle

USGS 7.5’ Series (1945), Calhoun County, Florida,

Chipola Formation; UF 114914, 2 shells, locality and

formation same as holotype; UF 114922, 3 shells, Ten-

mile Creek 04 (CA020) (= Tulane University locality TU

951), Clarksville Quadrangle USGS 7.5’ Series (1945),

aaee County, Florida, Chipola Ronnavon

Distribution: Although Opalia politesae is one of the

more common epitoniids found in the Chipola Forma-

tion, its distribution appears limited to the reef paleoen-

vironments found along Tenmile Creek and the Chipola

River.

Etymology: Named for Greta Polites, an avid student

and ballecor of Chipola and Shoal River formation fos-

sils.

Discussion: According to Clench and Turner (1950:

231) members of the genus Opalia have “Spiral sculpture

usually of exce edingly fine incised threads which may or

may not be finely pitted.” In addition, the genus also

possesses relatively low, heavy costae, no aanbilions and

a thick outer lip divided into a thin inner unsculptured

ring and a thicker finely pitted one. In the subgenus

Nodiscala the sutures are crenulated and the basal cord

is absent.

Gardner (1947) reported the subgenus Nodiscala from

Eocene deposits in Australia. However, there is no evi-

dence of the genus Opalia from the Eocene of the

United States. Machel and Dockery (1984) reported a

number of different species of Opalia from the lower

Oligocene Mint Spring Formation of Mississippi. Ac-

cording to Gardner (1947: 578), the subgenus Nodiscala

is * ‘peculiarly eee of the Miocene of central

Europe” and she also noted that several species have

been found in the Pliocene of Italy. Landau et al. (2006)

reported a number of fossil Nodiscala from a wide range

of European localities. These fossils have been found in

deposits that date from the early Miocene to the late

Pleistocene. DuShane (1979) lists Opalia borealis as pos-

sibly coming from Miocene deposits of the northeastern

Pacific region. All of the re maining Opalia studied by

DuShane (1979) came from Pliocene and Pleistocene

formations. In the Caribbean, Maury (1917) reported a

single specimen of Epitonium textuvestitum from

Pliocene deposits in Santo Domingo (Dominican Repub-

lic). The description of this species given by Maury

1917 7) clearly indicates that it belongs to the genus Opa-

lia. Both Campbell ( 1993) and Petuch (1994) have as-

signed this species to the genus Opalia. Campbell's

specimen came from Pliocene deposits in Hampton, Vir-

ginia and Petuch’s specimen was reported from the

Pliocene deposits of the former APAC shell pits in Sa-

rasota. Florida.

Opalia politesae is one of the more abundant species

scacchi. (Hoernes,

of Epitoniidae in the Chipola Formation where it is a

unique representative of this genus. In overall contour

and shape it bears some resemblance to the fossil O.

textuvestitum but it lacks varices and has fewer costae on

the body whorl. Among Recent species O. politesae is

most similar to Opalia burryi, which ‘ranges from south

Florida through the West Indies to Trinidad. However,

O. politesae is a much smaller and more slender species

that has fewer body whorls (S versus 9-11) and fewer

costae on the body whorls (12-13 versus 14) than its

Recent counterpart. The close similarity between O. po-

litesae and O. burryi suggests that O. polite sae is prob-

ably the ancestral form of O. burryi.

Opalia mica new species

(Figures 34-38)

Description: Shell short, stocky; one smooth proto-

conch whorl present, four teleoconch whorls. Nine to ten

costae on last teleoconch whorl. Spire angle 29°. Costae

sharply angulated, forming a node at the periphery of the

body whorl. Shell surface crossed with fine horizontal

and spiral threads that give the surface a pitted appear-

ance. Pitted sculpture covers entire surface of axial cos-

tae, Costae terminate at the suture creating a crenulated

ridge. Sutures moderately impressed. No basal ridge

present. No varices. No umbilicus. Aperture oblique,

subcircular, surrounded: by a thick lip. Inner a of

lip encircled by thin, smooth rim. Outer part of lip pos-

sesses fine vertical threads that radiate outward towards

the periphery. Vertical threads on lip overlapped by fine

threads that give the lip a pitted appearance. Columella

short and arched.

Holotype: UF 66077, maximum height 4.0 mm, maxi-

mum width 1.5 mm.

Type Locality: Shoal River Grotto (WLO004) (= Tu-

lane University locality TU 69A), New Harmony Quad-

rangle USGS 7.5’ Series (1987), Walton County, Florida,

Shoal River Formation.

Paratypes: UF 114924, I specimen, and UF 88160, 3

specimens, locality and formation same as holotype.

Distribution: —Opalia mica is a rare species but thus far

has only been obtained from the type locality.

Name alludes to its diminutive size.

Etymology:

Discussion: Opalia mica is structurally very different

from O. politesae. It is broader and sme ler than the

Chipola Formation species and possesses sharply angu-

lated costae. Gardner (1947) reported fragments of this

species from the Shell Bluff location along the Shoal

River in Walton County, Florida. The most comparable

fossil form to O. mica is the Miocene species Opalia ct.

1856) reported from Austria (Nord-

sieck, 1972). In size and overall form the two species are

very similar; however, O. mica has deeper sutures, lacks

rounded varices, and does not possess the sutural spiral

cords present in the European taxon.

Among extant forms O. mica is similar to O pumilio

Page 118

THE NAUTILUS, Vol. 121, No. 3

morchiana Dall, 1889. Both the Chipola Formation and

Recent species are small and have costae that are sharply

angulated at the periphery of the body whorl. However,

O. mica as varices, has a more acute spire angle (29°

versus 37°), and only has four teleoconch whorls com-

pared to seven for O. pumilio morchiana.

Genus Epitonium Réding, 1798

Type Species: by subsequent designation, Turbo sca-

laris Linnaeus, 1758.

Subgenus Asperiscala de Boury, 1909

Epitonium virginiae (Maury, 1910)

(Figures 39-46)

Description: Shell small, attenuate; 3 smooth, glossy,

protoconch whorls, 6-7 convex teleoconch whorls sepa-

rated by deep sutures. Spire angle 35°. 5-9 costae on last

teleoconch whorl. Costae prominent, blade-like, slightly

oblique with coronate shoulders. Fine spiral threads on

body whorl crossed by faint axial growth line. No varices.

Umbilicus absent. No basal cord. Aperture oval.

Holotype: PRI 3467 (formerly in Cornell University

collection), maximum height 3.75 mm, maximum width

1.5 mm.

Other Material Examined: UF 95695, 1 shell,

Chipola 13 (CA027) (= Tulane University locality TU

458), Clarksville Quadrangle USGS 7.5’ Series (1945),

Calhoun County, Blond. Chipola Formation; UF

89251, 4 shells, Chipola 28 (CA06G) (= Tulane Univer-

sity locality TU 545), Clarksville Quadrangle USGS

7.5’ Series (1945), Calhoun County, Florida, Chipola

Formation: UF 103784, 1 shell, Tenmile Creek 06

(CA023) (= Tulane Univ ersity locality TU 456), Clarks-

ville Quadrangle USGS 7.5" Series oe Calhoun

County, Florida, Chipola For ition; UF 72341, 1 shell,

Shoal River Grotto (WL004) (= Tulane Univ se locality

TU 69A), New Harmony Olenge USGS 7.5’ Series

(1987), Walton County, Florida, Shoal River Formation.

Type Location:

Florida.

Bailey's Ferry, Calhoun County,

Distribution: —Epitonium virginiae is one of the more

common species of Epitoniidae found in the Chipola

Formation. It is also found in the Shoal River Formation

where it appears to be extremely rare. A total of 59 speci-

mens of E. virginiae were examined from ten ( thipola

Formation collection sites in Calhoun and Liberty coun-

ties. The only Shoal River example of E. virginiae was

obtained from UF loc ality WI ree = TU 69A) in Walton

County.

Etymology: Unknown.

Discussion: Members of the genus Epitonium are

small, thin, generally slender turriculate shells with a

wide range of different sculptured characteristics. Some

have body whorls that are attached while others do not.

\ basal ridge may be present or absent, the costae may

be thin and blade -like or thick and rounded, and spire al

sculpturing may or may not be present. This high degree

of structural variability within the genus has Caused: re-

searchers to organize its members into numerous sub-

genera.

Epitoniidae with blade-like costae, spiral thread-like

cords on the body whorls, and an absence of a basal ridge

were assigned to the subgenus Asperiscala by de Boury

(1909). Members of the subgenus Asperisc cala have been

reported from deposits as old as Cretaceous (Wade,

1926).

Clench and Turner (1952) kept the subgenus As-

periscala and assigned to it all epitoniids with spiral cords

and either blade or cord-like costae. DuShane (1979: 91),

because of “Certain morphological differences from

those of Epitonium s.s.”, elevated Asperiscala to full ge-

neric rank when she described the family Epitoniidae’ in

the northeastern Pacific. According to ‘Kilburn (1985),

however, the type species of Asperiscala is not represen-

tative for this taxon. Kilburn reported that the type spe-

cies of Asperiscala described by de Boury (1909) had

cancellate sculpture. abe, Kilburn assigned

epitoniids with only spiral sculpture to the subgenus

Parviscala. Weil et al. (1999) and Nakayama (2003) re-

tained Asperiscala as a subgenus but limited its use to

epitoniids that resembled Parviscala that have an open

umbilicus and sutures with peaked costae. Herein, we

have retained the use of Asperiscala in the sense of

Clench and Turner (1952), pending resolution of the

previo cited above. But unlike its use in Weil et al. and

Nakayama, Asperiscala is herein used to represent mem-

ees of the genus Epitonium that have nee sculpture

that may or may not be intersected with faint axial

threads or narrow cords.

A more detailed analysis of larger specimens of E.

virginiae revealed the presence of fine vertical lines that

intersect the slender spiral threads on the body whorls

(Figure 46). The absence of this feature in Maury’s de-

scription was more than likely due to the small size of the

specimen she examined.

Of all the Chipola Formation species of Epitonium

examined, only EF. virginiae was encountered in the

younger Shoal River Formation. It is apparently a rare

species in this unit given that numerous collecting trips

to several different Shoal River Formation sites by dif-

ferent collectors have yielded liane one specimen. Close

examination of this specimen revealed that its shell struc-

ture is fundamentally the same as that of the Chipola

Formation species.

Several fossil species similar to E. virginiae have been

collected from different localities including the Carib-

bean, Central and South America, and Spain. Epitoniwm

(Asperiscala) venezuclense (Weisbord, 1962) from the

upper Miocene Mare Formation of northern Venezuela

comes closest in overall morphological characteristics to

E. virginiae, Both species have prominent blade-like,

slightly oblique costae with coroneted shoulders, and

both have spiral threads in the interspaces between the

axial costae that are crossed by fine axial filaments. The

G. W. Schmelz and R. W. Portell, 2007

Figures 39-46. Epitonium virginiae (Maury, 1910). 39-40. Apertural and abapertural views of holotype PRI 3467; maximum

ight 3.75 mm, maximum width 1.5 mm. 41. Magnified view of sculpture of teleoconch of PRI 3467. Scale bar = 1.0 mm, 42-45.

\pertur il, lateral, abapt rtural, and basal views of UF 95695; maximum height 6.9 mm, maximum width 2.45 mm. 46. Magnified vi

ot sculpture of teleoconch of UF 95695. Scale bar = 0.55 mm

Page 120

THE NAUTILUS, Vol. 121, No. 3

two species differ from one another in that E. virginiae

has more impressed sutures, the costae of E. virginiae do

not become obsolescent near the posterior suture, and

the angle of the spire is slightly wider in E. virginiae (35°

versus 32°).

Other fossil species similar to E. virginiae include Epi-

tonium loripanum Pilsbry and Olsson, 1941, from the

Pliocene of Ecuador, Epitonium amosbrowni. Pilsbry,

1921. from the Miocene of the Dominican Republic,

Epitonium ct. gabbi (de Boury) Woodring, 1959, from

the Gatun Miocene of Panama (Canal Zone), and Epi-

tonium muiricatoides (Sacco, 1891) from the early

Pliocene of Spain. The Chipola Formation species di ffers

from E. amosbrowni, which has 16 axial costae on the last

whorl versus 8-9 for E. virginiae, from E. cf. gabbi,

which has more pronounced and numerous spiral

threads on its body whorls, and E. muiricatoides, which

lacks spiral cords on the body whorls.

A close comparison of E. loripanum with E. virginiae

suggests that they may be the same species. Bok have

the s same body shape, the presence of faint spiral threads

on the body whorls, the same number of teleoconch

whorls, thin ribs with coronated shoulders, and lack an

umbilicus.

Among extant species, E. virginiae is most similar to

Epitonium denticulatum (Sowerby, 1844). However, E.

virginiae is more slender and has fewer axial costae.

Epitonium incomitatum new species

(Figures 47-51)

Description: Shell small, thin, turriculate. Protoconch

missing; 6 strongly convex teleoconch whorls separated

by deep sutures. Spire angle 31°. Thin, low, reflected

costae on last teleoconch whorl number ing 24. Costae on

each succeeding whorl slightly offset from the one above.

Numerous spiral cords on body whorl. No varices. No

basal ridge. Umbilicus present. Outer lip of aperture

thin. Aperture suboval.

Holotype: UF 91452, maximum height 5.0 mm, maxi-

mum width 2.2 mim.

Tenmile Creek 03 (CAQ17) (= Tulane

Clarksville Ouadenel

Calhoun County, Florida,

Type Locality:

University locality TU 546),

USGS 7.5' Series (1945), ¢

Chipola Formation.

Distribution: Only known from the type locality.

Etymology: The name is derived from the Latin word

incomitatus meaning unaccompanied or alone. It refers

to the unique specimen (holotype).

Discussion: = Epitonium incomitatum is an extreme ly

fragile shell. In shell sculpture it is similar to Parvise ala,

a subgenus retained by Weil et al. (1999) and N: tkayame U

2003). Gardner (1947) re porte da fragment of this shell

in the ( thipola Formation and assigned it to the subgenus

Crisposcala. A fragment similar to E. incomitatum was

also reported from the upper part of Gatun Formation in

Panama (Canal Zone). Woodring (1959) tentatively iden-

tified this species as Epitonium rushii (Dall, 1889). Weil

et al. listed E. rushii as synonym of Epitonium striatis-

simum Monterosato, 1878. Among fossil western Euro-

pean species E. incomitatum is most similar to Epito-

nium pulchellum (Bivona, 1832) which has been re-

ported from the middle Miocene in Italy (Cavallo and

Repetto, 1992). Epitonium incomitatum differs from its

European counterpart by having less elevated varices

and less prominent spiral cords on the body whorls.

Among extant species, E. incomitatum is most similar

to E. striatissimum, a rare species found in shallow water

off Cape Hatteras. se ae incomitatum differs from

E. striatissimum in that the body whorls are more in-

flated (the angle formed with the spire is 31° versus 25

for E. striatissimum) and the spiral cords on the body

whorl are much broader and less numerous.

Epitonium incomitatum also bears some resemblance

to Epitonium multistriatum Say, 1526, a species that

ranges from Massachusetts to Texas. It differs from this

species by ie: more numerous costae on the last

body whorl (25 versus 16-19) and by having broader

spiral cords on the body whorls. In addition, in E. in-

comitatum the costae are less abundant in the early

whorls, while E. multistriatum the costae are more nu-

merous in the early whorls (as many as 43 in some speci-

mens).

Epitonium regina new species

(Figures 52-56)

Description: Shell small, turriculate: three smooth,

glossy protoconch whorls, eight slightly angular body

whorls separated by a moderately deep suture. Spire

angle 23°. Eleven narrow, low, slightly reflected costae

on last teleoconch whorl. Costae occasionally offset with

costae on preceding whorl. Numerous faint, spiral cords

on body whorls intersected by faint spiral threads. No

basal cord. No umbilicus. No varices. Aperture thin, sub-

OV. al.

Holotype: USNM 534487, maximum height 6.1 mm,

maximum width 2.1 mm.

Farley Creek 03 (CAO009) (= Tulane

Clarksville Dlsdmel

Calhoun County, Florida,

Type Locality:

University locality TU 825),

USGS 7.5’ Series (1945),

Chipola Formation,

Paratype: UF 67498, I specimen, Chipola OL (CAOOL)

(McClelland’s Farm) (= Tulane University locality TU

457), Clarksville Quadrangle USGS 7.5’ Series (1945),

Calhoun C ounty, F lorida, € Chipola Formation.

Distribution: Collected only from the type locality.

Etymology: Named from the Latin word regina,

meaning queen, an allusion to the stately appearance of

this species.

Discussion: regina bears a

slight resemblance to Epitonium obliquum (Sowerby,

1$47), however E. regina has a cancellate pattern be-

Among Recent species, E.

G. W. Schmelz and R. W. Portell, 2007 Page 121

Figures 47-56. Epitoniwim incomitatiwm new species and Epitonium regina new species. 47-50. Epitonium incomitatum, ape

tural, lateral, abapertural, and basal views of holotype UF 91452; maximum height 5.0 mm, maximum width 2.2 mm. 51. Magnified

view of sculpture of teleoconch of UF 91452. Scale bar = 0.41 mm. 52-55. Epitonium regina: apertural, lateral, abapertural, and basal

vs of holotype USNM 534487: maximum height 6.1 mm, maximum width 2.1 mm. 56. Magnified view of s¢ ulpture of teleoconch

of USNM 5344587. Scale bar = 0.65 mm

Page 122

THE NAUTILUS, Vol. 121, No. 3

tween costae on the body whorls, lacks elevated costae

near the sutures, and has no umbilicus.

Among fossil species E. regina bares some resem-

blance to Epitoniwm smithfie Ider sis Mansfield, 1929 and

Epitonium dupliniana (Olsson, 1916). Epitonium smith-

fieldensis was reported from the Pliocene Yorktown For-

mation of Virginia. Like the Chipola Formation species it

is ornamented with marginally reflected slender varices

and the number of varices on the last body whorl is 12.

However, E. smithfieldensis does not have any spiral

sculpture and its varices are united at the suture. Epito-

nium dupliniana was reported from the middle Pliocene

Duplin Formation of North Carolina (Olsson, 1916).

Like E. regina, it has a thin shell with low varices and

spiral sculpturing on the body whorls that is intersected

by vertical threads. It differs from the Chipola Formation

species in that the varices on E. regina are broader and

more cord-like than they are in E. dupliniana. Also, the

upper shoulder on each of the varices of E. dupliniana

has a small hook-like projection, a feature not et on

the varices of E. regina.

Subgenus Epitonium Réding, 1798

Type of Subgenus: = Tubo scalaris Linnaeus, 1758 by

subsequent designation, Suter, 1913.

Epitonium contwaide new species

(Figures 57-66)

Description: Shell small, attenuate; 3.5 smooth proto-

conch whorls. Spire angle 30°. Six-and-a-half moderately

convex teleoconch whorls separated by a depressed su-

ture. Seven to nine low, moderately broad, T-shaped

costae on last body whorl. Costae gradually increase in

width as aperture ‘approache .d. Costae angled at shoul-

ders. Body whorls smooth but eroded specimens exhibit

numerous, fairly broad spiral cords on body whorls. No

basal cord, umbilicus, or varices Aperture suboval, outer

lip slightly expanded and thicketie d.

Holotype: UF 113894, maximum height 3.2. mm,

maximum width 1.2 mm.

Type Locality: Shoal River Grotto (WLO04) (= Tu-

lane University locality TU 69A), New Harmony Quad-

rangle USGS 7.5’ Series (1987), Walton County, Florida,

Shoal River Formation.

Paratypes: UF 67208, 9 shells, Shell Bluff (

(WL002) (= Tulane University locality TU 69), New ae

mony Quadrangle USGS 7.5’ Series (1987), Walton

County, Florida, Shoal River Formation; UF 89549, 1

same as preceding; UF

117092, 2 shells, locality and formation same as preced-

shell, locality and formation s

ing: UF $9638, 29 she lls, locality and formation same as

preceding; UF 72340, 1 shell, locality and formation

same as holotype; UF 88170 locality and formation same

as holotype

Distribution: Epitonium contwaiae is known only from

the type locality and from the Shell Bluff on Shoal River.

Etymology: Named for Wendy Conway a long time

field associate of the authors and an avid eallector of

Chipola and Shoal River formation fossils.

Discussion: —Epitoniwm with and without an umbilicus

and possessing smooth body whorls and no basal cord

were assigned to the subgenus Epitonium by Clench and

Turner (1951). Later researchers. including Weil et al.

(1999) and Nakayama (2003) subdivided Epitonium with

smooth body whorls into a number of subgenera. For

example, t those with smooth body whorls, no umbilicus,

and peaked costae were assigned to the subgenus Hir-

toscala, while Epitonium \y with smooth body whorls,

peaked costae and a slit-like umbilicus were placed into

the subgenus Lamelliscala. Additionally, those with

smooth body whorls, no umbilicus, and thick costae were

assigned to the subgenus Nitidiscala.

Although probably in the minority, we find the need

for splitting out the genus Epitonium into so many ad-

ditional subgenera based upon minor morphological dif-

ferences probably counterproductive to the establish-

ment of a more realistic classification system. Clench and

Tumer (1951) wamed researchers about this problem

when they reported the difficulties de Boury experienced

with his attempts to split out different members of the

family. Rather than add to the confusion that exists with

this taxonomic group, we have reverted to using the

broader subgeneric name of Epitonium as defined by

Clench and Turner (1951).

When compared to Recent species, non-eroded speci-

mens of Epitonium conwaiae (Figures 57-61) are most

similar to Epitonium humphreysii Kiener, 1838, which

ranges from Cape Cod Massachusetts south to Florida

(excluding the Florida Keys) and into the Gulf of Mexico

from Cape Romano to Texas. Fossil specimens of E.

humphreysii have also been reported from upper Mi-

ocene deposits of the Entrerriense Formation of the

Chubut Province, Argentina (Brunet, 1995). Both the

Shoal River Formation species and its Recent and fossil

counterpart have flattened costae which are variable in

width, both lack sculpturing on the body whorls, and

each species has a similar number of costae on the last

body whorl. The Shoal River Formation species differs

from E. humphreysii by having more deeply impressed

sutures, more angular costae on the shoulder of the body

whorls, and much smaller ave rage size.

In eroded specimens of E. conwaiae (Figures 62-66),

the body whorls reveal a sculpture of broad spiral cords

most similar to Epitonium championi Clench and

Turner, 1952, a rare epitoniid that inhabits intertidal and

near-shore waters from Cape Cod to North Carolina.

Both the Shoal River Formation form and E. championi

have flattened, cord-like costae which are variable in

width and both have spiral sculpturing which consists of

numerous flattened spiral cords. The Shoal River For-

mation species, however, differs from E. championi by

having more deeply impressed sutures, a greater number

of costae on the last body whorl (11 versus S or 9) and a

G. W. Schmelz and R. W. Portell, 2007

Figures 57-66. = Epitonium conwaiae new species. 57-60. Apertural, lateral, abapertural, and basal views of holotype UF 113594;

iaximum height 3.2 mm, maximum width 1.2 mm. 61. Magnified view of sculpture of teleoconch of UF 113894. Scale bar = 0.7

mm. 62-65. Apertural, lateral, abapertural, and basal views of paratype UF 72340; maximum height 4.2 mm, maximum width 2.0

61. Magnified view of s« ulpture of teleoconch of UF 723

AQ. Scale bar = 1.26 mm. UF 72340 is an eroded specimen figured

here to compare sculptural differences between it and unworn holotype UF 113894

greater angle between the spire and the outer shoulders

of the shell (30° versus 20

Among fossil species, well-preserved specimens of E

conwwaiae are most similar to Epitonium boltoni Gardner

1948, from the Pliocene Tar River deposits in North

Carolina Like E. conwaiae, E boltoni possesses smooth

spiral whorls, has thickened slightly raised costae an

lacks an umbilicus and basal cord. However, E. conwaia

differs from E. boltoni by having more impressed su

tures, more angular costae on the dorsal surface of each

body whorl and fewer numbers of costae on the last bods

whorl (9 versus 12

Page 124

THE NAUTILUS, Vol. 121, No. 3

The eroded form of FE. conwaiae also bears some simi-

larities to Epitonium alaquaense collected and described

by Mansfield (1935) from the late Miocene strata of

Vaughan Creek in Walton County, Florida. Gardner

(1947: 577) reported collecting a partial specimen from

Walton County from “a horizon that was slightly higher

than the typical Shoal River formation.” These i investiga-

tors were not able to locate Gardner's shell fragment but

were able to borrow Mansfield’s holotype (USNM

373149) for comparative purposes (see Figures 67-68).

Unfortunately, the holotype was broken, which made the

comparative study a bit more difficult than expected.

Nevertheless, examination of the shells showed that E.

conwaiae differs in several ways from E. alaquaense. Epi-

tonium conwaiae is more slender than E. alaquaense and

has 3.5 nuclear whorls versus 2.5 for E. alaquaense and

the number axial ribs on the post-nuclear whorls on E.

conwaiae range from 7-9 while the number of axial ribs

on E. alaquaense ranges from 9-12. Current evidence

suggests that that E. alaquaense is not found in the

younger Shoal River Formation since none of the speci-

mens examined were similar to Manstield’s shell.

The eroded form of E. conwaiae is also similar to the

fossil species Epitonium santodomingonum Pilsbry,

1921, from the Pliocene beds of Santo Domingo (Do-

minican Republic) and Epitonium antillarum fe Boury,

1909) from Pliocene beds in Virginia and North Carolina

(Gardner, 1948) and Florida (Olsson and Harbison,

1953). Like E. conwaiae, both fossil Pliocene species pos-

sess numerous spiral cords on the body whorls, lack a

basal ridge, and both have low, well de veloped costae on

the body whorls which increase in width near the aper-

ture. Epitonium conwaiae differs from E. santodomin-

gonum by having fewer costae on the last body whorl (9

versus 18) and an absence of varices on the last whorl of

the shell. Epitonium antillarum differs from E. conwaiae

by possessing fewer varices (7-9 versus 10-13), fewer

teleoconch whorls (6 versus S—9), thinner costae on the

body whorls, a thinner lip surrounding the aperture, and

a more acute spire angle (22° versus 30°). It should be

noted that E. antillarum is no longer considered a valid

species. In 1909, de Boury assigned the species antil-

larum to the epitoniid Sc alaria turricula Sowe rby, 1844.

However, the shell Sowerby (1844) described had al-

ready been named by 7’ nee (1842) as Sealaria can-

deana. Subsequently, Clench and Turner (1952) rectified

de Boury’s mistake by recognizing E. antillarum as Epi-

tonium candeanum. Of Favithe yr note, the shell described

by Clench and Turner (1952) as E. candeanum does not

fit the description of E. antillarum given by Gardner

1948) and later listed by Olsson and Harbison (1953) in

their treatise on Pliocene Mollusca of Southern Florida.

\ccording to Clench and Turner (1952), E. candeanum

has thinner and more numerous costae (18-25 versus

10-13) on the body whorls than does the species de-

scribed by Gardner (1945) as £. antillarum. In all prob-

ability the Epitonium described by Gardner (1948) is a

new spe cies. It is beyond the scope of this paper to rec-

tify this error and any effort to do so is being left to the

work of future investigators.

Epitonium hoerleae new species

(Figures 69-73)

Description: Shell medium height, sturdy, turriculate;

3.5 smooth, glossy, protoconch whorls. 7.5 convex teleo-

conch wars. separated by deep suture. Spire angle 24°.

Eight to nine thin costae on last body whorl. ‘Costae

slightly reflected backwards. At the whorl shoulder cos-

tae are slightly expanded and form a cusp-like node. Cos-

tae connected to one another at the suture, forming an

oblique angle to the shell’s central axis. Extremely ant

spiral thivends on glossy body whorls. No umbilicus. No

basal cord. Aperture suboval. Outer lip of aperture thin

and reflected backwards.

Holotype: USNM 534488, maximum height 8.8 mm,

maximum width 3.2 mm.

Type locality: Tenmile Creek 03 (CAQ17) (

University locality TU 546),

USGS 7.5’ Series (1945),

Chipola Formation.

Paratypes: USNM 534498, 1 shell, locality and forma-

tion same as holotype: UF 84579, protoconch missing

and aperture broken, Tenmile Creek 01 (CA002) (= Tu-

lane University locality TU 830), Clarksville Quadrangle

USGS 7.5’ Series (1945), Calhoun County, Florida,

Chipola Formation; UF 101862, 1 shell, protoconch

missing and aperture broken, Tenmile Creek 13 (CAQ58)

(= Tulane University locality TU 1097), Clarksville

Quadrangle USGS 7.5’ Series (1945), Calhoun County,

Florida, Chipota Formation; UF 99083, 1 shell, Chipola

23 (CA037) (= Tulane University locality TU 711),

Clarksville ae USGS 7.5’ Series (1945), Cal-

houn County, Blonde: C eae Formation; UF 94317, 2:

shells, Chipola 13 (CA027) (= Tulane University locality

TU 458), Clarksville Gundiancle € USGS 7.5’ Series

(1945), Calhoun County, Florida, Chipola Formation.

= Tulane

Clarksville Quadrangle

Calhoan County, Florida,

Distribution: Epitonium hoerleae is a ve ry common,

widely distributed species. It has been found at the type

locality of Tenmile Creek as well as along F Farley Creek

and the Chipola River.

Etymology: The species was named in honor of Shir-

ley Hoerle, one of the pioneer re searchers of C hipola

Formation mollusks.

Discussion: Clench and Turner (1951) placed e Saat

niids that possessed glossy whorls and extremely faint

spiral threads into the subge snus Epitonium. This classi-

fication has been retained with E. hoerleace.

Among recent species E. hoerleae is most similar to

Epitonium foliaceicostum VOrbigny, 1842, which inhab-

its offshore waters in the Caribbean and along Florida's

east coast. Both the fossil species and its Recent coun-

terpart have glossy body whorls with faint spiral threads,

approximately the same number of costae on the last

body whorl (S—9 versus 7—S), and a slightly reflected

G. W. Schmelz and R. W. Portell, 2007 Page 125

Figures 67-73. Epitonium alaquaense Mansfield, 1935, and Epitonium hoerleae new species. 67-68. Epitonium alaquaens«

ipertural and abapertural views of holotype USNM 373149 shown for comparison to E. conwaiae. 69-72. Epitonium hoerl

ypertural, lateral, abapertural, and basal views of holotype USNM 534488; maximum height $.8 mm maximum width 3.2 mm

Maenified view of s« ulpture of teleoconch of USNM 534488. Scale bar 0.95 mm

od.

Page 126

THE NAUTILUS, Vol. 121, No. 3

aperture lip. Epitonium hoerleae differs from the Recent

shell by having a greater number of nuclear whorls (3.5

versus 1.5), less alee ated costae and a more slender body

configuration.

A comparison of E. hoerleae with a number of fossil

species shows that it most closely resembles Epitonium

fargoi Olsson and Harbison, 1953, which was described

from the Plio-Pleistocene Caloosahatchee Formation of

south Florida as well as Epitonium proximus (de Boury,

1890) which was described from the early Pliocene of

France and the middle Pliocene of England (Harmer,

1920-1925). Both E. hoerleae and E. fargoi have convex,

glossy, body whorls, approximately the same number of

costae on the last body whorl (§ versus 9), the costae are

reflected backwards, and both have a small cusp-like

node on the upper shoulder. Epitonium hoerleae differs

from E. fargoi in that it has a more expanded reflected

outer lip, lacks a thickened cord on the inner lip, its

costae are thinner and more blade-like and the spire

angle in E. hoerleae is less acute (24° versus 20°) than its

Pliocene counterpart. The European species, E. proxi-

mus, differs from E. hoerleae in that it is larger and more

slender than its American counterpart and has more va-

rices on the body whorl (11-13 versus S—9).

Epitonium kallistos new species

(Figures 74-78)

Description: Shell small, turriculate; 3.5 smooth,

glossy, protoconch whorls, 6.5 convex teleoconch whorls

separated by a deep suture. Six moderately thickened,

slightly recurved costae on last body whorl. Spire angle

25°. Costae connected to one another at suture forming

an obliq ue angle with the central axis. Costae lack a cusp-

like bode on the shoulder of body whorl. Body whorls

smooth and glossy. No umbilicus; basal cord absent. No

varices. Aperture suboval, lip slightly thickened and re-

curved.

Holotype: UF 44614, maximum height 6.8 mm, maxi-

mum width 2.3 mm.

Type Locality: Tenmile Creek 02 (CA003) (= Tulane

University locality TU 70), Altha West Quadrangle

USGS 7.5’ Series (1982), Calhoun County, Florida,

Chipola Formation.

Distribution:

the type locality.

Epitonium kallistos is known only from

Etymology: Name is derived from the Greek word

kallisto meaning most beautiful.

Discussion: Among Recent and fossil epitoniids, E

kallistos is most similar to Epitonium unifasciatum (Sow-

erby, 1844), a species that today r anges from southern

Florida to the Lesser Antilles (Clench and Turner, 1951)

Epitonium unifasciatum has also been reported from the

upper Miocene deposits of the Entrerriense Formation

Argentina (Brunet, 1995). Like its

Recent and fossil counte rpart, E kallistos has smooth,

of Chubut Province

shiny, convex whorls, the aperture 1s suboval, there is no

basal ridge, and the outer lip of the aperture is reflected

backwards. Epitonium kallistos differs from E. unifascia-

tum by having fewer costae on the last body whorl (6

versus 7 — 9), a greater spire angle (25° versus 20°), more

deeply impressed sutures, and the costae on the body

whorls are more elevated.

Subgenus Gyroscala de Boury, 1887

Epitonium vokesae new species

(Figures 79-83)

Description: Shell small, turriculate; three smooth,

glossy, bulbous protoconch whorls, 5.5 moderately con-

vex ieee whorls. Suture moderately impressed.

Spire angle 27°. Eight to nine slightly raised, blade-like

costae on boty whorl. Costae sinuous, not joined at su-

ture with costae on preceding whorl. Distinct, narrow,

basal ridge present. No varices. No umbilicus. Outer lip

missing.

Holotype: UF 113595, maximum height 4.3 mm,

maximum width 1.6 mim.

Type Locality: Tenmile Creek 04 (CA020) (= Tulane

University locality TU 951), Clarksville Quadrangle

USGS 7.5’ Series (1945), Calhoun County, Florida,

Chipola Formation.

Paratypes: UF 114957, 1 shell, Tenmile Creek 03

(CAO017) (= Tulane University locality TU 546), Clarks-

ville Quadrangle USGS 7.5’ Series (1945), Calhoun

County, Florida, Chapels haiti, UF 99136, 1 shell,

spire missing, Chipola 23 (CAQ37) (= Tulane University

locality TU 711), Clarksville Quadrangle USGS 7.5’ Se-

ries (1945), Calhoun County, Florida, Chipola Forma-

tion.

Distribution: Epitonium vokesae has only been col-

lected along Tenmile Creek and the Chipola River.

Etymology: Named in honor of Dr. Emily Vokes, a

leading researcher of the family Muricidae and a mentor

to the many who have studied Chipola Formation fossils.

Discussion: Epitonium of the subgenus Gyroscala

possess a smooth protoconch of about three whorls, high

axial lamellae, gently convex smooth body whorls, a well

defined, thin basal cord and a thickened peristome (Kil-

burn, 1985).

Thiele (1929) and a number of other European work-

ers have made Gyroscala subordinate to Cirsotrema.

Clench and Turner (1951) and Abbott (1974) retained de

Boury’s (1887) taxonomic classification of Gyroscala as a

subgenus of Epitonium. Kilburn (1985) followed the lead

of Australian and Japanese malacologists and accorded

Gyroscala full generic status. According to Nakayama

(2003), the presence of a basal cord in Gyrose ala justifie s

raising this epitoniid to generic level. Once Gyroscala

was raised to generic status, a number of investigators

suggested establishing subgenera. Kilburn, for example,

suggested two subgenera (Boreoscala and Circuloscala),

while Nakayama suggested three (Fragiliscala, Pomi-

scala, and Circuloscala). As noted earlier, Nakayama al-

G. W. Schmelz and R. W. Portell, 2007 Page 127

Figures 74-83. Epitonium kallistos new species and Epitonium vokesae new species 74-77. Epitonium kallistos apertural lateral

ibapertural, and basal views of holotype UF 44614; maximum height 6.8 mm, maximum width 2.3 mm. 78. Magnified view of

sculpture of teleoconch of UF 44614. Scale bar = 0.51 mm 79-82. Epitonium vokesat apertural lateral abapertural and basal

of holotype UF 113898; maximum height 4.3 mm, maximum width 1.6 mm. 83. Magnified view olf sculpture of teleoconch of UI

113898. Scale bar = 0.37 mm

Page 125

THE NAUTILUS, Vol. 121, No. 3

ready used Boreoscala as a subgenus for Cirsotrema. To

further complicate the issue Weil et al. (1999) elevated

Gyroscala to the generic level but avoided using any

subgenera. Until a more comprehensive study has ‘Ge en

done with Gyroscala, we have decided to retain Gyro-

scala as a subgenus of Epitonium as suggested by de

Boury and retained by Clench and Turner.

Representativ es of the subgeneric group Gyroscala

are not very common. There are three reported living

species of the subgenus Gyroscala in the western Atlan-

tic. Epitonium late llosum Lamarck, 1822, is found from

Lake Worth, Florida, to the Lesser Antilles, as well as

from France to South Africa. Recently, Garcia (2002),

reported E. lamellosum from the Indo-Pacific to Califor-

nia. Epitonium rupicola Kurtz, 1860, is found from Prov-

incetown, Massachusetts, in the Atlantic to the Texas

coast in the Gulf of Mexico, whereas Epitonium xenicima

Melville and Standen, 1903, is a circum-global species

(Garcia, 2006). None of the Recent western Atlantic spe-

cies appear similar to the Chipola Formation species

which is much more slender, lacks varices on the last

whorl, has far fewer costae on the body whorls, and pos-

sesses a very distinctive, bulbous protoconch.

There are three fossil species similar to E. vokesae.

One is Epitonium aciculum (H. C. Lea, 1843) which,

according to Campbell (1993), was misidentified by

Gardner (1948) as E. pratti. The species comes from the

Pliocene Yorktown Formation in Virginia and North

Carolina as well as the Plio-Pleistocene Waccamaw For-

mation in North Carolina. Epitonium vokesae is more

slender than the Virginia and North Carolina species. In

addition, it has far fewer costae on the body whorl (9

versus 16-25), The other two fossil species similar to E.

vokesae are an unnamed specimen from the Miocene

Chagres Formation of Panama (Woodring, 1959) and

Epitonium magnolianum (Olsson, 1916) from the lower

to middle Pliocene deposits from North and South Caro-

lina and Georgia. Both the Chipola Formation and

Panama fossil species lack spiral sculpturing, have low

sinuous costae on the body whorls, and a suppressed

basal disk. The Chipola Formation species differs from

the Panama fossil in that it is much more slender and has

far fewer costae on the body whorl (9 versus 21). Epito-

nium vokesae is also a much more slender species than E.

magnolianum and lacks varices.

ACKNOWLEDGMENTS

We extend a special note of thanks to Burt Hayes, Cecil

Sexton, and William Tatum (Calhoun County, Florida)

and Robert Larson (Walton County, Florida) for allowing

access to collect on their properties, Thomas Waller and

Warren Blow (USNM), Bushra Hussani (AMNH), Paul

Callomon (ANSP), Warren Allmon and Jonathan Hen-

dricks (PRI), and Gustav Paulay and John Slapcinsky

FLMNH provided specimen loans or access to collee-

tions under their care Jose H. | eal (BMSM) kindly pro-

vided digital images of paratype BMSM 15301 (C. cirri-

tum). Andy Murray (Bradenton, FL), Greta Polites (Uni-

versity of Georgia, ee Kevin Schindler (Flagstaff,

AZ), and W ends Conway (Sun City, FL) helper with

specimen donations and field assistance. Sean Roberts

(FLMNH) assisted with digital photography using a Sony

DSC RI camera (10.3 megapixel resolution). ). Support for

field studies was provided by the McGinty Endowment

of the Florida Museum of Natural History and by Bar-

bara and Reed Toomey (both to R.W.P.). Thanks to the

anonymous reviewers who improved the final version of

the manuscript. This is University of Florida Contribu-

tion to Paleobiology 595.

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THE NAUTILUS 121(3):131-138, 2007

Page 13]

Molecular phylogeny of some Indo-Pacific genera in the

subfamily eee H. Adams and A. Adame. 1853 (1838)

(Gastropoda: Neogastropoda)

Francisco M. Heralde HI

National Institute of Molecular Biology

and Biotechnology

Maren Watkins

Marine Science Institute

University of the Philippines

Diliman, Quezon City

PHILIPPINES

Gisela P. Concepcion

Marine Science Institute

University of the Philippines

Diliman, Quezon City

PHILIPPINES

University of Utah

John-Paul Ownby

Pradip K. Bandyopadhyay

and Department of Biology

University of Utah

Salt Lake City, UT 84112 USA

Ameurfina D. Santos

National Institute of Molecular Biology

and Biotechnology :

University of the Philippines

Diliman, Quezon City

PHILIPPINES

Baldomero M. Olivera!

Department of Biology

Salt Lake City, UT 841 12 USA

[email protected]

ABSTRACT

We have carried out a phylogenetic analysis of a group of Indo-

Pacific species in the subfamily Turrinae (Swainson, 1840) us-

ing 12S mitochondrial ribosomal RNA gene sequences. Most of

the species analyzed are convention: ally assigned to a: of three

genera, Turris Roding, 1798, Gemmula Weinkauff, 1875 or

Lophiotoma Casey, 1904. The molecular analysis ers that

while the species of Turris and Gemmuila analyzed in this study

comprise monophyletic groups, the species presently assigned

to Lophiotoma definitely do not constitute a monophyle tic as-

semblage and can be separated into two very distinctive groups

of species based on the molecular analysis. The species pres-

ently designated as Lophiotoma tayabasensis Olivera, 2004,

Lophiotoma panglaoensis Olivera, 2004, Lophiotoma indica

(Réding, 1798) and Lophiotoma bisaya Olive pra, 2004, are re-

lated to Lophiotoma | (Une ‘dogemmula) unedo (Kiener, 1839 in

1S34-S0) by molecular criteria, and are clearly ence ly sepa-

rated from Turris, Gemmula or L ophiotoma (as redefined). We

propose that Unedogemmula (MacNeil, 1960) be recognized as

a full genus; Unedogemmula unedo (Kiener, 1839 in 1834—S0)

is the type species, and the species above are transferred from

Lophiotoma to Unedogemmula

INTRODUCTION

Venomous gastropods comprise three groups: the cone

snails, the auger snails or terebrids and the Ba

classically included in a single family, Turridae (H. Ad-

ams and A. Adams, 1853): these families are ann

assigned to the superfamily Conoidea (Ponder and

' Author for correspondence

Waren, 1988: Taylor, Kantor, and Sysoev, 1993). It has

been apparent for some time that the * otainaele? comprise

the largest species group in the superfamily (see for ex-

ample, ‘Powell: 1966): however, the groundbre: iking work

ot Bouchet and co-workers ( Bouchet et al., 2002: 2004)

in New Caledonia, has provided evidence that >90% of

Conoidean biodiversity probably resides in the “turrids”

(broadly defined).

“Turrids” are problematic at all levels: not only are

they a megadiverse group (>10,000 species) with a large

fraction of species that remain undescribed, but the phy-

logenetic relationships within the group are poorly un-

derstood. The number of different “turrid” genera that

have been proposed is >600; although in traditional mol-

luscan taxonomic work all turrids had been assigned to

the family Turridae, in most of the more recent system-

atic treatments, the group has been split into 5-6 ‘differ-

ent families (Taylor et al., 1993; Bouchet and Rocroi,

2005). However, some standard taxonomic treatments

retain the old nomenclature (see for example Kohn,

1998S).

To complement our ongoing study of turrid venoms

(see for example Watkins et al., 2006) we initiated « a study

of “turrid” molecular phylogeny: the first results at the

generic level are reported below. The genus Turris

Réding, 1798, is the nominate genus of the family Tur-

ridae, with Turris babylonia (Linnaeus, 1758) as the type

species. This is an e xclusive ly Indo-pacific ge nus. How-

ever, there has been inconsistency in the definition of

those genera which are traditionally oe together

with Turris in the subfamily Turrinae. Two other groups

largely from the Indo-Pacific, Lophiotoma Weinkaut!

1875, and Gemimula Casey, 1904, are included in the

Page 132

THE NAUTILUS, Vol. 121, No. :

subfamily by most workers; in the western Atlantic aa

eastern Pacific, the genus Polystira Woodring, 1928,

also thought to have a close affinity to Turris (the es

species for these genera are Lophiotoma acuta (Perry,

1811), Gemmula hindsiona (Berry, 1958) and Polystira

albida (Perry, 1811)) Additionally, a number of other

groups (such as Tur ridh upa (Hedley, 1922)) are regarded

as turrine by most workers.

A major motivation for these studies is to investigate

the toxin genes expressed in the venom ducts of

gastropods. Among the different groups of

conoidean <¢

turrids, we have initially coucentrated on studying the

gene products expressed i in the venom ducts of species in

the subfamily Turrinae, since they are larger and more

easily collected than are most other forad groups. We

hope to correlate the gene families expressed in venom

ducts with the nielecular phylogeny of the species ana-

lyzed. Thus, the molecular analy sis reported below has

focused on lar ger Indo-Pacific species in the subfamily

Turrinae, e.g. Turris, Lophiotoma and Gemmula spp.

The data that we present below demonstrates that two

groups of species presently assigoned to the genus

Lophiotoma, which appear to be dloscly related by shell

morphology, prove to be unexpecte dly divergent when

evaluated by molecular criteria and need to be placed in

different genera.

MATERIALS AND METHODS

Specimen Collection: — Species analyzed in this study,

shown in Table 1, were mostly collected by commercial

shell collectors in the Philippines, except for Polystira

albida (Perry, 1811), a generous gift of Drs, Estuardo

Lopez Vera and Ed Heimer, and Lophiotoma cerithifor-

mis (Powell, 1964), which was collected in Oahu, Hawaii.

Specimens of each were preserved either in RNAlater®

(Ambion Inc., Tx) or 95% ethanol, and DNA extracted as

described below. In most cases, the digestive gland was

used as the source of DNA; however, for alcohol pre-

served specimens where the shell had not been cracked,

the digestive gland was often degraded, and DNA was

extracted from foot tissue.

Identification and sequencing of clones encoding 12S

mitochondrial rRNA gene segments: Genomic DNA was

prepared from tissue (~20 mg) from each turrid species

using the Gentra PUREGENE DNA isolation kit (Gen-

tra Systems, Minneapolis, MN) according to the manu-

renomic DNA from each

species (~lOng) was use dias a template for polymerase

fac ture rs standard protocol. ¢

chain reaction (PCR) with oligonucleotides correspond-

ing to 12S-I (5' TGC CAG CAG YCG CGG TTA ) and

12S-III (5' AGA GYG RCG GGC GAT GTG T) mito-

chondrial rRNA segments (Oliverio and Mariottini,

2001). The 5’ and 3’ primers included adapte rs GGAGA-

CAU and GGGAAAGU respectively for annealing to the

cloning vector pNEB206A. The POR cycling profiles

vere as follows: initial denaturation (95°C, 60s): followed

hy 40 cycles of denaturation (95°C, 20s): annealing ( 55°C.

20s) and extension (72°C, 30s), The PCR products were

Table 1. List of species analyzed in this study.

Species

Locality

Lophiotoma acuta (Perry,

1811)

Lophiotoma bisaya® Olivera,

2004

Lophiotoma cerithiformis

(Powell, 1964)

Lophiotoma cingulifera

(Lamarck, 1822)

Lophiotoma indica®

(Réding, 1798)

Lophiotoma Jickelii

(Weinkautf, 1875)

Lophiotoma olangoensis

Olivera, 2002

Lophiotoma panglacensis®

Olivera, 2004

Lophiotoma polytropa

(Helbling, 1779)

Lophiotoma tayabasensis®

Olivera, 2004

Lophiotoma unedo®

(Kiener, 1839 in

1834-80)

Gemmula speciosa (Reeve,

1543)

Gemmula diomedea Powell,

1964

Gemmula rosario Shikama

and Hayashi, 1977

Gemmula lisajoni Olivera,

2000

Gemmula sogodensis

Olivera, 2005

Turris garnonsii (Reeve,

1$43)

Turris grandis (Gray, 1833)

Turris normandavidsoni

Olivera, 2000

Turris babylonia (Linnaeus,

1758)

Turris spectabilis (Reeve,

1S43)

Turris totiphyllis Olivera,

2000

Polystira albida (Perry,

1811)

Drillia regius (Habe and

Murakami, 1970)

Buenavista, Marinduque,

Philippines

Batangas, Philippines

Oahu, Hawaii

Cawoy, Olango Island,

Philippine S

Aligway Is. Dipolog

Philippines

Cawoy, Olango Island,

Philippine S

Cawoy, Olango Island,

Philippines

Panglao Is. Bohol, Philippines

Bataan, Luzon, Philippines

Sogod, Cebu, Philippines

Panglao Is. Bohol, Philippines

Batangas, Philippines

Sogod, Cebu, Philippines

Cawoy, Olango Is and,

Philippines

Sogod, Cebu, Philippines

Cawoy, Olango Island,

Philippines

Cawoy, Olango Island,

Philippines

Cawoy, Olango Island,

Philippines

Bay of Campeche, Mexico

Panglao Is., Bohol,

Philippines

° These species a

re proposed to be transferred from

Lophiotoma to Unedogemmula (see text).

purified using the PureLink PCR Product Purification

Kit (Invitrogen Life Technologies, Carlsbad, California)

following the manufacturer's suggested protocol. The

eluted DNA fragments were digested with uracil specific

excision reagent, annealed to pNEB206A vector

(USER™ Friendly Cloning kit, New England BioLabs,

Inc., Beverly, Massachusetts) and the resulting products

tr wastorine d into « competent DEH5e cells ( Samnfrcle anc

F.M. Heralde III et al., 2007

Russell, 2001). Plasmid DNAs were isolated from ampi-

cillin resistant colonies and the nucleic acid sequences of

the inserts determined using AB] DNA sequencer with

ABI Big Dye chemistry (Foster City, DNA se-

quences have been ie re to GenBank and the

accession numbers are: EF467333, EF467333

EF467335, EF467336, EF467337, E splints

EF467339, EF467340, EF467341, EF46734

EF467343, EF467344, EF467345. ae

EF467347, EF467348, EF467349, EF467350,

EF467351, EF467352, EF467353, EF467354,

EF467355, and EF467356

ies

»)

Sequence Analysis: — Nucleic acid sequences (the long-

est of which had 593 nucleotides) were aligned manually

using MEGA version 3.1 (Kumar, 2004). One tree was

created from two independent runs using the software

program MrBayes (Huelsenbeck, 2001: Ronquist, 2003).

5,000,000 trees were made in each run, 50,000 of which

were saved. Two hundred and fifty of each of those

50,000 were also discarded as burn-in. Each run had four

chains (one cold and three heated). The two independent

runs were combined into a single tree where branches

were preserved if they were found in 70% or more of

those trees not discarded. The standard deviation after

5,000,000 generations was 2.401 x 10~°. A general time

reversible (GTR) model was used, with the rate variation

of some sites kept invariable and the remaining rates

drawn from a gamma distribution. The other tree was

created using the software program PHYML (Guindon,

2003). A thousand trees were obtained using non-

parametric bootstrap analysis and combined into a single

tree where branches were preserved if they were found

in 70% or more of the given trees. A G TR model was

used, with the base frequency estimates found empiri-

cally and the proportion of invariable sites estimated.

Four substitution rates were used, with the gamma dis-

tribution parameter estimated.

RESULTS AND DISCUSSION

PCR Amplification of 12S Sequencing: The se-

quences of 12S <7 from 23 species in the subfamily

Turrinae (see Table 1) were obtained as described above.

The 12S sequence of a Drillia species, Drillia regius

Habe and Murakami, 1970), was used as the outgroup

for the phylogenetic analysis. The sequences obtained

are shown in Table 2: these were aligned for maximal

overlap.

Phylogenetic Analysis: — A phylogenetic tree, shown

in Figure 1, was constructed as described under Meth-

ods. The species that are presently assigned to two of the

major turrine genera, Turris and Gemmula, appear as

monophyletic clades in the phylogenetic tree obtained

through Bayesian methods. However, the Lophiotoma

species analyze d clearly split into two distinct, well-

separated groups.

Thus. the species presently assigned to Lophiotoma

analyzed in this study do not appear to constitute a

monophyletic assemblage. A large separation is found

between two groups of Lophiotoma species; one group

includes Lophiotoma unedo (Kiener, 1839 in 1834-80),

Lophiotoma tayabasensis Olivera, 2004, Lophiotoma

panglaoensis Olivera, 2004, Lophiotoma indica (Réding,

1798), and Lophiotoma bisaya Olivera, 2004. These spe-

cies appear to be much more divergent from the Turris

and Gemmula branches than the other group of

Lophiotoma, which includes the type rie of

Lophiotoma, Lophiotoma acuta (Perry, 1811); the latter

comprises two cre branches, one branch pheal spe-

cies such as Lophiotoma cingulifera (Lamarck, 1822),

assigned by many syste matists to the sub yFenus Xenuro-

turris Iredale, 1929, which is regarded as a separate ge-

nus by some workers (Powell, 1966).

Generic Classification and Nomenclature: = The un-

expected phylogenetic separation between two groups of

species conventionally assigned to the genus

Lophiotoma, makes the present assignment of these spe-

cies into the conventional Indo- Pacific turrine genera,

Turris, Lophiotoma, and Gemmutla inconsistent with the

phylogenetic tree shown in Figure 2. One potential so-

lution would be to lump these Indo-Pacific genera to-

gether under one genus, Turris, and use subgeneric des-

ignations for each large clade of species (this might be

called the “Conus alternative”; the major group of Indo-

Pacific Turrinae comprise a phylogenetic branch that

does not appear to be more dive ergent by molecular cri-

teria than is the divergence within the species presently

assigned to the genus Conus; Espiritu, 2001). ). Although

this altermative may have some merit, the substantial lit-

erature referring to species in the traditional genera Tur-

ris, Gemmula, and nuns (including a significant

paleontological component of the research literature)

would make this a radical (and probably impractical) al-

ternative.

Because Lophiotoma acuta is the designated type spe-

cies for the genus Lophiotoma, the species presently in

Lophiotoma that are in the branch not including L. acuta

require a new generic designation. There are two ge-

neric/subgeneric designations potentially available for

the group. One is a name proposed originally by Powell,

Lophioturris Powell, 1964; Powell envisione d Lo} hiotur-

ris as a genus allied to Lophiotoma with Loptanionis

indica (Réding, 1798) as type. Lophioturris was set up

specif Really { for forms that have blunt paucispiral proto-

conchs; since most of the species in this clade have po-

lygyrate (multispiral) protoconchs, Lophioturris does not

seem to be an appropriate taxonomic designation (see

Powell, 1964, for a discussion of differences in proto-

conch morphology).

The other available generic name for this group of

species (which would have priority) is Unedogemmula

MacNeil 1960: as originally proposed, Unedoge ~mmula

was a separate genus, with l indogemmula une do (Kiener,

1839 in 1534-80) as type. However, P< well relegated

Unedogemmula to be a subgenus of Gemmula. Subse-

THE NAUTILUS, Vol. 121, No. 3

Page 134

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[900] WLL OVWOVLLVYY OVLLLLIOWY LOOVOLODLO LLIOOLYLYLS LIODWOLISL OLLLLLLIOD WWLOOVIVLL SOYODLOVLY DOLYVLYOID LOLIDYYODD OVLIOYOOYO VLLLISLOD) SOLIOYOOWY WOLOWWWVLI LYVYVLISIO LYYOLYLOWL SYOYOOWLLL WY-LLLOWYL LYWWLIVLYOD LIOLIYLLY If

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(900) WLL OVWOWOOVYY OVLLLLIOVY LODYOLODLO LISOLVLYLO LIOOWILLIO OLLLLLLIDD YOLIIVLVID ODYODLOVLY OOLYYLVILD LOLIDYYDOO OVLLLYOIVO WLLLLYLOOD SOLLOVOOWW WOLIVYYWLL LYVWWLVLY- LYWOLYLOVL OVOLOIYLLL Y-VLLLYWYL LYVVIOLWDO LIDLI¥IOW) PT

[909] WLL OVYOWLOVVY SVLLLLLOVY LODVOLODLO LISOLYLVIO LIDOWILLLO SLLLLLLIOD YOLIDWOVLL OIWILOVLY OOLYYLYOLD LOLIDVYDOD OVLLLYOOWO WLLLLYLOOD OOLLOYOOWW WOLOYYYWLL LYYYWOVLY- LYYOLYIOVL OYOOODVLLL Y-VLLLY¥¥L Lyv¥lolvod LIdLIviov) ct

(90%) VLL OVVOVLLYYY OVLLLLIOVY LODYOLODLO LLOOLYLVLO LIOSWOLLID SLLLLLLIOD YWLODVOVLL SDWIDLOWLY OOLYYLYOLD ISLIIVWOOO OWLLLYIOWD VLLLLYLOSD SOLIOVODYY WOLOVYYWLL LYVWWOWOY- LYWOLYDOVL OVSLODWLLLE Y-VLLLYYYL LYYVIVLLOD LIDLivlow CT

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[900] VWLL SYYDYLLYYY OVLLLLIOVY LOOYOLODLO LLOOLYLYLO LLLOWOLLIO OBLLIOLLIOD WWLODVOVLL OOWDILYWLY OOLYVIVLLD LOLIOWYODD OVLLLVIOYD VLLLLOLODD OPLLOYOOWY WOLOYYWYLL LYYYYLYIOY LYVOLYIOWL OVOWDOWLLL WYYY-LYYY- LYY¥LYLIOS IODLIVLLYD oT

[90%] VLL OVVOVLLYYY OVLLLLIOVY LOOYOLOOLD LIODLVLYLO LLLSYOLLIO OLLIOLLIOD WWLOOVOVLL OOWVDOLYVLY ODLYVIVLID LOLIDYYOOD OVLLLYIIYD VLLLLOLOSD SOLLOVDOWY WOLOWWWWLL LYVYWLYLOY LYVOLYLOWL OWVOWDIVLLL YW-WWLYYY- LYYYLYLLOD IDDLI¥IDYD 6

[900] VLL SYWDWLLYYY OVLLLLIOVY LOOYDLODLO LLODLVLYLO LLLOYOIOLD SLLIOLLIOD YWLODVOVLL SOWODLYYLY SOLYULYLIO LOLIOWYDOD OVLIOVOOVO WLLLYALODD OOLLIOVOOYY WOLOWWWWLL LYVYYLYLLY LYVOLYLOWL SYOWOIVLLL YY-VLLYY¥- LY¥V¥LYLIO9 IODLIYLIWD 8

[900] VLL OYOOWLLYVY DVLLLLIOVY LOOYOLOOLD LLDDLVLVLS LIOOWOLLID OLLLLLLIOD VYLOOVLVLL SOWODLYWLY OOLYWLYOILD LOLIOVYODD OVLLOVOIYD VLLLLVLO9D SALIOYOOWY VOLOYWYWLL LYYVYLYO-Y LYVOLVIOVL SYOVIOVLLL WYWLLLYYYL OYWWLVLYSS LOOLLWILYS ¢

[900] WLL OVVOVLLYVY OVLLLLLOVY LODWOLOOLD LISDLYLVLD LIOSYOLLLO OLLLLLLIOD YYLOOVOVLL SOWODOLYVLY SOLYYLYOIO LOLOIOVYOOD OVLLLYIOVD VLLLLVLOOD SDLIOVOOWY VOLOWVWWLL LYYYVLYL-Y LYVOLYIOVL OYODOOVLIL YYWLLLVWYL LYYYIOLYDS LODLIYLLYD 9

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[90] VLL OYYOWLOWVY OVLLLLIOVY LOOVOLODLO LLOOLYLYLO LIOOWOLLLD OLLLLLLIOD YVLODVIVLL SOWDOLOVLY SOLWYLYOLD LOLOOYYDOD OVLLLYIDYO VLLLLVIODD SOLIOVOOYY VOLOVYYWLL LYYWYLYOVS LYVOLYLOWL OYOWIIVLEL YOVLLIYVYOL OYYYLOLYSD DOoLivLliva ©

[90%] VLL SYYOWLOWYY OVLLLLIOVY LODYOLODLO LLODLWLVLO LL-SVOLIOL OLLLLLLIOD YWLOOWLVOL OOWOOLOVLY OOLYWLYOLD LOLIIYYOON OWLLLYOIYO VLLLLOLODS POLLIOVOOWY WOLOWYYWLL LYYWYLVLYY LYVOLYDOVL OYOVIOVLLL WWWLOLYYYD LYV¥LOLYOD LOQLI¥LOW ¢

[900] VWLL OYVOVLOVWY OVLLLLIOVY LODYOLOOLOS LIODLYLYLS LIOOVOLLIO SLLLLLLIDD YWIOOVLOLL SOWIDLOVLY OOLYVLOILO LOLIDVYOSO OVLLLYIDYD WLLLLVLOOD SOLLOYOOYY WOLOVWYWLL LYYYWLYOVY LYVOLYLOWL SVOWVOOWLEL YYYLIOWYWL OYYWISLYOD IODLIYLLIYD T

[€0Z] DDD WLYDYLOYDD OLOYYYDOOY OLLOLVYYWY LVLOLYYOLL WYV-LYWLL- -OWWLYYVVY DOVDVLLLLI WWWLYLYVYL LLIVYYWLOVY YYLVLYLLLO OLYWD-LLLL L---SOWLYY W-WLYY-LLY ----- LYOVL LOWLYOWWYY WLODOV-LLL -LYLYDLOWY LLYYWOYVWIO YYYLIODIOD SOYODWOOSL bt

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(E07) DDD VLIVOWLLYOD OLOVYYOSOY NOLODOYOWS LOLOLYWOLL DYYYWOOOV- WYYYLYYLLL L-OLY-LOL- LVWLVIVLISL LLYYVWLOOY WYYWLYLLLI 39-V¥-LOWL L--LOOWWLL LYLLLVY-WL¥Y ----- DWOVL LOVOWOWWYY VLOODWWLYY -LULYOLOYY LLOVOYYYDD YOVLIONDOO LOYSOVIOOL 0%

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F. M. Heralde III et al., 2007 Page 135

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Figure 1.

Phylogenetic tree of species listed in Table 1. A tree based on 12S rDNA sequences was constructed (see Methods)

Species from three Indo-Pacific families in the subfamily Lei were analyzed (Turris, Gemmula, Lophiotoma). In addition, one

Atlantic aes Polystira albida and a species in the subfamily Drillinae (or Clavinae), Drillia regius were included. Note that the

species of Lophiotoma are split into two widely separated groups. As discussed in the text, species marked by an asterisk are proposed

to be moved to another genus. Lophiotoma jickelii and Turris grandis are non-standard taxomomic assignments : the figured

specimens: these are widely regarded as synonymous to Lophiotoma acuta and Turris crispa, respectively (see Powell, 1964).

However, the molecular data clearly se parates Lophiotoma acuta and Lophiotoma jickelii, and the type of Turris crispa is sufficiently

divergent from the form shown that using Turris grandis seems a preferable name for this species.

quently, Kilburn (1983) suggested that based on general

shell morphology, Unedogemmula seems much closer to

species traditionally assigned to Lophiotoma, and that it

should more appropriately be regarded as a subgenus of

Lophiotoma; this suggestion has been adopted in most of

the recent taxonomic treatments of turrine genera.

The molecular results reported above demonstrate

that Unedogemmula unedo is indeed closely related to

some of the spe cies most taxonomists prese mitly include

in Lophiotoma, (such as Lophiotoma indica, Lophiotoma

bisaya. and Lophiotoma tayabasensis). Furthermore, the

molecular analysis clearly shows that there is no justifi-

cation for designating Unedoge mmula as a subgenus of

either Gemmula or Lophiotoma, since Unedogemmula

unedo is in a very divergent branch of the phyloge netic

tree. Thus, our results support the original designation of

Unedogemmula as a full genus, although the species

comprising the genus need to be somewhat redefined.

Unedogemmula unedo is the type species, and the larger,

strongly maculated forms previously assigned to

Lo} shiotote (such as Lophiotoma indica, Lophiotoma

ee nsis, and Lophiotoma bisaya) are transferred to

Unedogemmula from Lophiotoma. A recent analysis of

Philippine forms related to these species, clarifying the

relationships between these forms and the Unedogem-

mula unedo group, was recently published (Olivera,

THE NAUTILUS, Vol. 121, No. 3

ee eee ee ee ee Drillia regius

Polystira albida oii i \o

Unedogemmula panglaoensis aa

98 70

Unedogemmula unedo

96 91

100

Turris babylonia ah

Seca

Unedogemmula tayabasensis

Unedogemmula indica =X.

Turris garnonsii Aa

Turris grandis —-=

Turris spectabilis al

100

Turris totiphyllis =:

Gemmula diomedea = [we

100 86

Gemmula speciosa

100

85 400

100

a |

0.1 expected substitutions per site

Moure 2

Figure 2.

generations). Note that all the major groups are highly supp

assigned to Lophiotoma is now regarded as a full genus, Unedogemmula, with Unedogemmula unedo as type

figure, these species (st

2004

. ated in that work a

nedogemmutla, as rede fine: d below, ae this should

it seems likely ong all of the species level taxa

» properly placed in the genus

be veritied by obtaining molecular data for those species.

Genus Unedogemmula MacNeil, 1960

30-LO5mm, fusiform with

anterior canal unnotched

Shell large

straight,

Description:

tall s

toconc h variable

often with the transitional part of the larval shell deco-

pire long Pro-

from blunt paucispiral to multispiral,

rated with brephic axials or axially costate whorls. Sinus

ill labeled Lophiotoma in Figure 1) are

Gemmula sogodensis ~~ ap

Gemmula lisajoni ee NN

Gemmutla rosario

100

The phylogenetic tree in Figure 1 is re-plotted, except that the confidence limits are included (calculated after 10 million

wrted; as is explained in the discussion, one of the groups formerly

species; thus, in this

designated as Unedogemmula spp.

is peripheral, deep, and narrow at the termination of the

sinus rib.

Remarks: ~The genus has shell morphology with strong

similarities to Lophiotoma: most spt ae s have a smooth

peripheral keel but, in some there are distinct

peripheral granulations. In other species, these granula-

tions continue to the body whorl, but these tend toward

obsolescence in most forms. In contrast to Lophiotoma,

the peripheral keel does not consist of two raised cords at

spe cle S,

the edges with a depress sd area in the center; rather,

F. M. Heralde III et al., 2007

Page 137

some forms, there is a single smooth rib exhibiting a

variable level of peripheral granulation.

In order to test the veracity of the proposals based on

the Bavesian analysis and outlined above, a second ap-

proi ach to the phy logenetic an: ysis of the sequences was

also carried — A Maximum Likelihood method was

employed as described under Me thods. The results of

se analysis are shown in Figure 3. The separation of the

species described above origin: ily in Lophiotoma into

two distinct groups is strongly sup porte ed using this analy-

. Thus, both methods support raising Unedoge mmula

The analyses differ,

Gemmula; the maximum

. a full genus, as described above.

however, in the results with

likelihood method does not group all of the species ana-

lyzed into a single monophyletic clade, but into two

groups of species. Thus, given this discrepancy between

the two methods, the monophyly of Gemmula clearly

requires further investigation.

A brief summary of a proposed revision of Indo-Pacific

genera in the subfamily Turrinae is given in Table IIL.

The cladogram in Figures 2 and 3 give support to Turris

and Unedogemmula. However, Lophiotoma (redefined

to exclude the species transferred to Unedogemmutla) has

two branches, both strongly supported; the conjoining of

the branches has less than 90% support in the Bayesian

analysis, and is not supported above the cutoff level

Drillia regius

Polystira albida

Unedogemmula panglaoensis

Unedogemmula tayabasensis

Unedogemmula unedo

87 100

Unedogemmula indica

Unedogemmula bisaya

84 Turris normandavidsoni

Turris garnonsii

Turris grandis

Turris babylonia

Turris spectabilis

Turris totiphyllis

Gemmula diomedea

Gemmula sogodensis

Gemmula speciosa

100

Figure 3. Phylogenetic tree of species listed in Table 1

the tree in Figure 1, but was constructed using the PHYML

This tree was made using the same sequence a

software

Gemmula lisajoni

Gemmula rosario

Lophiotoma polytropa

Lophiotoma jickelii

97 Lophiotoma acuta

Lophiotoma olangoensis

Lophiotoma cerithiformis

Lophiotoma cingulifera

lgnment as that used for

program

- > IAQ

Page 135

THE NAUTILUS, Vol. 121, No. 3

(70%) in the Maximum Likelihood analysis. Given these

data, the solution would be to split Lophiotoma into two

separate genera, Lophiotoma and Xenuroturris. We feel

that at this time the more conservative approach of re-

taining the genus Lophiotoma, and dividing it into two

subgenera, Lophiotoma (s.s.) and Xenuroturris (with

Lophiotoma (Lophiotoma) acuta and Lophiotoma (Xe-

nuroturris ) cingulife ra as types, respectively) is prefer-

able until a wider range of species has been analyzed.

There are a number of species (presently in Lophiotoma)

that are problematic to assign (such as Lophiotoma ruth-

veniana (Melvill, 1923)), and we believe that a molecular

analysis of these forms needs to be carried out before we

fully understand the relationship between Lophiotoma

(s.s.) and Xenuroturris. It may well turn out that, when

the eae is completed, the separation between the two

branches (Lophiotoma and Xenuroturris) will be defini-

tive; at that point, separating the two groups of species

into different genera will be justified.

The major conclusion from this work is that

Unedogemmula should be recognized as a full genus, and

is a sister group to the major branch that includes Turr is,

Gemmula, and Lophiotoma (as redefined).

ACKNOWLEDGMENTS

We thank Sean Christensen for help with experiments

Bradford Stevenson for help with the preparation of phy-

logenetic trees and Drs. Edgar P. Heimer de la Cotera

and Estuardo Lopez-Vera for providing the Polystira

sample. This work was ea ed in part by a Program

Project grant from the National Institutes of Healthy,

GM48677.

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Caledonia site. Biological Journal of the Linnean Society

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Bouchet, P. and J. P. Rocroi, 2005. Classification and nomen-

clature of gastropod families. Malacologia 47: 1-397.

Bouchet, P., A. Sysoey, and P. Lozouet. 2004. An inordinate

fondness for Turrids. In: Molluscan Megadiversity: Sea,

Land, and Freshwater, [Abstracts of the] World Congress

of Malacology, Perth, Abstracts, p. 12.

ee D. J. D., M. Watkins, V. Dia-Monje, G. E. Cartier,

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Guindon, S. a O. Gascuel. 2003. A simple, fast and accurate

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lihood. Systematic Biology 52: 696-704

Huelsenbeck, J. P. and F. Ronquist. 2001. MRBAYES: Bayes-

ian inference of phylogeny. Bioinformatics 17: 753-755.

Kilburn, R. N. 1983. Turridae (Mollusca: Gastropoda) of south-

ern Africa and Mozambique. Part 1. Subfamily Turrinae.

oo of the Natal Museum 25: 549-585

—_ A. J. 1998. Superfamily Conoidea. In: Beesley, P. L.,

>. J. B. Ross, and A. Wells (eds.) Mollusca: The Southern

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Kumar, S., K. Tamura, and M. Nei. 2004. MEGA3: Integrated

Software for Molecular Evolutionary Genetics Analysis

and Sequence Alignment. Briefings in Bioinformatics 5;

150-163.

Olivera, B. M. 2004. Larger forms in Lophiotoma: Four new

species described in the Philippines and three from else-

where in the Indo-Pacific. Science Diliman 16; 1-28.

Oliverio, M. and P. Mariottini. 2001. A molecular framework

for the phylogeny of Coralliophila and related muricoids.

Journal of Molluscan Studies 67: 215-224.

Ponder, W. F. and A. Warén. 1988, Classification of Caenogas-

tropoda and Heterostropha—A list of family-group names

and higher taxa. Malacological Review, Suppl. 4: 288-328.

Powell, A. W. B. 1964. The family Turridae in the Indo-Pacific.

Part 1. The subfamily Turrinae. Indo-Pacific Mollusca 1:

297-346.

Ronquist, F. and J. P. Huelsenbeck. 2003. MRBAYES 3: Bayes-

ian phylogenetic inference under mixed models. Bisse

matics 19: 1572-1574.

Sambrook, J. and D. W. Russell. 2001. Molecular Cloning—A

Laboratory Manual. Cold Spring Harbor Laboratory

Press, Cold Spring Harbor, New York.

Taylor, J. D., Y. I. Kantor, and A. V. Sysoev. 1993. Foregut

anatomy, feeding mechanisms, relationships, and classifi-

cation of the Conoidea(=Toxoglossa)(Gastropoda). Bulle-

tin of the Natural History Museum, London (Zoology) 59:

125-170.

Watkins, M., D. R. Hillyard, and B. M. Olivera. 2006. Genes

expressed in a Turrid venom duct: divergence and simi-

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247-256.

THE NAUTILUS 121(3):159-145, 2007

Page 139

Reproductive biology of the nudibranch Doris fontainei

VOrbigny, 1835 (Gastropoda: Opisthobranchia) from the

M agellanic Region

Le Co

Claudia Muniain'

Museo Argentino de Ciencias Naturales

“Bernardino Rivadavia”

Avda. Angel Gallardo 470

C1405D]R, Buenos Aires

ARGENTINA

[email protected]

Valdivia, CHILE

Carlos S. Gallardo

Instituto de Zoologia E.

Universidad Austral de Chile

Pablo E. Penchaszadeh

Museo Argentino de Ciencias Naturales

“Bermardino Rivadavia”

Avda. Angel Gallardo 470

C1405D]R, Buenos Aires, ARGENTINA

and

Departamento de Biodiversidad y

Biologia Experimental

Facultad de Ciencias Exactas y

Naturales

Universidad de Buenos Aires

Buenos Aires, ARGENTINA

F. Kilian

ABSTRACT

The present study describes the egg deposition, embryonic and

larval development, and reproductive behavioral traits of the

South American dorid nudibranch Doris fontainei dOrbigny,

1835, observed in laboratory under controlled conditions. Be-

havior during copulation and spawning was recorded using a

digital video camera. Copulation and spawning lasted 24 h and

22-93 h, re spectively. The spawned mass reached 1.80 m in

length and was coiled in a counter-clockwise direction from the

centre with one edge affixed to the substrate. The ribbon con-

tained numerous small capsules, each having 2-4 eggs of about

S6 ym in diameter. Intracapsular egg development lasted

about 14 days at 14.5°C, culminating with the release of up to

1.25 million veliger larvae per egg mass. The new veliger larvae

showed incipient development of the foot and a relatively small

protoconch (ca. 158 jm), indicating they enter a planktotrophic

phase. A comparison with other nudibranchs, and particularly

with dorids, suggests that D. fontainei has an annual cycle

whose egg mass fits a morphological pattern typical of the fam-

ily. The characteristics of its encapsulated development show it

is one of the most fecund species among those having this

pattern, which also explains, in part, its geographical dispersion

from Peru in the Pacific to northern Argentina in the Atlantic.

Additional Keywords: Nudibranchia, reproductive behavior,

multiple embryos, planktotrophic development

INTRODUCTION

An interesting characteristic of nudibranch gastropods is

their capacity to produce ee and delicate egg

masses, which can be obtained in laboratory through ae

equate maintenance of adult individuals. The pattern of

early development of these egg deposits is a basic trait in

: :

Author for correspondence

the life histories of the different species. These patterns,

which can be de termined through observations under

controlled conditions, may also provide new elements to

better typify and distinguish species whose taxonomic

status is still debated.

Hurst (1967) has grouped the egg deposits of opistho-

branchs into three morphological types in accordance

with the morphology of the ribbon, mode of attachment,

and alignment of the capsules. However, Thompson

(1967) defined three developmental larval strategies:

planktotrophy, lecithotrophy, and direct dev elopment.

Although there is general knowledge about egg depo-

sition and definition of some dev elopme ntal parameters

for different species worldwide, the priate biology

of the rich and diverse nudibranch assemblages inhabit-

ing the coasts of South America is still poorly known.

Doris fontainei dOrbigny, 1835, is a widely distributed

and common guiiscach from the South American

coast. This sea slug inhabits the extensive littoral fringe of

the SE Pacific (Peru and Chile), spanning the entire

Magellanic region, to latitude 38°S in northern Argen-

tina, following cold Antarctic currents to depths of 70 m

(Muniain et al., 1991; Muniain, 1997; Schrédl, 1997a, b:

2000). As with other dorids occurring along this stretch

of coastline, D. fontainei has been subjected to several taxo-

nomic revisions, including a recent re-assignment based

on detailed anatomical studies (Valdés and Muniain,

2002).

An exploratory sampling for nudibranchs on the coast

near Valdivia, Chile (40°S), allowed us to collect and

maintain living specimens of D. fontainei in the labora-

tory and observe their reproductive behavior, including

copulation, spawning, some aspects of individuals behav-

ior after completion of egg deposition, and plank-

totrophic larval stage.

Page 140 THE NAUTILUS, Vol. 121, No. 3

C. Muniain et al., 2007

Page 141

MATERIALS AND METHODS

A total of six specimens of Doris fontainci were collected

from Los Molinos (40°S), Valdivia, (Chile), by scuba div-

ing to depths of 3-6 m a 4-16 March 2001. The water

temperature at the sampling site was 14-15°C. To obtain

information on the copulatory and reproductive behavior

of Doris fontainei, the specimens were maintained in

aquaria containing aet rated seawater at approximately

14.5°C, at the Zoological Institute of the Universidad

Austral de Chile, Valdivia. Egg masses deposited on the

aquarium walls were cultured in the original container

without dislodging the egg mass. Routine isothermal

seawater changes were made daily with water filtered to

1 um. Egg dev elopment was monitored until hatching.

Egg samples were preserved in 6% formalin in seawater

for further microscopic studies, which included counting

and measuring the capsules and the eggs and embryos

contained inside them. Most of the information pre-

sented in this study was recorded by photography and

digital video during maintenance of living individuals.

Complementary bhsernations were made using pre-

served material. Voucher material was deposited in the

Museo Argentino de Ciencias Naturales “Bernardino

Rivadavia” under accession number MACN 36542.

Egg deposits were described from macroscopic obser-

vations as well as using a Zeiss Axiostar stereoscopic mi-

croscope equipped with a digital camera and imaging

software at Museo Argentino de Ciencias Naturales

“Bernardino Rivadavia”. The number of eggs per capsule

were determined using repeated counts in randomly

chosen locations throughout of the egg ribbon. Develop-

ment was observed under microscope and each develop-

mental stage illustrated, particularly that of the troco-

phore and middle vel iger; mean numbers of embryos per

capsule, and their size relation to the size of the capsule

were ca Iculated. Capsular volume (V) was determined

(in wm”) based on measurements of capsular length and

width, assuming that the depth axis was equiv alent to the

width axis. The radii (r), ro, r3), and the volume was

calculated using the formula:

= 4/37 x 1 x Yo X Is.

Sixty capsules previously separated in groups according

to the number of embryos they contained were 1 randomly

extracted from the gelatinous matrix and measured as

mentioned above.

The diameters of 40 eggs at the unsegmented zygote

stage were measured. Descriptions of ihe dev elopment

included three embryonic stages: trochophore, middle to

advanced veliger, and pre-hatching veliger; each stage

was photographed and measured (n = 30). The total

number of eggs in a deposit was estimated indirectly by

counting egg capsules in two different 5 mm-wide seg-

ments of the egg ribbon. For this purpose representative

segments of ihe ribbon were isolated by cutting them

W ith a scissor and carefully extracting individual capsules

from the gelatinous mi itrix of each segment. The mean

number of eggs per cm of eg¢ ribbon was determined,

and the total number of eggs in the deposit was calcu-

lated based on the length of the entire ribbon. The rib-

bon length was detesmiadd by measuring the length of

its whatiached free edge from the photogr aphic images,

taking into account the photo scale. The approximate

number of embryos in the egg ribbon was estimated

multiplying the number of egg capsules by the average

number of embryos they éontained: the total number or

embryos was estimated following the same procedure as

for eggs. The development period was determined in an

egg deposit maintained under controlled temperature,

and was considered to start at egg deposition and finish

at the initiation of hatching. Recently hatched larvae

were observed under stereoscopic microscope.

RESULTS

Copulatory and Spawning Behavior: Copulation

and post-copulation behavior of two of the six speci-

mens collected and maintained in aquaria was observed

(Figures 1-8). During copulation, nudibranchs had the

rignt sides in contact at the level of the genital See

with the heads in opposite directions (Figure 1). Copu-

lation lasted between 2 and 4 hours, with the animals in

total contact; penises were not visible because they were

obscured by the mantle, although there was a conspicu-

ous penial papilla, as described by Muniain et al. (1991)

and Valdés and Muniain (2002). One individual 76.2

mm in length began to spawn in a counter-clockwise

direction eraistial) ) at 16:00 h on 13 Mar. 2001 and fin-

ished at 15:00 h on 14 Mar. 2001 (23 h) at a temperature

of 14.7°C. During spawning, movement was very slow,

with the rhinophores contracted within the sheaths and

the branchial tuft exposed but motionless. Following

deposition of the egg mass in the center of the aquarium,

the nudibranch slowly moved to one side. A second

specimen, 64 mm in length, began to spawn two hours

after collection on 16 Mar. 2001, showing sinistral ovi-

position, and terminated at 11:00 on a Mar. 2001, after

a total spawning time of 22 h (Figure 2). Following com-

plete spawning, the slug cemmained ss oa arebile:

contracting and completely invaginating very slowly, with

only the posterior portion of the foot adhered to the

Figures 1-8.

Doris fontainei 7 Orbigny, 1835, reproductive behaviour captured from digital video. 1. Copulation maintained in

aquaria. Scale bar = 1 cm. 2-3. Specimens showing sinistral oviposition (counter-clockwise direction). Scale bars: Figure 2 = 1 cm;

Figure 3 = 1.5 cm. 4. Detail of the post-spawned behavior. Scale bar = 1 cm. 5. After half and hour immobile, the nudibranch slow]

returning to a normal position. Scale bar = 0.5 mm. 6. Detail of rhinophores and gills exposed again. Scale bar = 1 cm. 7. Detail of

the free border and scattered spaces of the spiral ribbon. Scale bar =

ri, rhinophores; fo, foot

em. Abbreviations: gi, gills: ma, mantle: mo, mouth: ri

2 cm. 8. Complete spawn, specimen (64 mm). Scale bar = 1.5

‘ > 9

Page 142

THE NAUTILUS, Vol. 121, No. 3

” lll *

Figures 9-13. Doris fontainei @ Orbigny, 1835, 9-12. Details of the ovoidal capsule with 2 and 3 eggs. The volume of the capsule

increases with the number of embryos, 3 embryos is the most common number. Scale bar = 45 jam. 13. Hatching planktotrophic

larva, right lateral view. Typical coiled shell (type 1, Thompson, 1961). Scale bar = 25 pm.

operculum; rm, retractor muscle; se, statocyst; v, velum.

substrate (Figures 3, 4). The ventral surface of the

mantle comple tely covered the organism, which raised so

as to show the anterior portion of the foot. The nudi-

branch remained in this position for about half an hour

(Figure 5), slowly returning to a normal position, extend-

ing the rhinophores, cinnalline the mantle, and slowly

crawling away from the egg deposit ( (Figure 6). This be-

havior was completely re corde d with digit: u video from

11:00 to 12:10 hrs on 17 Mar. 2001 (Figure 8).

Egg Mass and Larval Development: The egg mass

formed a spiral ribbon of concentric rings consisting of

five to seven revolutions around a central point (Figures

The free border of the ribbon was undulated, un-

like the attached border, which was straight (Figure 7).

The undulations of the free border coincided with con-

centric rings of the ribbon. Both the egg deposits ob-

tained in the laboratory and those collected in the field

were pink. The latter were typically positioned under

rocks or in shaded areas.

The egg mass ae some mucous areas without

egg capsules (Figure 7). The egg capsules were arranged

line arly along the el abn. and were joined by a fine

cord (chalaza) that served to maintain the spiral configu-

ration. Randomly scattered spaces without capsules oc-

curred in the egg ribbon and in its different median

portions. The egg de posit from the 64-mm specimen was

formed in six revolutions around the center, with a maxi-

mum diameter of 10 cm; it was 13.4 mm in height, and

an estimated 1.8 m in length (Figure 8).

Mean egg diameter was 86.5 + 4.7 xm. The capsules

contained multiple embryos, although single embryos

appeared occasionally The number of e smbryos in cap-

sules ranged from 2 to 5,3 e ee . ing ‘clearly the

most common number ( Figures 9— The first count

based on 338 capsules, containing fe ee gave an

average number of 3 embryos per capsule. A second

count, of 361 capsules (containing intermediate veligers),

1300

Abbreviations: int, intestine; op,

produced an average number of 3.18 embryos per cap-

sule. The average of the 2 counts was about 3.1 embryos

per capsule.

The a tended to be ovoid, exhibiting some

more irregular shapes when crowded. The volume of an

egg capsule increased with the number of embryos it

contained (Figures 9, 11). As expected, the eggs demon-

strated typical spiral cleavage. Measurements of the

different stages were as follows: trochophore = 99. 35, -

5.5 jum (n = 20), advanced encapsulated veliger = 116.25 +

125 55 pm (n = 20), and hatched veliger larva = 158.1 +

7.16 wim (n = 10). Hatching of typically planktotrophic

veligers becha 14 days after incubation, at an avet rage

temperature of 14. 5°C. These veligers measured a mean

of 150 jzm (n = 20) in prostomial length, with the pro-

toconch sinistrally coiled, with 4 to 1 whorl at hatching.

At hatching, the larvae had no eyes nor propodium ru-

diment, although the velum, velar retractor muscle, lar-

val kidney, nephrocy sts, operculum, and cephalopedal

alimentary apparatus were developed (Figure 13),

The total number of larvae released from the egg mass

was estimated indirectly through observations under

controlled conditions. Microscope- aided counts carried

out on 2 transects of the egg ribbon showed an average

of 2236 egg capsules per cm. With an estimated length of

ca. 1.8 m (indicated above), the entire egg deposit con-

tained about 402450 capsules, and with the above esti-

mate of 3.1 embryos per cé ipsule, the entire egg deposit

contained about 1.25 million embryos, resulting in the

same number of developed planktotr yphic veliger larvae

a few days later.

DISCUSSION

The present study describes some of the traits that typify

the the history of D. fontainci, specifically in regard to its

reproductive pattern. Its relatively large body size sug-

C. Muniain et al., 2007

Page 143

gested a priori that the species would have an annual

evcle, with one generation per year, and the spawning

characteristics fell into Class A, ‘following the classifica-

tion and corresponding attributes of dorid nudibranchs

prov ided by Hurst (1967)

The characteristics of the e neapsulated development

described in the present study agree with the embryonic

developmental pattern observed in most of the Dorida-

cea, based on information summarized by Thompson

(1967) that typifies the three basic developmental stages

observed in that group of nudibranchs. The embryonic

development observed in D. fontainei fits Type-1 pattern

of this scheme. Free planktotrophic larvae are released at

hatching, there are multiple relatively small embryos per

capsule, and the period of e smbryonic development lead-

ing to hatching of weakly dev eloped veliger larvae is

comparatively short.

Based on information on related species (Thompson,

1958: 1967: Strathmann, 1987: Goddard, 2005) most

dorids with planktonic life cycles have eggs ranging be-

tveen 60 and 130 jm in size, many of which are in the

70-90 zm range, in agreement with values found for D.

fontainei. There is also agreement in the number of em-

brvos produced per spawning, with values as high as

several hundred thousand embryos per spawning in the

most fecund species, such as Acanthodoris pilosa ( coe

gaard, 1789) and Archidoris pseudoargus (Rapp, 1 gh

(both reviewed by Thompson. 1967). peer

fontaine is a highly fecund dorid, given that it ee

over a million eggs per spawning. This notably exceeds

the numbers reported for other species in the group; the

closest species is A. pseudoargus, which spawns 645000

eggs (Colgan, 1914, cited in TI soot a 1967). The du-

ration of embryonic development (pre-hatching) in

planktotrophic species can be as short as 6 to 10 days,

and rarely exceeds one month (Thompson, 1967). The

development period of 15 days found for D. fontaine:

falls within the range expected for dorids with plank-

totrophic dev elopment. Another trait coinciding with the

Type-1 pattern of development are the characteristics

and degree of development of the veliger larvae hatched

from the egg deposits of D. fontainei. As Thompson

(1967) and Todd et al. (2001) mentioned for species

showing this pattern, typical characteristics include ab-

sence of eyes or propodial rudiment, but presence of a

deve sloped velum, retractor muscle, larval kidney, oper-

culum, and cephalopedal digestive system, which were

the most notable traits observed by us.

Based on observations of lecithotrophic dorids of

the genera Adalaria, Dendronotus, Discodoris, Tritonia

(Thompson, 1958; 1961; 1962; 1976; Gohar and

Abul-Ela, 1959: Roginsky, 1962; Todd, 1979) and of the

direct-developing species Cadlina laevis Linnaeus, 1767

Roginsky, 1962: Thompson, 1967), the eggs are 200 and

400 j.m in size, respectively, and the total number of

embryos per spawn is clearly lower (aprox. 15000 to

50000), reaching only a few ‘hundreds in species with

direct development.

As indicated above, it is clear that D. fontainei has

multiple embryos per egg capsule, but given the lack of

information, we do not iow if this condition is common

within Doridacea, although Thompson (1967) mentions

it as a frequent characteristic among the opisthobranchs

that have planktotrophic larvae and Type-1 develop-

ment. Among the studies of reproductive aspects in other

species of M: agellanic opisthobranchs, the presence of

multiple embryos was noted in the saccoglossan Elysia

patagonia Muniain and Ortea, 1997, which completes

development with the formation of planktotrophic larvae

(Muniain and Ortea, 1997; Muniain and Penchaszadeh,

2000; Muniain et al., 2001).

Given the large number of embryos that D. fontainei

produces, eadlonine of more than one embryo per egg

capsule is probably an efficient mechanism for reducing

the energetic cost of these egg masses compared to the

relatively large benefits of maximizing the numbers of

larvae gener rated per spawning,

Although there is little information available on some

species in this region, the development of D. fontainei

can be compared with that of other dorids that oceur on

the Chilean coast, such as Peltodoris marmorata (Bergh,

1898) (cited as Anisodoris rudberghi). It has eggs of SO

wm in diameter and a larger number of eggs per capsule

(5-23), as well as a pre- hatching veliger of 125 jum in

length, and its embryonic development occurs in only 10

days at 14-16°C (Brokordt, 1992).

The large number of larvae produced by D. fontainei

suggests a possibly high degree of larval mortality and,

aesouiated with this, a potentially effective mechanism

for extensive larval dispersal. Wide dispersal can main-

tain gene flow among populations as well as the extensive

geographical dispersal (Schrédl, 1997a, b; 2000; Valdes

and Muniain, 2002) shown by this nudibranch in com-

parison with other species on the South American coast.

There are no records in the literature of detailed ob-

servations on the behavioral mechanisms accompanying

spawning of nudibranchs. The intriguing post-hatching

behavior shown by a specimen of D. fontainei in the

present study deserves further laboratory research with a

higher number of individuals to determine whether be-

havior has a given pattern at this stage of the reproduc-

tive process.

The duration of the spawning process (22-23 h) asso-

ciated with the relatively long, highly fecund egg ribbon

of D. fontainei may imply the generation of some facts of

phy siological imbalance, whieh the animal strives to over-

come through the behavioral event observed at the cul-

mination of oviposition. Whether this behavior results

from an inordinate energy demand compared to that

occurring in other species of nudibranchs is unknown.

Evaluations made in prosobranchs have shown he high

relative energetic cost involved in making benthic egg

masses (Perron, 1981; 1982).

Studies on the chemical ecology of this species have

shown that it biosynthesizes repellent compounds that

accumulate only in the mantle tubercles (Muniain, 1997

Page 144

THE NAUTILUS, Vol. 121, No. 3

Gavagnin et al., 1999). If this behavior is confirmed un-

der laboratory conditions, further studies should also

evaluate its possible occurrence in natural conditions,

where the nudibranch would be exposed to predation

throughout the entire stage.

Nudibranchs can be considered semelparous (despite

spawning repeatedly in a season) in that once they have

reached maturity, their period of spawning is inevit ably

followed by apparent genetically programmed post-

reproductive e death (Todd et al., 2001). There fore, repro-

duction must be fully successful in the single opportunity

presented to these animals, which will probably not be

able to survive until another reproductive season to re-

peat the process. Such is the high cost of reproductive

activity for the individual. The bvoad geographic distri-

bution of D. fontainei makes it a useful specie s for future

studies of pc yssible geographic variations in its pattern of

embryonic development. Further studies including re-

lated nudibranch species in the region should be con-

ducted. For example, studies of the dorid Adalaria

proxima (Alder and Hancock, 1854) have shown that the

egg diameter and other embryonic and larval traits could

show intraspecific variation, reflecting the adjustment of

its populations to a given range of variability in environ-

mental conditions where the species is distributed (Jones

et al., 1996; Todd et al., 2001).

ACKNOWLEDGMENTS

This study was funded by the GEF-BIRF 28385/AR (A-

CB-51), CONICET (PIP 5301), ANPCyT (PICT 34111)

and DID-UACH (S2005-12) ) proje cts. C.M. and P.E.P

are Research Members of the National Research Council

of Argentina (CONICET).

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eee uw de tres especies de nudibranquios presentes en la

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Gavagnin, M., N. Uneur, F. ¢

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lusean Studies 68: 345-351,

THE NAUTILUS 121(3):146-158, 2007

Page 146

Two new modern records of the southern oyster drill

Stramonita haemastoma floridana (Conrad, 1837) in

Chesapeake Bay, USA

Juliana M. Harding!

Department of Fisheries Science

Virginia Institute of Marine Science

Gloucester Point VA, 23062 USA

[email protected]

M. G. Harasewych

Departinent of Invertebrate

Zoology, MRC-163

National Museum of Natural History

Smithsonian Institution,

Washington, DC 20013-

[email protected]

P.O. Box 37012,

7012 USA

ABSTRACT

Live southern oyster drills, Stramonita haemastoma floridana

(Conrad, 1837), have been collected from two Chesapeake Bay

western shore tributaries. Four specimens were collected be-

tween Brown Shoal and Thomas Rock in the lower James River

in February 2005. Thirteen live southern oyster drills were

collected from Back River in April 2006, Identification of these

drills as Stramonita haemastoma floridana has been confirmed

using DNA bar-coding data. Southern oyster drills collected in

Chesapeake Bay are genetically nearly identical to populations

from the Atlantic coast, and differ signific: uitly from popula-

tions from the Azores and from the Gulf of Mexico. These

collections mark the first reported records of live southern oys-

ter drills from within the Chesapeake Bay estuary. It is un-

known if these drills represent isolated introductions or expan-

sions of the northern range this species. Water temperature

patterns in Chesapeake Bay and the Mid-Atlantic Bight from

1990-2005 are similar to those observed in the late 1950s when

Stramonita haemastoma floridana was first found living in

Chincoteague Bay, Maryland.

Additional Keywords: Range extension, water temperature,

Cape Hatteras, zoogeography, DNA bar-coding

INTRODUCTION

The southern oyster drill Stramonita haemastoma flori-

dana (Conrad, 1537) is a predatory gastropod typically

found in association with populi itions of the oyster Cras-

sostrea virginica Gmelin, 1791 (e.g., Burkenroad, 1931;

sutler, 1955), A population from the Atlantic coast of the

United States was described as Purpura floridana Con-

rad, 1837, but later regarded to be a subspecies of the

Author for correspondence

broadly ranging Stramonita haemastoma (Linnaeus,

1767), a morphologically diverse taxon that has been re-

ported to span the temperate and tropical coasts of the

eastern and western Atlantic as well the eastern Pacific.

The name Stramonita haemastoma hacmastoma (type

locality: Tenerife, Canary Islands, fixed by Clench, 1947:

76) has been applied to the populations ranging from the

Channel Islands southward to Senegal, the western

Mediterranean, the Madeira, Canary and Cape Verde

Islands in the eastern Atlantic (Clench, 1947: 76; Poppe

and Goto, 1991: 141), the Azores (Morton et al., 1998),

Trinidad southward to Uruguay in the western Atlantic

(Clench, 1947: 76), and from Baja California southward

to Peru in the eastern Pacific by some (e.g., Clench,

1947) but not all (e.g., Keen, 1971: 549) researchers.

Stramonita hae Se floridana (type locality: Hy-

poluxo Island, Lantana, Florida, see Clench, 1947: 77)

was reported to range from North Carolina southward to

Yucatan and the West Indies, and throughout the Ca-

ribbean as far south as Trinidad (Clench, 1947: 76),

extending to Brazil (Abbott, 1974: 180), The name oe

monita hae smastoma canaliculata (Gri Ly, 1839) (type lo-

cality originally but erroneously listed as China) has been

applie 1d to a distinctive morphological variant prevalent

throughout the Gulf of Mexico (e.g., Abbott, 1974: 180).

Some three dozen names have been variously partitioned

and synonymized for the many geogr iphically circum-

scribed shell phe ce be longing to the Stramonita

hacimastoma complex (see Rose abe ra, 2005).

North American records of Stramonita have generally

been divided into two subspecies, Stramonita haema-

stoma floridana from the Atlantic coastline, and Stra-

monita haemastoma canaliculata from the Gulf of

Mexico; such division is based on shell characters (Ab-

bott, 1974: Butler, 1985). Stramonita haemastoma

canaliculata has been differentiated morphologically on

J. M. Harding and M. G. Harasewych, 2007 Page 147

Figures 1-9. Stramonita spp. 1-6. Stramonita haemastoma floridana (Conrad, 1837). 1. USNM 618840, Willis Wharf, Hog Island

Bay, Northumberland Co., Virginia, trapped in 2.4 m depth. September 15, 1955, ex-F. W. Sieling. 2. USNM 1091020, oyster reefs

just below Deep Creek, Lower James River, Virginia, in approximately 3 m, February 2005. 3. Male dnd, female specimen, USNM

1091021. Back River, Virginia, in commercial crab pots set in 3 m, April 2006. 5. USNM 416874, Fort Macon, North Carolina. 6.

USNM 1099258, Sand bar on north side of Fort Pierce Inlet, E of Little Jim Island, Ft. Pierce, Florida. 6 August 2004. 7. Stramonita

canaliculata (Gray, 1839), USNM 1099259, St. Andrews Bay, Florida, rip-rap near campsites. 8. Stramonita haemastoma haysae

(Clench, 1927), USNM 568834, Barataria Bay, Louisiana. 9. Stramonita haemastoma haemastoma (Linnaeus, 1767), USNM

1099260, intertidal rocks, Vila Franca do Campo, Sao Miguel, Azores, 26 July 2006.

Page 148

THE NAUTILUS, Vol. 121, No. 3

Atlantic Ocean

Figure 10. Map of the Atlantic Ocean depicting the southeastern coast of the United States, Chesapeake Bay region [inset], and

the Azores Islands, showing collection sites for Stramonita specimens and water-temperature data. Sites are identified as follows:

Delaware lightship and buoy stations (DE), Chincoteague, VA (A1), Chesapeake lightship and buoy stations (C), Fort Pierce Inlet,

FL (A2), St. Andrew's Bay, FL (A3), Pensacola, FL (A4), and Sao Miguel, Azores (A5). Within the Chesapeake Bay region

Chincoteague, VA (B1), Maryland DNR monitoring station NBM1S49 (B2), Maryland DNR monitoring station NBM 1301 (B3),

Virginia Institute of Marine Science, Gloucester Point, VA (B4), Back River, VA (B5), James River near Deep Creek (B6).

the basis of its larger size, presence of strong, rugose

shoulder nodules, and its deeply channeled suture. Sev-

eral authors (e.g., Butler, 1954; Gunter, 1979: Walker,

1982) had reported that differences in shell morphology

are neither consistent nor concordant with geographical

patterns and concluded that subspecific distinction was

not warranted. More recently Liu et al. (1991) con-

firmed that characters of shell and radula were not taxo-

nomically informative, but demonstrated that Stramonita

from the northern Gulf of Mexico could be differentiated

from populations of the Atlantic coast at “a level that is

characteristic of congeneric species” using allozyme elec-

trophoresis. Other recent studies have explored the ef

fectiveness of the east Florida ecotone as a barrier to

gene flow between Atlantic and Gulf coast populations

for a variety of taxa (for reviews, see Avise, 2000; Wise et

al., 2004). Vermeij (2001; 701) reviewed the Recent and

fossil species of Stramonita, and suggested that the Stra-

J. M. Harding and M. G. Harasewych, 2007

Table 1. Samples of Stramonita spp. used in this study.

Taxon Locality

Page 149

Voucher Number of

(1) Pensacola, FL

St. Andrew's Bay, F

Sao Miguel, ees

Stramonita canaliculata

Stramonita haemastoma haemastoma

Stramonita haemastoma floridana

Deep Creek, VA

Back River, VA

Ft. Pierce Inlet, FL

specimens specimens GenBank

USNM sequenced accession number

SSST09 1 US6330

1099259 1 EU073061

1099260 2 EU073051—EU073052

1099258 2 EU73053-EU073054

1091020 3 EU073055—E U073057

1091021 3 EU07305S—E U073060

" Sequence data from Harasewych et al., 1997

monita haemastoma complex may consist of as many as

four western Atlantic and two eastern Atlantic species.

He also noted similarities (bifid crenulations along the

outer lip) between some specimens of S. canaliculata and

S. bifida Vermeij, 2001, from the Cantaure Formation

(early Miocene, Venezuela).

Although the northern limit of Stramonita along the

eastern United States has generally been reported as Or-

egon Inlet, North C selina (Wells and Grey, 1960; But-

ler, 1985), living individuals (Figure 1) and recent his-

cee specimens (empty shells) were collected from lo-

cations in the Maryland and Virginia waters of

Chincoteague Bay (Figure 10, Al) in 1955 and 1956

(Sieling, 1955; 1960), extending the northern range

boundary for this animal along the US Atlantic coast.

Sieling (1960) suggested that the southern 6yster drills

were ceded. into Chincoteague Bay with trans-

planted oysters. Subsequent surveys of Assate: ague Island

and Chincoteague Bay by Counts and Bashore (1991),

Table 2.

Homer et al. (1997), and Prezant et al. (2002) reported

living S. haemastoma floridana from these habitats as

febenily as 1996 (Prezant et al., 2002). Counts and

Bashore (1991) categorized S. haemastoma floridana as

“rare” a linked the relative decline in abundance and

distribution of this drill with the decline in local oyster

resources from 1960 to 1989. Prezant et al. (2002) report

living S. haemastoma floridana in their survey and col-

lected specimen(s ) from Memorial Park, Maryland.

Living species ’ Stramonita have not previously been

reported from the interior of the Chesapeake Bay (An-

drews, 1956; Wass, 1972) in modern time, although Sh.

canaliculatum had occurred in Chesapeake Bay, Mary-

land, and S. h. floridana had reached southern New Jer-

sey during the exceptionally warm Sangamonian Stage

(80, 000 to ca. 220,000 yr BP) of the Pleicracene | Petuch

1997: 53). Ruiz et al. (2000) ) reported S. haemastoma as

an eceblehed aes in the Chesapeake Bay region on

the basis of the Chincoteague Bay, Maryland- Virginia,

Summary of water temperature data sources from 1950-2005 for Delaware (DE, Figure 10) and Chesapeake (C, Figure

10). Sea surface temperature (SST), bottom temperature (BT) and air temperature at the water surface (AT) were used.

Station Year(s) Data Station type Depth Source

Delaware/Winter Quarter 1955-60 SST, BT Winter Quarter lightship 24-29 m ]

1961-70 SST, BT Delaware lightship 30 m 2

1970-S4 SST One degree quadrangles NA 3, 4,5

Chesapeake 1984-2005 AT, SST NOAA buoy 44009 28 m 6

1950-197] SST Chesapeake lightship 20 m 7,8

1971-84 SST One degree quandrangles NA 3, 4,5

1985-2005 AT, SST NOAA buoy CHLV2 11.6m 9

1. Winter Quater lightship data archive. East Coast USCG lightship/Lightstations. MBLWHOI Library data archives, Woods

Hole, MA. http://dlaweb.whoi.edw/lightship/lightships_winterqtr.html

2. Delaware lightship data archive. East Coast USCG Lightship/Lightstations. MBLWHOI Library data archives, Woods Hole,

MA. http://dlaweb.whoi.edu/lightship/lightships_delaware_html

3. 1966-1974. The Gulf Stream. U.S. Naval Oceanographic Office, Vols 1-9.

4. 1975-1980. gulfstream. U.S. Dept. of Commerce, NOAA, National Weather Service. Vols. 1-6.

5. 1981-94. Oceanographic Monthly Summary. U.S. Dept. of Commerce, NOAA, National Weather Service/National Earth

Satellite Service. Vols. 1-14.

6. Delaware Bay, Buoy 44009. http:/Avww.nodc.noaa.gov/BUOY/44009. html.

7. Bumpus, D. 1957. Surface water temperatures along Atlantic and Gulf coasts of the United States. U.S. Fish and Wildlife

Service Special Scientific Report—Fisheries No. 214.

5. Chesapeake lightship data archive.

MA. http://dlaweb.whoi.edu/lightship/lightships_chespstn.html

East Coast USCG Lightship/Lightstations. MBLWHOI Library data archives, Woods Hole

9. Chesapeake Light. VA, Buoy CHLV2. http:/Avww.node.noaa.gov/BUOY/chlv2. html.

Page 150

THE NAUTILUS, Vol. 121, No. 3

SS SS SS eS eee

[ 2005: James River, VA

WB 2006: Back River, VA

0 } lI

[tet

64 "66 68 70 72 74 76 78 80 "82 "ga Ps 88 "90 a ea

Midpoint of shell length class (mm)

Number of snails collected

Figure 11. Length frequency distribution of live Stramonita

haemastoma floridana specimens collected from Back River,

VA (2006) and the James River, VA near Deep Creek (2005).

and Hog Island Bay, Virginia, collections reported by

Sieling (1960) (P. Forowolk Smithsonian Environmental

Research Center, personal communication). Distribution

patterns of many molluscan species along the US Atlantic

coast are directly related to water temperature with Cape

Hatteras acting as a natural zoogeographic boundary

(Franz and Mervill, 1950a, b). Wells and Gray ( (1960)

reported S. h. floridana on two subtidal shipwrecks south

of Cape Hatteras, North Carolina, but found none on the

four shipwrecks they examined north of Cape Hatteras.

Wells (1961) described re soular collections of resident

S. h. floridana from inte srtidal oyster reefs in the vicinity

of Beaufort and Cape Lookout, North Carolina.

Southern oyster drills begin laying eggs at water tem-

peratures between 20 and 30°C (Stickle, 1999). At water

temperatures less than 10-12.5°C southern oyster drills

stop feeding on bivalves, bury into the substrate, and

become inactive (Bulter, 1954; Gunter, 1979; Garton and

Stickle, 1950; Stickle, 1999) until water temperatures

rises. While the upper lethal thermal limit for southern

oyster drills is 35 to 40° C (Brown and Stickle, 2002), the

lower lethal thermal limit for southern oyster drills is

unknown.

This report documents the first collections of living

Stramonita haemastoma floridana from within the

Chesapeake Bay estuary. We characterize both oe of

southern oyster drills collected in the Che ssapeake Bay

genetically (cytochrome c oxidase I “DNA bar-coding”)

aad compare ‘these data to se quences derived from rep-

resentative spe cimens from the southeastern (Indian

River Inlet) and northwestern (St. Andrew's Bay, Pensa-

cola) coasts of Florida, as well as from samples from Sao

Miguel, Azores. The presence of these drills in Chesa-

peake estuarine habitats is analyzed in the context of a

long term (1950-2005) coastal ‘bottom water tempera-

ture data set and two independently collected estuarine

bottom water temperature data sets (Chincoteague Bay,

Maryland and Virginia, 1951-56; McGary and Seiling,

1953; Seiling, 1957; York River, Virginia, 1986-2005:

VIMS, 2006) in order to relate the observed Chesapeake

collections with environmental/climate conditions in

known drill habitats along the Atlantic coast and in Chin-

coteague Bay.

MATERIALS AND METHODS

Sample Collection: Four living S. haemastoma (Fig-

ure 2) were collected in the lower James River, Virginia,

in the Che ssape rake Bay from oyster (Crassosteea vir

ginica) reefs just below Deep Creek (Figure LO, BG) at a

depth of approximately 3m in Fe bruary 2005 and were

tured in to the Virginia Institute of i Science

(VIMS) rapa whelk [Rapana venosa (Valenciennes,

1846)| bounty program (see Harding a Mann, 2005,

for bounty program details). All four Stramonita speci-

Table 3. Nucleotide (above diagonal) and amino acid (below diagonal) ) ditfe srences between samples in the portion of the cytochrome

c oxidase I gene

obsoleta {Genbank NC 007781]})

(591 aligned positions, corresponding to positions 73 to 664 in the complete mitochondrial genome of Iyanassa

Sample l 2 3 5 6 7 8 9 10 lI 12

1) S. canaliculata Pensacola, FL — 0) 126 126 123 123 123 122 124 124 123 122

2) S. canaliculata St. Andrew’s, FL 0) — 126 126 123 123 123 122 124 124 123 122

3) Sh. h aemastoma Azores | s S — 5 56 55 55 58 56 56 56 56

1) S. h. haemastoma Azores 2 i 7 | — 57 56 56 61 57 57 DT 57

5) S. h. floridana Ft. Pierce Inlet, FLI 7 7 3 2 ~ 2, 2 7 3 5 3 3

6) S. h. floridana Ft. Pierce Inlet, FL2 7 if 3 2 0) 0 5 | 3 | |

7) S. h. floridana Deep Creek, VAI 7 7 3 2 0 0 — 5 | 5 | |

5) S.h. floridana Deep Creek, VA2 S S 2 3 | | | —- 6 S 6 6

9) Sh. floridana Deep Creek, VA3 7 4 3 2 0 0 0 | = { 2 2

10) S. hh. floridana Black River, VAI v4 7 3 2 0 0 0) | 0 — { |

11) S.h. floridana Black River, VA2 7 7 3 yy) 0 0 () | 0) 0 = 2

12) S. h. floridana Black River, VA3 7 7 3 2 0) 0) 0 | 8) 0 0

J. M. Harding and M. G. Harasewych, 2007

Page 15]

: licul

S. canaliculata PenFL Gulf of

Mexico

S. canaliculata SAnFL

S. haemastoma

haemastoma Azores 1 Azores

S. haemastoma

haemastoma Azores 2

S. haemastoma Fort

floridana FPIFL1 Pierce

S. haemastoma Inlet, FL

floridana FPIFL2

S. haemastoma

floridana DCVA1

Deep

S. haemastoma Creek

floridana DCVA2 VA ,

S. haemastoma

floridana DCVA3

S. haemastoma

floridana BRVA1 Back

S. haemastoma River

floridana BRVA2 VA

S. haemastoma

floridana BRVA3

Figure 12. Strict consensus of four most parsimonious trees

(1 = 176; ci = 0.955; ri = 0.960) resulting from an exhaustive

search using maximum parsimony (PAUP 4.0b10) based on 591

bp of cytochrome c oxidase I sequences. Branch lengths from

one of the four most parsimonious trees are above the

branches, bootstrap (bold) and jackknife (bold italic) propor-

tions are below the branches.

mens were collected in a single oyster dredge tow. Thir-

teen living S. haemastoma (Figures 3-4) were collected

in Back River, Virginia, (Figure 10, B5) in April 2006 and

were also turned into VIMS via the rapa whelk bounty

program. The 2006 collection was made using commer-

cial crab pots (wire mesh cubes with approximately 0 0.5m

sides) baited with northern quahogs |Mercenaria merce-

naria (Linnaeus, 1758)| deployed at a depth of 3 m. Shell

lengths of specimens (mm, maximum dimension tip of

the spire to bottom of the siphonal canal) were measured

upon receipt at VIMS. Voucher specimens from both

Chesapeake collections have been deposited in the Na-

tional Museum of Natural History. Smithsonian Institu-

tion (USNM, Table 1) ;

DNA Bar-coding: Three specimens from each Chesa-

peake collection were frozen and transported to the labo-

ratory, where DNA was extracted from red buccal

muscles of each animal using Qiagen DNEasy kits ac-

cording to the manufacturer's protocol. A portion of the

mitochondrial gene for the DNA bar-coding region of

the cytochrome c oxidase I gene was amplified using

Sigma JumpStart eee Re sadly Mix and Meyer's (2003)

degenerate sd versions of the Folmer et al. (1994) HCO

and LCO primers for the samples listed in Table 1. The

resulting PCR products were cleaned using magnetic

beads [Agencourt, manufacturer's protocol] and se-

quencing reactions were run on ABI 3730 sequencers

that were set up according to manufacturer's instruc-

tions. Sequences were manually checked/corrected and

assembled using Sequencer™ 4] (Gene Codes C Jorp.)

then aligned wih Clustal W (Higgins et al., 1994) using

default settings, and their pace analyzed using

PAUP 4.0b10 (Sivoftord. 2002). Pairwise comparisons of

sequences and amino acids were performed using Mega

3.1 (Kumar et al., 2004).

Water Temperature Data: CoastaL HaAsirats: Wa-

ter temperature data from 1950-2005 were used to de-

scribe mean monthly bottom water temperatures (BT,

C°) for two stations in the Mid-Atlantic Bight (Delaware/

Winter Quarter and Chesapeake) (Figure 10, Table 2).

Daily sea surface and bottom water temperature data

from the Delaware/Winter Quarter (1961-70) and

Chesapeake (1958-71) lightship stations were used to

calculate the average monthly observed difference be-

tween sea surface tempe rature (SST) and bottom water

temperature (BT) on a site-specific basis. The observed

average monthly SST-BT differences from the daily

lightship data were used to estimate average monthly B’ r

at these sites during years after 1971 when only SST was

available (Table 2).

ne air temperature (AT) and SST data from

NOAA buoys (1985-2005) were used to calculate the

observed dteronoe between average monthly AT and

average monthly SST for Delaware Light and Chesa-

peake I Light. The observed av erage aioe AT-SST dif-

ferences from the buoy data were use sd to estimate

hourly SST for these sites during months after 1954

when only AT was available.

Average residuals for annual BT from the long-term

(1950-2005) average annual BT were calculated for each

site in which at least 9 months of data were available.

Monthly average bottom water temperatures (with stan-

dard error of the mean) were calculated for each of 12

months for all sites. Monthly BT estimates from 1970 to

1983 (Delaware) and 1984 (Chesapeake Light.) use a

single published monthly average ( Table 2) while

monthly BT estimates from NOAA buoys (Table 2) are

averages calculated from hourly readings with n values

>400 per month.

Estuarine Habitats: Water temperature data col-

lected at Public Landing, Maryland, in Chincoteague

Bay from McGary and Sieling (1953) and Sieling (1957)

were used to calculate average monthly water tempera-

tures (with standard error of thé mean) during the period

1951-1956. Modern (2001-2004) water temperature

data from Chincoteague Bay stations south of Robins

Marsh (XBMS8149) (Figure 10, B2) and near the MD-VA

border (XBM1301) (Figure 10, B3) were obtained from

Page 152

THE NAUTILUS, Vol. 121, No. 3

A. Delaware

Feb 1961 - Dec 1970

102, 138170 148

1 2 3 4 5

B. Chesapeake

Jan 1958 - Dec 1971

Average monthly SST-BT difference (°C)

n= 202 188 174 180 169

1 2 3 4 6)

eae e 13.

162-164

7 8 9g 10 11 12

) e

145 197 159 173

7 8 i) 10 11 12

Month

ae rage monthly difference between sea surface temperature (SST) and bottom water temperature (BT) from lightship

data (Table 2) for Delaware (A) and Chesapeake (B) lightships. Error bars indicate standard error of the mean. N values (number

of sail, SST-BT pairs used to estimate the SST-BT differences) at each site are presented above the X axis in each panel.

the Maryland De Lain nt of Natural Resources Eyes on

the Bay web site (http://mddnr.chesape: akebay. net/

bay_cond/). Water de pth at all (historical and modern) of

these Chincoteague Bay sites is 3 m or less. Seiling

(1954) reported less than a degree Celsius variation be-

tween surface and bottom water temperatures and the

data presented in McGary and Seiling (1953) and Seiling

(1957) from. sites throughout the estuary confirm the

well-mixed nature of these non-channel habitats.

The Virginia Institute of Marine Science (VIMS) at

Gloucester Point, Virginia (Figure 10, B4) maintained a

water temperature monitoring station from 1986 through

September 18, 2003, (arrival of Hurricane Isabel) that

recorded bottom water temperatures (°C) ata depth of

2-3 m. The VIMS Molluscan Ecology program has main-

tained an environmental monitoring station located

within 200 m of the original VIMS station since January

2005 that records bottom water temperatures (°C) at

depths of 2 m. Hourly water temperature averages were

obtained from the VIMS data archive (1986 to Se p 2003)

and the VIMS Molluscan Ecology program environmen-

tal data archive (2005) and used to calculate monthly

residuals from the 1986-2005 mean and average monthly

bottom water temperatures (with standard error of the

mean).

RESULTS

aa ad The southern oyster drill specimens col-

lected from the James River in Febru: wy 2005 ranged in

size from 66.4 to 75.6 mm shell leneth with an average

shell length of 69.0 + standard error of the mean 2.9]

mm. Sointhe rm oyster drills collected in April 2006 from

Back River had shell le neths ranging from 73 to S9 mm

with an average shell length of $3.3 + 1.41 mm. Size

frequency distributions for both collections ( Figure 11)

indicate that all specimens were adults (sexually mature;

Burkenroad, 1931; Butler, 1985) and represent the up-

per end of the size distribution typically found in Loui-

siana habitats (Brown and Richardson, 1987; Brown et

al,, 2004). These large individuals have probably reached

a size refuge from most local predators including blue

crabs (€ allincetes sapidus, see the work of Turra et al.,

2005, using C. eve preying on S. hacmastoma) and are

certainly capable of cating oysters >50 mm shell height

(Garton, 1986; Brown and Richardson, 1987).

J. M. Harding and M. G. Harasewych, 2007

A. Delaware

Jan 1984 - Dec 2005

n= 10261 10050 12342 11677 12880 12493 13543 14049 12911 13168 11876 11666

2 3 4 5

B. Chesapeake

Jan 1985 - Dec 2005

)

Average monthly SST-AT difference (C)

7 8 ) 10 11 12

n= 10317 9982 11059 10995 11258 12175 12422 11965 11671 11434 12023 11430

1 2 3 4 5

Figure 14.

(Table 2) for Delaware (A) and Ches: apeake (

7 8 9 10 11 12

Month

Average monthly difference be Ae en sea surface temperature (SST) and air temperature (AT

) from NOAA buoys

3) buoys. Error bars indicate standard error of the mean. N values (number of daily

SST-AT pairs used to estimate the SST-AT he at each site are presented above the X axis in each panel.

DNA Bar-coding: Comparisons of a 591 base-pair

portion of the auitechotd! il gene for cytochrome c oxi-

dase I from the samples listed in Table 1 revealed that

specimens of Stramonita collected within Ches sapeake

Bay were genetically very similar or identical to speci-

mens colle ecte 2d from Fort Pierce Inlet, on the southeast-

ern coast of Florida, but differed substantially from

Azorean samples and even more so from specimens from

the northwestern coast of Florida (Table 3). Maximum

parsimony analyses of the 144 phylogenetically informa-

tive sites using the exhaustive search algorithm yielded

four equally parsimonious trees (length = 176; ci = 0.955;

ri = 0.960). Figure 12 illustrates strict consensus tree of

these four trees, including results of bootstrap and jack-

knife analyses.

Water Temperature Data: =CoastTaL Hapirats: Aver-

age annual bottom water temperatures in the period

1950-2005 for the two stations along the US Atlantic

coast followed a latitudinal trend and were lower

9.81°C. SEM = 0.15, 600 months of data) at Delaware

than at Chesapeake (12.30°C, SEM = 0.21, 596 months

of data). The differences between SST and BT recorded

by the lightships (approximately 1956-71) show the sea-

sonal deve thom nt of the thermocline at Delaware and

Chesapeake beginning in April and persisting until Oc-

tober with the most pronounced differences between

surface and bottom water temperatures occurring in July

and August (Figure 13) when these stations experience

surface temperatures that are at least 7° C higher than

bottom temperatures. Air temperatures re eee d by =

Delaware and Chesapeake NOAA buoys (T: able 2

are 1-2° warmer than SST from April through July (Fig.

ure 14).

Examination of annual average residual bottom water

temperatures from the average long-term (1950-2005)

bottom water temperatures at coastal stations (Figure

15) shows that both sites see above average wa-

ter temperatures during the late 1950s. Multiple con-

secutive years between 1970-1980 and, again, in the pe-

riod 1995-2005. Estimated bottom water te mperatures

for 2002 were among the highest observed during the

period 1950-2002 for Delaware and Chesapeake Bays

Delaware and ¢ Jhesapeake, both north of ¢ vape Hatteras,

experience water temperatures of 10°C or less eight and

five months out of the year, respectively (Table 4A

Page 154 THE NAUTILUS, Vol. 121, No. 3

—_

2 A. Delaware

—_

8 2

€

2 1

co}

5 O

aI -1

j=)

(o>)

= -2

je)

3 (3

oO on TOO ON TOWONTSOWDWDIONTGOAOGOA TLC OM]M OA +t

ag ouonwnonwnonwnooeDnoogoaorRrenRRRRWWAADANAD DD DAD OS OF

DADDADAADA AAD AAA ADAAAADAAAAAAADAADAADAA DAA OO AS

Sar oe el er ool, so ei, a Ton, a Se, od, Seal Sen a a ee eR co on oe, ec ed Te) |

~ B. Chesapeake

ae P

—

o 2

= i

fe)

co}

> 0

—

So -1

fo)

2 2

(eo)

Go -

g -3

Ke)

oO oNWNnNWONMONWOODADAARe RE RERR DWWADWDADWANANBDDDDHD OO OS

w DOOD ADADAADA ADA AA ADA A DAA AAAAAADAAAAIAAADAA HA OOO

TS SS YS wwe SSS SS SS SE SS SS SS SS SS SE SES ON NN

Year

Figure 15. Annual average residual bottom temperature (BT) from the long term (1950-2005) average bottom temperatures for

Delaware (A) and Chesapeake (B). Long term average data for each site are presented in Table 4. The error bars represent the

tandard error ol the mean mM degrees ¢ elsius

J. M. Harding and M. G. Harasewych, 2007

Table 4. Summary of average monthly bottom water temperatures for coastal stations (A, 1950-2005) and estuarine stations (B)

shown in Figure 10 and discussed in text. Standard error of the mean for each value is given in parentheses.

Month Delaware Chesapeake

Jan > as (0,22) 6.96 (0.24)

Feb 5 (0.20) 5.60 (0.23)

Mar 5 58 (0.18) 6.12 (0.21)

Apr 7.02 (0.15) 8.31 (0.20)

May bora 29) 11.02 (0.21)

Jun 3 (0.18) 14.34 (0.21)

Jul 9 45 (0.15) 14.93 (0.20)

Aug 10.69 (0.20) 16.72 (0.25)

Sep 14.62 (0.29) 20.27 (0.20)

Oct 16.56 (0.20) 19.11 (0.21)

Nov 13.57 (0,15) 11.02 (0.21)

Dec 10.06 (0.18) 10.41 (0.23)

Chincoteague Bay, Chincoteague Bay, Chincoteague Bay, York River,

Public Landing, MD XBMS8149 XBM1301 Gloucester Point, VA

Month (1951-56) (2001-04) (2001-04) (1956-2005)

Jan 4.14 (0.36) 5.3 5.7 5.32 (0.36)

Feb 6.22 (0.33) 5.25 5.38 5.55 (0.47)

Mar 9.76 (0.35) 9.65 99 8.46 (0.33)

Apr 14.57 (0.43) 15.53 15.85 13.61 (0.30)

May 20.1 (0.38) 18.65 18.87 18.86 (0.26)

Jun 24.08 (0.38) 2120 26.33 23.91 (0.26)

Jul 27.77 (0.32) 26.1 25.85 26.96 (0.18)

Aug 27.2 (0.26) 27.42 27.65 27.11 (0.18)

Sep 23.6 (0.40) 22.95 22.93 23.99 (0,59)

Oct 17.32 (0.50) 14.55 14.6 19.28 (0.33)

Nov 10.72 (0.38) 15.1 15.68 13.56 (0.24)

Dec 5.64 (0.39) 5.2 5.5 $8.36 (0.54)

EsTUARINE Hapirats: Annual average residual bottom

water temperatures from the York River at Gloucester

Point, Virginia were higher than the 1986-2005 annual

average (16.23 + 0.56°C) in 10 of the 1S years for which

data are available (Figure 16) including 2005.

The relatively shallow (3 m or less) estuarine habitats

examined in Chincoteague and Chesapeake Bays expe-

rience a seasonal average monthly water temperature

cycle appropriate for their latitudes (Table 4B) with wa-

ter temperatures less than 10°C observed December

through March. The same pattern was observed in Chin-

coteague Bay during 1951-1956 (Table 4B).

DISCUSSION

The collections of live Stramonita from the James River

(2005) and Back River (2006) mark the first modern

record of this predatory gastropod from the Chesapeake

Bay interior. The morphology of these specimens, par-

ticularly their larger size, deeply channeled suture, and

the presence of strong, rugose shoulder nodules in the

Back River samples, is similar to that of some populations

from the northern Gulf of Mexico [particularly the form

named Stramonita haemastoma haysae (Clench, 1927

(Figure 8)|. This, in tum, has raised questions about

whether these animals were introduced into Chesapeake

Bay, and the possible source of such introductions. How-

ever, the results of the “DNA bar-coding” study clearly

uicaeate that the Chesapeake Stramonita haemastoma

floridana are genetically very similar to, and in one case

indistinguishable from, a population of Stramonita hae-

mastoma floridana trom southeastern Florida. There

seems little doubt that the source of the Chesapeake

Stramonita haemastoma floridana populations lies along

the eastern seaboard of the United States. Less clear is

whether these populations were introduced into Chesa-

peake Bay with oysters, as had been suggested by Sieling

(1960) for the Chincoteague Stramonita, or if their pres-

ence is due to a northward expansion of the range of

Stramonita haemastoma floridana due to warmer ocean

temperatures.

The “bar-coding” data also indicate that the east-coast

Stramonita are well differentiated from, yet more similar

to Stramonita from the Azores than to specimens from

the northern Gulf of Mexico (Figure 12), Provisional

taxonomic consequences of this study are to recognize

the Azorean populations as Stramonita haemastoma hae-

mastoma, to retain the usage of Stramonita haemastoma

Page 156

THE NAUTILUS, Vol. 121, No. 3

3 York River, Gloucester Point, VA

-2

Residual from 1986-2005 average bottom temperature (C)

(jo)

1950

1952

1954

1956

1958

1960

1962

1964

1966

1968

1970

1972

1974

Figure 16. Annual average residual bottom temperature

(16.23 °

of the mean in degrees Celsius.

floridana for the populations from the eastern United

States, and to distinguish the populations from the north-

erm Gulf of Mexico as Stramonita canaliculata,

cated by Liu et al. (1991). Detailed analyses of this spe-

cies complex over its entire ge ographic range are clearly

required to better unde rstand its bioge ography and tax-

as ac lve )-

onomy.

Water temperature patterns in Chesapeake Bay habi-

tats and along the coast of the lower Mid-Atlantic Bight

the late 1990s have average as were

water temperature trends observed in the period 1955—

1957 when live Stramonita were collected on the Atlantic

coast of Maryland and Virginia (Sieling, 1960). Although

the lower thermal lethal limit for southe ‘mm oyster Sills is

the

om water temperatures in Chincoteague Bay during the

1951-56 are within 1 to 2°

Fibs ape aa observed in the lower York River from

1986

ave seasonal te mperature regime »5 conduc ‘ive to survivi al

since been above

unknown, fact that observed seasonal trends in bot-

yeriod C of bottom water

2005 sugge sts that Che 'sapei ake Bay tributaries may

Living southern oyster drills were col-

Bay by Prezant et al. (2002)

of this animal

ected from Chincoteague

during surveys between 1991 and 1996, some 40 years

after the initial collection (Sieling, 1956). The very low

sopulation levels (“rare”, Counts and Bashore, 1991) of

Bay

in local

L991) rather than

southern oyster drills observed

19SS—89 have

ovster re

in’ Chincoteague

been attributed to a decline

Ince

sources (¢ ounts and Bashore

owmoonrwtowdoondriewrTraoewWdeiondrst

KER WAWDWDAWDAADADABDWDIWAABDHA GOO

DAA AADAAADAA DAA HD OO OO

bl ee ee el ee el ee ee ee ee oe

Year

(BT) from the long term (1986-2005) average bottom temperatures

C, standard error of the mean 0.56 °C) recorded at Gloucester Point, Virginia. The error bars represent the standard error

unfavorable water temperatures. The tributaries of the

lower Ches: ape rake Bi My have resident populations of bi-

valves commonly eaten by Stramonita including mussels

and oysters ( (Butler, 1985: Garton, 1986: Brown and Ri-

chardson, 1987). Several of these tributaries are also sites

of focused oyster restoration efforts that incorporate ad-

dition of either spat on shell or broodstock oysters fur-

ther expanding the potential prey field for southern oys-

ter drills.

If these collections represent the beginning of an in-

vasion into Chesapeake Bay tributaries, the persistence

of Stramonita haemastoma floridana in Chesapeake Bay

will be dictated by the rmal tolerances as subjected to

seasonal temperature cycles, while the geographic distri-

bution will be set by si linity tolerances of adult and lar-

vae. Adult Stramonita haemastoma may survive at salini-

ties as low as 5-7 ppt (Gunter, 1979; Stickle, 1999). Stra-

monita haemastoma egg capsules survive and release

viable larvae at salinities down to at least 7.5 ppt and

possibly as low as 3.5 ppt (Stic ‘kle, 1999). Veliger larvae

may survive up to 5 days when exposed to LO ppt (Wells,

1961) but Roller and Stickle (1989) reported very low

survival at salinities less than 15 ppt. In June 1972, heavy

rains from Tropical Storm Agnes in combination with

summer temperatures killed essentially all of the native

oyster drills [Urosalpinx cinerea (Say, 1822), Eupleura

caudata (Say, 1$22)| in the upper re saches of Ches: apeake

Bay tributaries and reset the distributional range of the

J. M. Harding and M. G. Harasewych, 2007

Page 157

native drills to the high salinity, lower reaches of the

Chesapeake Bay where natural oyster populations cur-

rently persist in only limited regions and numbers. Re-

establishment of these native gastropod species to their

former habitats is occurring slowly over decadal time

scales and is limited by the Trek of planktonic larvae for

both Urosalpinx cinerea and Eupleura caudata. Reinva-

sion of their historic habitats by the native drills is con-

founded by the fact that the historically widespread dis-

tribution of oyster reef habitat has been drastically re-

duced in areal cov erage since the early 1960s by a

combination of diseases and environmental degr: adation.

Competition for the niche vacated by the native drills

during Tropical Storm Agnes already includes one large

non-native gastropod, the veined rapa whelk (Rapana

venosa) (Harding and Mann, 1999; 2005), which has

planktonic veliger larvae like Stramonita haemastoma

and is equally long- lived. Regardless of how these Stra-

monita haemastoma got to these C hesapeake tributaries,

the presence of acide: individuals in these tributaries adds

yet another competitor for this vacant niche as well as

additional predation pressure on local oyster resources.

Successtul invasion of a habitat requires a breeding

population and the presence of all life history stages in

the new habitat (Williamson, 1996). Southern oyster

drills live from 5-20 yr in Florida (Butler, 1985) ‘with

generation times on the order of 12 months (Burken-

road, 1931; Butler, 1954). The small number of speci-

mens collected to date combined with the absence of

these animals in annual fishery independent surveys of

oyster reefs in the James River conducted by Harding

and colleagues at VIMS and the Virginia Marine Re-

sources Commission since the early 1990s may be an

indicator that the observed southern oyster drills speci-

mens represent small, isolated introductions that have

not yet established local populations.

ACKNOWLEDGMENTS

Special thanks are extended to all local watermen who

have donated Stramonita haemastoma and Rapana

venosa to the VIMS research programs. Karen Caposella,

Christina Conrath, Meredith Fagan, Adriana Picariello,

and Matt Robinson assisted with local whelk collections

by VIMS. Dr. Gregory Herbert, University of South

Florida, kindly provided specimens of Stramonita from

St. Andrew’s Bay, Florida. Azorean specimens were col-

lected during a workshop in Vila Franca do Campo, Sao

Miguel, Azores, hosted by The University of the Azores.

The Workshop was a joint organization of Sociedade

Afonso Chaves and the Department of Biology of the

University of the Azores. Support from FLAD (Portu-

guese- American Foundation for Dev elopment) is grate-

fully acknowledged. Specimens from the Indian River

Inlet were collected during a workshop supported by the

Smithsonian Marine Station at Fort Pierce. Dr. Katherine

Farnsworth, Indiana University of Pennsylvania, provided

GIS assistance with maps. This is Contribution Number

2859 from the Virginia Institute of Marine Science, Glouc-

ester Point, Virginia, and Smithsonian Marine Station at

Fort Pierce Contribution Number 696.

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THE NAUTILUS 121(3):159-162, 2007

Page 159

Brachycythara beatriceae, a new species from the

Alboran Sea and the eastern Atlantic Ocean

(Gastropoda: Neogastropoda: Conidae)

Paolo Mariottini

Dipartimento di Biologia

Universita di “Roma Tre”

Viale Marconi 446, 00146

Roma, ITALY

[email protected]

ABSTRACT

Based on shell characters, Brachycythara beatriceae, a new

gastropod species of the family Conidae from the Alboran Sea

and the eastern Atlantic Ocean, is here described. The new

taxon, represented by five specimens dredged along the Span-

ish Mediterranean and the West Sahara coasts, is compared

with the similar Brachycythara atlantidea (Knudsen, 1952), a

species that occurs in the same geographical area. This new

species is conchologically distinct and can be identified on the

basis of its teleoconch shape, rib count and microsculpture,

protoconch morphology, and shell color.

Additional Keywords: Mangeliinae, turrid, shell morphology.

INTRODUCTION

Traditionally, the epithet “turrid” has been used as a

general term referring to the numerous species belong-

ing to the family-group Turridae H. Adams and A. Ad-

ams, 1853, sensu lato. The new taxon described in the

article is assigned to the family Conidae Fleming, 1822,

subfamily Mangeliinae Fischer, 1883, genus Brachy-

cythara W oodring, 1928, in accordance with the revision

of the super rfamily Conoidea ( = Toxoglossa) Rafinesque,

1815, as proposed by Taylor, Kantor, and Sysoey (1993).

The genus was re-described by Powell ( 1966: 117), and

consists of small Recent and Tertiary species that aa 7

occur in the Caribbean area. Brachycythara has been

represented in the eastern Atlantic Ocean by only one

species, Brachycythara atlantidea (Knudsen, 1952) (see

Roldan and Otero-Schmitt, 1999), which has recently

been reported from the Alboran Sea by Smriglio et al.

2007). I had the opportunity to examine five shells of a

species that, in spite of their similarity with B. atlantidea,

could not be considered that species. These shells, col-

lected along the southern Spanish (Costa del Sol) and

West Sahara coasts, show features typical of the genus

Brachycythara. After a comparison with the similar and

sympatric B. atlantidea, I believe they represent an un-

described species.

Abbreviations used are: MZB, Laboratorio di Malaco-

logia, Museo di Zoologia dell Universita di Bologna,

Italy. Private collections cited in this article are: CS-PM,

Carlo Smriglio and Paolo Mariottini (Rome, Italy); SR

Stefano Rufini (Rome, Italy); FG, Franco Gubbioli

(Marbella, Spain). SEM photographs were carried out at

the LIME (Interdepartmental Laboratory of Electron

Microscopy), University “Roma Tre”, Rome, Italy.

SYSTEMATICS

Superfamily Conoidea Ratinesque, 1815

Family Conidae Fleming, 1822

Subfamily Mangeliinae Fischer, 1883

Genus Brac hycythara Woodring, 1928

Type Species: — By original designation, Cythara gibba

Guppy, 1896. Woodring, 1925, Carnegie Institute

Washington of Publications 385, p. 175.

Brachycythara beatriceae new species

(Figures 1-9; 13-20)

Description: Shell rather small, to 10.9 mm in length,

holotype 8.6 x 3.4 mm, biconical, elongate-fusiform,

solid, spire about half total height. Protoconch multispi-

ral, dome shaped, of 3-3.5 strongly convex whorls, first

1.5-2 whorls smooth, remainder wheds reticulated with

oblique axial costae crossed by spiral ribs of about equal

width; protoconch indicative of planktotrophic larval de-

velopment. Diameter of protoconch about 700-750 jum.

Protoconch-teleoconch transition not well marked. Te-

leoconch of 5-6 whorls, weakly angulate near middle of

spire, sutural ramp gently concave, whorl sides gently

convex; last whorl about 2/3 shell length. Axial sculpture

consisting of 7-S prominent, slightly opisthocline, flexu-

ous, and narrowly rounded axial folds: folds regularly

spaced, with much broader inte rspaces, ret aching from

suture to suture on spire but fading out after crossing the

Page 160

THE NAUTILUS, Vol. 121, No. 3

Figures 1-12. Shells of Brachycythara species. 1-3. Brachycythara beatriceae new species, holotype, 5.6 « 3.4 mm, MZB 31032,

Spain, Alboran Sea, off Marbella, 36°25’ N, 4°52’ W, 30-40 m. 4-6. Brachycythara beatriceae new species, paratype A, 10.9 x 4.5

mm, CS-PM, Spain, Alboran Sea, off Malaga, 36°33’ N, 4°22' W, 50 m, 7-9. Brachycythara beatriceae new species, paratype B, 9.3

3.7 mm, FG, coast of West Sahara, 50-60 m, 10-12. Brachycythara atlantidea (Knudsen, 1952), specimen L, 9.5 x 3.6 mm, FG,

coast of West Sahara, 30-60 m. Scale bars: 2 mm

shouldes slope and at about the middle of the base. Spira

sculpture of numerous, very fine threads that densely

alternate with bigger interspaces; the subsutural threa

shows well-marked axial denticles. At higher magnifica-

tion, it can be observed that each interspace consists Oo

several up to tive) row of rounded tiny granules, each

one linked axially to the uppel and lower thread by a very

Ine connection \pe rture narrow, ovate, about one thir

of the shell height Siphonal canal short, narrow, anc

opel Inne lip vith a moderately developed pariecta

| | Outer lip thin or variced according to the

we of growth, whether the lip coine ides with an axia

1 1 interspace Anal sinus marked, arcuate on

ope. Shell color white vith a brown band on

lower half of body whorl (juvenile and subadult shells

uniformly white); a darker spot present on the outer lip

at the boundary of the white and the brown colors

Type Material: Holotype (Figures 1-3), 8.6 « 3.4

min, MZB 31032, Spain, off Marbella, 36°28’ N 4°52’ W,

30-40 m; Paratype A (Figures 4-6), 10.9 x 4.5 mim, CS-

PM, Spain, off Malaga, 36°33" N 4°22" W, 30-40 m;

Paratype B, 8.3 x 3.1 mm, SR, Spain, off Malaga, 36°33'

N 4°22" W, 30-40 m; Paratype C (Figures 7-9; 13-20)

9.3 «3.7 mm, FG, West Sahara, 50-60 m: Paratype D

8.2 «x 2.6 mm, FG, West Sahara, 50-60 m

Other Material Examined: Shells of Brachycythara

atlantidea (Knudsen, 1952): Specimen A, 10.2 « 4.1 mm

P. Mariottini, 2007 Page 16]

te wed

Roe

Figures 13-28. Shell morphological details of Brachycythara species by SEM photographs. 13-14. Brachycythara beatriceae new

species, paratype B. 15-20. Details of the larval whorls and the shell sculpture. 21-22. Brachycythara atlantidea (Knudsen, 1952

spec. F. 23-28. Details of the larval whorls and the shell sculpture

Specimen B, 9.2 7 mm; Specimen C, 7.5 x 3.4 mm mens A—G, N are deposited in CS-PM collection, speci-

Specimen D, $8.2 x 3.6 mm; Specimens A—D from Spain, mens H—M in FG collection

off Estepona, 36°25’ N, 05°09" W, 150-250 m; Specimen

E. 7.7 x 3.4 mm, from Spain, off Adra, 36°45’ N, 03°01’

W. S0-150 m; Specimen F (Figures 21-28), 7.4 x 3.2

mm: Specimen G, 10.2 x 4.4 mm; Specimens F—G from Distribution: Alboran Sea (Costa del Sol Spain and

Spain, ott Malaga, 36°33’ N, 04°22’ W,50 m: Specimen eastern Atlantic Ocean (West Sahara

H, 10.5 x 4.3 mm; I, 9.3 x 3.6 mm: Specimens H—I from

Spain. off Marbella. 36°28’ N, 04°52’ W, 30-40 m

Specimen L (Figures 10-12), 9.5 x 3.6 mm; M, 8.2 x 3.4

mm: Specimen N, 6.3 x 2.6 mm; Specimens L-N Etymology: This species is named afte: the author's

dredged along the coast of West Sahara: 30-60 m Speci- daughter

Ww OO

Type Locality: Alboran Sea, Spain, off Marbella

36°28' N, 4°52’ W, 30-40 m depth

Habitat: The dredged specimens were from muddy

bottoms

Page 162

THE NAUTILUS, Vol. 121, No. 3

Remarks: Brachycythara beatriceae new species

shows several shell diagnostic features (shape and sculp-

ture of the protoconch and the teleoconch) that match

the ones described by Powell (1966: 117, plate 18, fig. 7)

for the genus Brachycythara, an amphiatlantic group of

Recent to Miocene/Pliocene species whose distribution

ranges from the Caribbean to the coast of West Africa

(Powell, 1966; Rolan and Otero-Schmitt, 1999). Brachy-

cythara atlantidea (Figures 10-12, 21-28) has been the

only species ae to this genus known to occur in

the eastern Atlantic Ocean (Rolén and Otero-Schmitt,

1999) and in the Alboran Sea (Smriglio et al., 2007).

Brachycythara beatriceae and B. atlantidea show close

similarities, but the two taxa can be easily separated on

the basis of several shell morphological differences, as

summarized in Table 1. Furthermore, at high magnifi-

cation the complicated spiral microsculpture of B. bea-

triceae shows finer threads, smaller and more numerous

granules in each interspace. The finding of three speci-

mens of B. beatriceae from two localities off the Spanish

coast (Alboran Sea) indicates that this species can be

added to the Recent Mediterranean malacofauna, while

the collecting depth suggests that it is a circumlittoral

species. As a final conclusion, the genus Brachycythara is

at the present represented by two sympatric species dis-

tributed in the Alboran Sea and the eastern Atlantic

Table 1. Comparisons of shell features between B. beatriceae

and B. atlantidea

B. atlantidea

Morphological characters B. beatriceae

Prontoconch diameter size 700-750 600-650

(in jzm)

Protoconch number of whorls 3-34 3

Teleoconch axial folds 7-8 9-10

(last whorl)

Shell color white with a

basal brown

uniformly

yellowish-

band brown

Ocean, Brachycythara atlantidea and Brachycythara

beatriceae.

ACKNOWLEDGMENTS

I am grateful to Mr. Carlo Smriglio and Dr. Marco Ol-

iverio (Dipartimento di Biologia Animale e dell Uomo,

Universita di Roma “La Sapienza”, Rome, Italy) for their

critical comments and advice on the present paper. I

would like to express my deep gratitude to Mr. Franco

Gubbioli (Marbella, Malaga, $ Spain) and to Dr. Stefano

Rufini (Dipartimento di Biologia, Universita di Roma

“Tor Vergata’, Rome, Italy) for kindly supplying type

material of B. beatriceae and specimens of B. atlantidea.

Dr. Andrea Di Giulio (Dipartimento di Biologia, Univer-

sita di “Roma Tre”, Rome, Italy) is acknowledged for

SEM photographs, which were carried out at the LIME

(Interdepartmental Laboratory of ie Microscopy,

Universita di “Roma Tre”, Rome, Italy). I am very in-

debted to two anonymous referees for Shr suggestions

and corrections to the manuscript.

LITERATURE CITED

Powell, A. W. B. 1966. The molluscan families Speightiidae and

Turridae. Bulletin of the Auckland Institute and Museum

5: 1-157.

Rolan, E. and J. Otero-Schmitt 1999. The F roa Turridae s..

(Mollusca, Neogastropoda) in Angola, 2. Subfamily Man-

geliinae Fischer, 1883. Argonauta 13: 5-26.

Smriglio, C., A. Di Giulio, F. Gubbioli, and P. Mariottini 2007,

Brachycythara atlantidea (Knudsen, 1952) (Gastropoda,

Neogastropoda, Conidae), first report from the Western

Mediterranean Sea. Basteria, 71(1—3): 1-4.

Taylor, J. D., Y. I. Kantor, and A. V. Sysoev. 1993. Foregut

anatomy, feeding mechanisms, relationships and classifi-

cation of the Conoidea (=Toxoglossa) (Gastropoda). Bul-

letin of the Natural History Museum (Zoology) 59: 125—

170.

Woodring, W. P. 1928. Miocene Mollusks from Bowden, Ja-

maica. Part Il, Gastropoda and discussion of results. Car-

negie Institute, Washington, Publication 85, 564 pp.

MBL WHO!

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CONTENTS

VIC t

Volume 121, Number 4

December 21, 2007

ISSN 0028-1544

Thomas J. DeVries

John Slapcinsky

Robert Lasley

Lennie Rotvit

Jorgen Litzen

Ase Jespersen

Thomas Fox

Eliane P. Arruda

Osmar Domaneschi

Jonata de A. Francisco

José Carlos N. de Barros

Donn L. Tippett

Late Cenozoic Tegulinae (Gastropoda: Trochidae) from southern Peru... .

Three new species of Paryphantopsis (Gastropoda: Pulmonata:

Charopidae) from the Nakanai Mountains, New Britain,

Papua: New Guineas... 5-06 eee eh ee dae Gees cee Seeds ase tbe

Mysella gregaria new species, a bivalve (Galeommatoidea: Montacutidae )

to) D

commensal with an intertidal burrowing sea anemone from

North ‘Carolina, USAs «cis c0ch e008 eo gawas teehee ose h ew DEE HM

Corbula tarasconii, a new species of Corbulidae (Bivalvia) from

offshore: Brazil. p< ct a a eee odd Ghat oe wake eee baw ee ded Be hdd

Two new gastropod species (Neogastropoda: Drilliidae, Turridae) from

the western Atlantic Ocean . 0.0

163

182

190

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THE NAUTILUS 121(4):163-1S1, 2007

Page 163

Late Cenozoic Tegulinae (Gastropoda: Trochidae) from

outhern Peru

Thomas J. DeVries!

Burke Museum of Natural History and

Culture

University of Washington

Seattle. WA 98195 US. A

ABSTRACT

Four new fossil tegulines (Gastropoda: Trochidae) are de-

scribed from southern Peru [Chlorostoma quipua new species

(late Miocene to late Pliocene), Intistoma pirqua new genus,

new species (early Pliocene), Cantallocostoma panistostum new

genus, new species (late Miocene to early Pliocene), Tegula

(s.l.) masiasi new species (early to middle Miocene)], as are

Pliocene and Pleistocene occurrences of the extant Chloro-

stoma atrum (Lesson, 1830), C. luctuosum (d’Orbigny, 1841),

Cantallocostoma quadricostatum (Wood, 1828), Agathistoma

patagonicum (dOrbigny, 1835), T. (s.1.) melaleucos (Jonas,

1844), and T. (s.l.) tridentata (Potiez and Michaud, 1838).

These data show that the Peruvian chlorostomine group is too

ancient to be a Pliocene sister group to Caribbean-Atlantic

agathistomines: indicate additional eastern Pacific groups of

tegulines exist with roots reaching into the Miocene; and fur-

ther demonstrate the success of A. patagonicum as a wide-

spread and long-lived teguline in austral waters. A small radia-

tion of Peruvian chlorostomines during the late Pliocene coin-

cided with a molluscan mass extinction event in the Peruvian

Faunal Province.

Additional Keywords: Mollusk, Tegula, Chlorostoma, Agathi-

stoma, Miocene, Pliocene, Cenozoic, Pisco Basin, Taxonomy

INTRODUCTION

Seven extant teguline species (Gastropoda: Trochidae)

inhabit the cool coastal waters of the Peruvian Faunal

Province. They are, according to their traditional nomen-

clature, Tegula atra (Lesson, 1830), T. luctuosa

(dOrbigny, 1841), T. ewryomphala (Jonas, 1844), T. ig-

nota ae 1976, T. tridentata (Potiez

and Michaud, 1838), T. quadricostata (Wood, 1528),

and T. patagonica (dOrbigny, 1535) (Alamo and

Valdivieso. 1997: Véliz and Vasquez, 2000). An eighth

species, Tegula melalewcos (Jonas, 1544), a species from

warmer waters of northern Peru, does appear rarely at

southern Peruvian latitudes. This teguline fauna is as

diverse as that of the warm-water Panamic Faunal Prov-

* Mailing address: P.O. Box 13061, Burton, WA 98013 USA

ince (Keen, 1971; Alamo and Valdivieso, 1997), although

with mostly different species, and is more speciose than

the teguline f auna of the Magellanic Faunal Province

(four species), which is a subset of the Peruvian fauna

(Forcelli, 2000).

The geological record of tegulines from the Peruvian

Faunal Province is meager. Tegula luctuosa, T. atra, and

T. tridentata are listed in Herm’s (1969) study of

Pliocene and Pleistocene mollusks from Chile. Tegula

luctuosa and T. melaleucos were reported from upper

Pliocene and Pleistocene beds of northern Peru

(DeVries, 1986). Four new species of tegulines, consid-

ered to have early to middle Miocene age (DeVries and

Frassinetti, 2003: Finger et al., 2007), have been de-

scribed from the Navidad Formation of central Chile by

Nielsen et al. (2004).

This paper documents four new fossil species and sev-

eral previously known Recent species of tegulines in

Neogene deposits of southem Peru, including the oldest

leewn teguline from Peru, the early Miocene Tegula

(s.l.) masiasi new species; creates two new genera of

tegulines, Cantallocostoma, new genus, and Intistoma,

new genus, each with a newly described Neogene fossil

species in the Peruvian Faunal Province, Cantallocos-

toma panistosum new species and Intistoma pirqua new

species, and each having a modern representative in the

eastern Pacific Ocean, the Peruvian C. quadricostatum

and Californian I. aureotinctum (Forbes, 1852); adopts a

full generic status for Tegula (Chlorostoma); and pro-

vides evidence for a late Miocene origin of a western

South American group of Chlorostoma species.

GEOLOGY

The late Cenozoic marine stratigraphy of southern Pe-

ruvian forearc basins has been described by Dunbar et al

(1990) and DeVries (1998). Teguline-bearing deposits

crop out west of Nazca and throughout the Sacaco Basin

(Fig. 1). These bioclastic conglomerates and sandstones

which were assigned to the La Planchada and Pisco for-

Page 164

THE NAUTILUS, Vol. 121, No. 4

Camana Basin’

1 | 1 l 1 |

Figure 1. Location of onshore portions of Cenozoic forearc

basins in southern Peru. New fossil species of Tegula are from

Cenozoic deposits west and south of Nazca.

mations by Beaudet et al., 1976, and Muizon and

DeVries, 1985, respectively, represent high-energy fore-

shore and intertidal environments lying eleee to moun-

tainous paleo-shorelines and quieter forediore and inner

shelf environments lying hundreds to thousands of

meters from paleo- shorelines defined by the beginning

of the Andean foothills.

MATERIALS AND METHODS

Most Peruvian specimens described in this study were

found by the author. Comparative material was studied

at the Natural History Museuin of Los Angeles County,

Los Angeles, California, USA (LACM). ‘Locality and

sample descriptions are listed in the appendix. Lengths

(L) and widths (W) are measured in millimeters, with

dimensions of broken specimens enclosed by parenthe-

ses. A non-standard orientation for photographs of some

specimens has been necessary to reveal important char-

acters. Some figured specimens were coated with ammo-

nium chloride. ‘Type s and numbered specimens, includ-

ing those figured, are deposited at the Orton Geological

Museum, Ohio State University, Columbus, Ohio USA

(OSU): the Departamento de Paleontologia de Vertebra-

dos, Museo de Historia Natural, Universidad Nacional

Mayor de San Marcos, Lima, Peru (MUSM INY); and

the Burke Museum of Natural History and Culture, Uni-

versity of Washington, Seattle, Washington (UWBM).

The prefixes “DV” refers to DeVries localities, “|M” to

localities of J. Macharé (Instituto Geolé6gico Minero y

Metalirgico, Lima, Peru), and “WIZ to localities of W.

J. Zinsmeister (Purdue University, Indiana, USA).

SYSTEMATICS

Superfamily Trochoidea Rafinesque, 1815

Family ‘T Crochidae Rafine ssque, LS15

Subfamily Tegulinae Kuroda, Habe and Oyama, 1971

Genus Tegula Lesson, 1835

Type Species: Trochus elegans Lesson, 1835 (by

monotypy) (= Trochus pellisserpentis Wood, 1828). Re-

cent, Pacific coast of Central America.

Remarks: Resolving the difficult subfamilial place-

ment of Tegulinae wathib Trochoidea (Hickman and

McLean, 1990: Bouchet and Rocroi, ‘ 2005) is beyond the

scope of this paper. Tegula itself has been assigned to

three different subfamilies over the past century, but

most taxonomists now refer it to the undiagnosed Teg-

ulinae Kuroda, Habe, and Oyama, 1971 (Hiclanen and

McLean, 1990). Only one extant teguline species, Tegula

pellisserpentis (Wood, 1828), pertains to Tegula a 'S5)

(Keen, 1971). Its combination of foavares: (densely

packed and heavily beaded spiral cords, tightly twisted

columella, and numerous teeth extending from the col-

umella across the apertural floor) is not seen in other

species assigned to Tegula, so T. pellisserpentis is herein

considered to be the monotypic representative of Tegula

(s.s.), an opinion shared by J]. H. McLean (pers. comm.,

2006). Taxa often considered as subgenera of Tegula,

e.g., Chlorostoma and Agathistoma, are elevated to ge-

neric status in this paper in accord with the practice of

some authors (e.g., Ammold, 1907; Higo et al., 1999) and

the opinion of J. HH. McLean (pers. comm., 2006).

Genus Chlorostoma Swainson, 1840

Type Species: §Trochus argyrostomus Gmelin, 1791

(by subsequent designation, Herrmannsen, 1546). Re-

cent, Japan.

Remarks: Swainson (1840) erected Chlorostoma as a

subgenus of Trochus Linnaeus, 1755, to include species

with a “remarkably oblique” aperture, a very deep um-

bilicus, oa one or two teeth on the outer (= lower part

of inner ?) lip. He assigned two species s to Chlorostoma:

Ti me (Chlorostoma) argyrostoma (= Tegula argyros-

toma of authors) and Trochus (Chlorostoma) umbilicaris

|= Gibbula umbilicaris (Linnaeus, 1758) |. Herrmannsen

(1546) implicitly limited the definition of Chlorostoma by

choosing T. argyrostoma as the type species. Examina-

tion of LACM material from the western North Pacific

Ocean [C. argyrostoma, C. lischkei (Tapparone-Canefri,

1874), C. rusticum (Gmelin, 1791), C. turbinatum (A.

Adams, 1853), C. xanthostigma (A. Adams, 1853)] shows

that adult chlorostomine umbilici can be either open or

closed. In the view of Grant and Gale (1931), Chloros-

toma should be further restricted to toothed species hav-

ing ventricose whorls, a nacreous interior, and a dark

purplish exterior. This diagnosis makes possible the iden-

tification of Recent chlorostomine taxa from both sides of

the North Pacific Ocean and in the eastern South Pacific

Ocean.

Chlorostoma pia (Lesson, 1830)

(Figures 2-4, 7-9, 13)

Trochus ater Lesson, 1830, vol. 2, pt. 1, p. 344, Mollusques, pl.

16, fig. 2: d Orbigny, 1S40: 409; Philippi, 1544, Abbildun-

T. DeVries, 2007 Page 165

Figures 2-4, 7-9, 13. Chlorostoma atrum (Lesson, 1830). 2. UWBM 97863, DV 1372-1, Recent, oblique spire view, width = 9.8

mm. 3. UWBM 97855, DV 395-1, Recent, basal view, width = 30.7 mm. 4. UWBM 97863, oblique basal view. 7. MUSM INV 126,

DV 1252-1, early Pleistocene, oblique spire view, width = 27.1 mm. 8. UWBM 97855, i aalote view. 9. UWBM 97860, Paracas

Hotel, Recent, lateral view, length = 22.2 mm. 13. UWBM 97860, oblique basal view, width = 23.4 mm. Figures 5,6. © Chlorostoma

funebralis (A. Adams, 1855). UWBM 97562, south of La Jolla, California, Recent, width = 13.4 mm. 5. Oblique spire view. 6. Oblique

basal view. Figures 10-12, 14-19. Chlorostoma luctuosum (d’Orbigny, 1841). 10. UWBM 97864, Paracas Hotel, Recent, lateral

view, length = 20.2 mm. Il. UWBM 97865, Paracas Hotel, Recent, apertural view, length = 20.5 mm. 12. UWBM 97866, Hueco

La Zorra, Peru, Recent, apertural view, length = 21.6 mm. 14. UWBM 97864, oblique basal view, width = 23.7 mm. 15. UWBM

97865, basal view, width = 24.4 mm. Figure 16. UWBM 975866, oblique spire view, width = 28.5 mm. 17. UWBM 97567, Lomas.

Peru, Recent, oblique spire view, width = 26.4 mm. 18. UWBM 97867, oblique basal view. 19. UWBM 97868, Chile, Pleistocene,

oblique basal view. width = 37.3 mm. Figures 20, 21. Chlorostoma ao aay (Jonas, 1844). UWBM 97871, DV 1599-1,

Recent, length = 26.0 mm. 20. Apertural view. 21. Lateral view. Figures 22, 23. Chlorostoma ignotum (Ramirez-Bohme, 1976).

UWBM 97872, Pellehue, Chile, Recent, width = 17.5 mm. 22. Oblique spire view. 23. Basal view. oo 24-26. Chlorostoma

quipua new species. 24. UWBM 97873, DV 1254-Bal 6, late early Pliocene, basal view, width = 17.3 mm. Note faint protractive

stripes on base. 25. MUSM INV 136, DV 571-1, syntype, late Miocene, oblique spire view, ak “170 mm. 26. UWBM 97873

oblique spire view.

Page 166

gen und beschreibungen neuer oder wenig gekannter

Conchylien, v. 1, p. 188, pl. 5, fig. 6; Philippi, 1846, Die

Kreiselschnecken oder Trochoideen, p. 195, pl. 30, fig. 1;

Hupé, 1854, p. 142, Malacologia, pl. 4, fig. 2.

Monodonta atra Lesson.—Potiez and Michaud, 1838: 319, pl.

29, figs. 14, 15.

Tegula atra Lesson.—Dall, 1909: 239, pl. 24, fig. 4; Carcelles and

Williamson, 1951: 262; Aldea and Valdovinos, 2005: fig. SB.

Morch, 1850: 20,

Tegula (Chlorostoma) atra (Lesson, 1830).—Marincovich,

1973: 24, fig. 42; Alamo and Valdivieso, 1997: 13, fig. 25;

Guzman et al., 1998: 35, fig. 22; Forcelli, 2000: 61, fig. 87;

Véliz and Vasquez, 2000: 759, fig. 1B.

Trochus moestus Jonas, 1844: 113; Philippi, 1846, Abbildungen

und peschreibungen neuer oder wenig gekannter Con-

chylien, v. 2, pl. 6, fig. 5; Philippi, 1 1846, Die Kre-

isclsthnedlen’ oder Trochodecn. p. 199, Pr 30, fig. 2;

Hupé, 1854: 147, Malacologia, pl. 4, figs. 3, 3a, 3b.

Tegula moesta (Hupé, 1854).—D. all, 1909: 239: Alamo and

Valdivieso, 1997: 14.

Chlorostoma minor Morch, 1850: 20.

Diagnosis: Shell width about 40 mm; last whorl

broadly rounded, including shoulder; keels lacking. Um-

bilicus of juvenile and adult shells white, closed; meni

cus with two well exposed spiral cords, the adaxial cord

terminating in a columellar tooth.

Material Examined: UWBM 97855, DV 398-1, Re-

cent, L = 25.0, W = 30.7; UWBM 97856, DV 398-1, lot

of 2; UWBM 97857, DV 1252-1, L = 11.8, W = 18.6;

UWBM 97858, DV 1418-1, latest Pliocene, L = (22.2),

W = 30.2; UWBM 97859, DV 463-1, late Pleistocene, lot

of 2; UWBM 97860, Paracas Hotel, Recent, L = 22.2; W

= 23.4; UWBM 97861, Ipun, Chile, Recent, lot of 2;

UW BM 97863, DV 1372-1, Recent, L = 8.0, W = 9.8;

MUSM INV 126, DV 1252-1, early Pleistocene, L =

(19.2), W = 27.1; MUSM INV 127, DV 1418-1, L = 31.4,

W = 36.6; MUSM INV 128, DV 463-1, lot of 2.

Occurrence: Late Pliocene to middle Pleistocene:

southern Peru to southern Chile. Late Pleistocene:

southern Peru, Chile, southern Argentina. Recent:

northern Peru to southern Chile, southerm Argentina (G.

Pastorino, pers. comim., 2002).

Remarks: Specimens of Chlorostoma atrum can ex-

ceed 40 mm in width and are generally smooth-shelled

and purple-black, either entirely or dorsally, only. The

last whorl is always broadly rounded; it lacks the keeled

spiral cords present on specimens of C. luctuosum. The

base on some specimens of C. atrwm has weak spiral

threads; the spire of some also has one or more narrow

spiral grooves that produce as many as 15 intervening low

broad spiral cords (Figure 7), not unlike the spiral sculp-

ture of the Californian C. funebralis (A. Adams, 1855)

(Figures 5, 6),

The adult shell of Chlorostoma atrum is usually dis-

tinguished from that of other Peruvian chlorostomines by

its closed umbilicus. [G. Collado (pers. comm., 2005)

notes that juveniles of C. luctwosum and adults of the

small Tegula (s.1.) tridentata occasionally have closed

umbilici.| The white umbilical area on specimens of C.

atrum has two spiral cords. A white inner cord rises from

THE NAUTILUS, Vol. 121, No. 4

beneath the umbilical callus and terminates on the edge

of the columella as a thickened tooth. A weakly devel-

oped white outer cord traces the boundary of the um-

bilical area and becomes flattened on the columella, not

quite protruding far enough to produce a tooth. Rarely,

one or two weak spiral spurs develop between the two

umbilical cords in a nacreous area that lies adaperturally

of a thin, glossy, umbilical veneer. They, too, do not

extend far enough to produce columellar teeth.

Chlorostoma luctuosum (VOrbigny, 1841)

(Figures 10-12, 14-19)

Trochus luctuosus d’Orbigny, 1841, v. 5, p. 409, pl. 76, figs.

16-19; Philippi, 1846, Die Kreiselschnecken oder Tro-

choideen, p. 153, pl. 25, figs. 4, 5; Hupé, 1854: 143.

Tegula luctuosa Orbigny.—D Dall, 1909: 239: DeVries, 1986:

512, pl. 27, figs. 3, 4; Guzman et al., 1995: 36, fig. 23; Véliz

and Vasquez, 2000: 762, fig. 1D: Alamo and Valdivieso,

2000: 14; Aldea and Valdovines, 2005: fig. SE.

Diagnosis: Shell width to 35 mm. Last whorl with one

to three spiral cords or keels. Adult umbilicus open; ju-

venile umbilicus usually open; umbilicus with two well

exposed umbilical spiral cords, the adaxial cord terminat-

ing in a columellar tooth.

Material Examined: OSU 37596, DV 240-23, latest

Pliocene, L = 19.5, W = 23.9: UWBM 97864, Paracas

Hotel, Recent, L = 20.3, W = 23.7; UWBM 97865, Pa-

racas Hotel, Recent, L = 20.5, W = 24.4: UWBM 97866,

Hueco La Zorra, Recent, L = 21.6, W = 28.5; UWBM

97867, Lomas dump, Recent, L = (20.6), W = 26.4:

UWBM 97868, W]Z 345, Chile, Pleistocene, L = 26.8,

W = 37.3; UWBM 97869, DV 382-1, Pleistocene, L =

(20), WwW. 7 98.4: UWBM 97870, WIZ 345, L = (35.0), W

= 43.1; MUSM INV 129, DV 382-1, L = 20.6, W = 31.1;

MUSM INV 130, JM 8220, Pleistocene, lot of 2.

Occurrence: Late Pliocene to upper Pleistocene:

northern Peru to Tongoy, central Chile. Recent: Galapa-

gos Islands to Concepcion, central Chile (southern limit

from LACM collections).

Remarks: Specimens of Chlorostoma luctuosum are

large and purple-black, either entirely or dorsally, only.

Adult specimens are generally distinguished from speci-

mens of C. atrum by having an open cambaeus and from

both C. atrum and C. euryomphalum by having one to

three primary spiral cords or keels: one near a base of

the whorl, forming the periphery (Figure 12); another

about one quarter of the distance anteriorly from suture

to suture (Figure 12); and a third occasional] y dev eloped

just anterior to the periphery (Figure 19). Some speci-

mens of C. luctuoswm are also cov ened with tertiary spiral

threads (Figures 17, 1S). The thin umbilical veneer and

columellar teeth are identical to those on specimens of

C. atrum, as are the umbilical cords, except that they are

exposed coiling deep into the umbilicus.

Chlorostoma euryomphalum (Jonas, 1844)

(Figures 20, 21)

Trochus euryomphalus Jonas, 1844; 113; Philippi, 1844, Abbil-

dungen und beschreibungen neuer oder wenig gekannter

T. DeVries, 2007

Page 167

774250 W

sxOIH NVOIANVNVd

Aguada

de Lomas

Pacific

Ocean

e@ DV _ locality—sample

0 Skm

SCALE

Contour interval is 100 m

Figure 41.

new species.

Type locality (DV 571-1) of Chlorostoma quipua

Conchylien, v. 2, p. 27, pl. 6, fig. 4; Philippi, 1846, Die

Kreiselschnecken . Wi schotdeen, p 15h aL 25, fig. 7.

Tegula euryomphala (Jonas, 1844).—Carcelles and W illiamson,

1951: 262.

Tegula euryomphalus [sic] (Jonas).—Dall, 1909: 239; Alamo

and Valdivieso, 1997: 14.

Tegula euryomphala (Jones, 1844) [sic] —Guzman et al., 1998:

36, fig. 25; Véliz and Vasquez, 2000: 762, fig. LE; Aldea

and Valdovinos, 2005: fig. SC.

Trochus kieneri Hupé, 1854, p. 144, Malacologia, pl. 4, figs. 1,

la. 1b.

Diagnosis: Shell width to 35 mm. Last whorl broadly

rounded. Umbilicus white, open; umbilicus with two we il]

exposed spiral cords, the adaxial cord terminating in a

columellar tooth.

Material Examined: U\WBM 97871, DV 1599-1, Re-

cent, L = 26.0, W = 29.9.

Occurrence: Late Pleistocene: Northern to central

Chile. Recent: Southern Peru to central Chile.

Remarks: Specimens of Chlorostoma euryomphalum

are large, purple-black, and characterized by a broad

open umbilicus and broadly rounded whorls. They differ

from specimens of C. atrum. which have a closed umbi-

licus, and C. luctuosum, which have one or more angular

spiral cords or keels. On some specimens of C. luctio-

sum, however, including Recent Peruvian and Chilean

examples from LACM collections, Pleistocene Chilean

specimens from WJZ collections, and upper Pliocene Pe-

ruvian specimens from northern Peru, the spiral cords

are so weak that assigning the material to C. luctuosum or

C. euryomphalum is problematic.

Chlorostoma ignotum (Ramirez-B6hme, 1976)

(Figures 22, 23)

Tegula ignota Ramirez-Bohme, 1976: 3, figs. 1-6; Forcelli,

2000: 61, fig. 8S; Véliz and Vasquez, 2000: 762, fig. 1F;

Aldea and V idoanos 2005: fig. SD; Collado ond Brown,

2005: 131.

Diagnosis: Shell width to 30 mm. Outer layer slate

colored. Sculpture consists of several well developed, un-

beaded, primary spiral cords. Umbilicus open.

UWBM 97872, La Rinconada,

11.1, W = 17.5; UWBM

29.4, W = 32.6.

Material Examined:

Pelluhue, Chile, Recent, L =

97906, Ipun, Chile, Recent, L =

Occurrence: Recent: central to southern Chile.

Remarks: = Chlorostoma ignotum is known only as a

Recent species from Chile (e.g., Collado and Brown,

2005). Specimens of C. ignotum differ from those of

other extant chlorostomines of the Peruvian Faunal

Province in two significant respects: they lack the purple-

black outer shell jan er of C. atrum, C. luctuosum, and C.

euryomphalum, being rather slate colored, and they have

numerous, prominent, well-differentiated primary and

secondary spiral cords between the periphery and suture

and to a lesser extent on the base. The well-exposed

umbilical cords, thin umbilical veneer, and columellar

teeth are identical, however, with those of other species

of South aac Chlorostoma, and the strong spiral

cords (Figure 22) are like those seen on rare specimens

of early Plioc ‘ene C, quipua new species (Figures 25, 26;

see below)

Chlorostoma quipua new species

(Figures 24—40)

Diagnosis: Adult whorls purple-black dorsally; spire

and base usually light brown, commonly with dark brown

protractive stripe s or mottling. Umbilicus narrow, open.

Description: Shell up to 30 mm wide. Spire angle

about S0 degrees. Periphery near base, sharply rounded

to slightly angul uw. Sutures appressed. Protoconch un-

known: te Jeoconch with five flat-sided to slightly convex

whorls. Axial sculpture absent or rarely with rugose pro-

tractive ribs. Thin growth lines strongly prosocline. Spiral

sculpture of 20 muted spiral thre ads: poste rior to periph-

ery; rarely with three to five spiral grooves separating

four to six broad low spiral cords or mthioul spiral sculp-

ture. Twenty to 30 evenly spaced spiral threads on base

of juvenile specimens, muted or obsolete on adult

whorls. Outer shell layer purple-black on adult whorls,

tan or light brown on spire whorls and base. Protractive

and rarely retrotractive wrinkled brown stripes usually

Page 168 THE NAUTILUS, Vol. 121, No. 4

Figures 27-40. Chilorostoma quipua new species. 27. MUSM INV 131, DV 571-1, syntype, spire view, width = 25.4 mm. 28.

MUSM INV 133, DV 1635-2, early Pliocene, basal view, width = 16.0 mm. 29. UWBM 97880, DV 1635-2, basal view, width = 16.6

mim. 30. UWBM 97879, DV 1635-2, basal view, width = 18.5 mm. 31. UWBM 97590, DV 1598-1, early Pliocene, oblique spire, width

= 11.2 mm. 32. UWBM 97883, DV 1254-1, Pliocene, oblique basal view, width = §.9 mm. 33. UWBM 97876, DV 571-1, syntype,

apertural view, length = 22.7 mm. 34. UW BM 97880, oblique spire view. 35. MUSM INV 133, apertural view. 36. UWBM 97581,

DV 1029-1, early Pliocene, spire view, width = 17.4 mm. 37. UWBM 97879, oblique spire view. 38. MUSM INV 135, DV 1029-1,

oblique spire view, width = 11.3 mm. 39. UWBM 97878, DV 809-1, Pliocene, spire view, width = 5

809-1, basal view, width = 19.0 mm.

2.1 mm. 40. MUSM INV 132,DV

present on base and less often on spire. Inside edge of Type Material: (All DV 571-1, all syntypes) UWBM

outer lip smooth. Umbilicus open, narrow; umbilicé al ve- 97876, L = 22.7, W = 27.3; UWBM 97877, L = (17),

neer thin. Columella with thick inner tooth at end of W = (25.1); MUSM INV 131, L = (18.9), W = 28.4;

well-exposed white umbilical cord. Weak outer MUSM INV 136, L = (10.2), W = 17.0.

tooth adjacent to floor of aperture at end of thin spiral Other Material Examined: UWBM 97873. DV1254-

cord following outer margin of umbilical area. Small pa- Bal 6, late early Pliocene: 1, = (11.0): W = 17.3: UWBM

rietal flange barely ove thanging umbilicus. 97874. DV 1254-Bal 10, late Pliocene, L = (4.8), W =

Type Locality: DV 571-1, Alto Grande, about one km 10.4: UWBM 97875, DV 1254-Bal 10, L = (14.1), W =

south of intersection with abandoned paved road to San 24.4: UWBM 97878, DV SO9-1, Pliocene, L = 15.4, W =

Juan de Marcona, on south-facing hillside west of Pan- 22.) pe 97879, DV 1635-2, early Pliocene, L

american Highway; one of several shell banks of the 12.2. W = 18.5: UWBM 97880, DV 1635-2, L = (11.3),

Pisco Formation (Figure 41). Locality inaccurately re- W = 16.6; UWBM 97881, DV 1029-1, early Pliocene,

ferred to as El Jahui w in Muizon and DeVries (1985). L = (12.3), W = (17.4); UWBM ae DV 1029-1. L =

15°26'57"S, 74°52'06"W (Acari 1:100,000 quadrangle), (12), W = 16.7; UWBM 97883, DV 1284-1, Pliocene,

Middle upper Miocene, L = (5.2), W = 8.9; UWBM 97890, DV 1598-1, early

T. DeVries, 2007

Page 169

Pliocene, L = 8.4, W = 11.2; MUSM INV 132, DV SO9-1,

= gr W = (19.0); MUSM INV 133, DV 1635-2, L =

11.2, W = 16.0; MUSM INV 134, 1635-2 lot of 2;

MUS INV 135, DV 1029-1, L = 6.9, W = 11.3

Occurrence: Middle late Miocene to early late

Pliocene: southern Peru.

Etymology: “Quipua,” Latinized version of “quipu,”

Inca counting device of braided and knotted strings,

evoked by the wrinkled brown stripes on the base and

spire of this species.

Remarks: Specimens of Chlorostoma quipua difter

from those of C. euryomphalum and C. luctuosum by

having a narrower cabied area, smaller parietal f flange,

and protractiv' e brown stripe Ss. Specimens of C. quipua

lack the keeled spiral cords of C. luctuosim and closed

umbilicus of C. atrum. Some specimens of C. quipua,

both Miocene and Pliocene, have broad spiral cords (Fig-

ure 25) like those seen on the juvenile whorls of some

specimens of C. atrum (Figure 7). A single specimen

from upper Pliocene beds above Playa Huacllaco ( Figure

26) has spiral cords as pronounced as the raised spiral

cords on specimens of the modern Chilean C. ignotuim

(Figure 22). Some lower Pliocene specimens near Yauca

(Figure 31) have coarse protractive axial ribs like those

on some Asian chlorostomines.

Specimens of Chlorostoma quipua superficially re-

semble those of C. gallina (Forbes, 1852), a Pliocene-to-

Recent species from California and Baja California

(Grant and Gale, 1931; McLean, 1978), and C. rugosiwm

(A. Adams, 1853), a Recent species from the Gulf of

California (Keen, 1971). Specimens of all three species

have some degree of purple-black color and protractive

stripes on the spire and/or base. Specimens of C. gallina

and C. rugosum, however, are more ventricose laterally

and bz sally and have weak to prominent protractive axial

ribs and stripes posterior to the base. Specimens of C,

gallina usually have a closed umbilicus.

Shells of Chlorostoma quipua are found in upper Mi-

ocene beach deposits near Alto Grande (DV 571-1; see

Muizon and DeVries. 1985) with specimens of Chorus

frassinetti DeVries, 1997, and Acanthina obesa DeVries,

2003 (DeVries, 1997, 2003). Lower Pliocene specimens

of C. quipua occur together with specimens of the mu-

ricid gastropods, Concholepas kieneri Hupé, 1854; Xan-

thochorus ochuroma DeVries, 2005; and Herminespina

saskiae DeVries and Vermeij, 1997: and the turbinid gas-

tropod, Prisogaster mcleani DeVries, 2006 (DeVries,

2005, 2006; DeVries and Vermeij, 1997)

Genus Cantallocostoma new genus

Type species: Trochus quadricostatus \Vood, 1828.

Recent, Peru and Chile.

Diagnosis: White to brown outer shell layer. Three to

five beaded primary spiral cords. U mbilicus open, broad,

Two adaxially situated spiral umbilical cords terminating

in columellar teeth. Parietal wall vertical, without pari-

etal flange overhanging umbilicus.

Description: Shell wp to 35 mm in diameter. Whorls

ventricose to quadrate; periphery weakly bicarinate. Spi-

ral sculpture of three to five primary spiral cords bro-

ken into beads; interspaces with two to five continuous

or weakly beaded tertiary threads. Base with four to

five beaded or non-beaded primary spiral cords and in-

tervening secondary cords and tertiary threads. Um-

bilical area white, tabulate, sh: ply de fined, with margin

of umbilical area flaring towards aperture as steep-

ly inclined wall. U salslious open, broad, with two spiral

umbilical cords situated adaxiz ily, the innermost

thicker; each cord terminating in columellar tooth.

Third tooth sometimes present at base of columella. Um-

bilical veneer variably developed. Columella thin,

upright, without parietal flange overhanging umbili-

cus. Floor of aperture with le -dge but without teeth. In-

ner lip sometimes with four to six closely spaced low

teeth.

Occurrence: Late Miocene to Pleistocene: southern

Peru. Recent: northern Peru to Chile.

Etymology: “Cantalloc,” site near Nazca, Peru, where

subterranean aqueducts are reached from ground level

by pre-Incaic stonewall-lined spiral paths that resemble

the spiral umbilical cords of this genus.

Remarks: Specimens of Cantallocostoma differ from

those of Asian, Californian, and Peruvian Chlorostoma by

having beaded spiral cords and two adaxially situated

amabileeal spiral cords. They differ from specimens of

Intistoma new genus, by having closely spaced beads and

lacking a subsutural band of well-deve loped protractive

nodes, Specimens of Cantallocostoma fer from teg-

ulines traditionally assigned to Agathistoma by lacking

hallmarks of that genus: “narrow open umbilicus, a

smooth or finely beads d spiral sculpture, and a varie-

gated surface coloration” (Olsson and Harbison, 1953:

351).

Cantallocostoma quadricostatum (Wood, 1825)

(Figures 42-45, 47-49)

Trochus quadricostata Wood, 1828: 16, pl. 5, fig. 16.

Trochus quadricostatus Wood.—Philippi, 1546, Die krei-

selschnecken oder Trochoideen p. 154, pl. 25, fig. 6.

Tegula quadricostata (Wood, 1828).—Véliz and Vasquez, 2000

759, fig. LA; Aldea and Valdovinos, 2005: SF.

Tegula quadricostata Gray.—Dall, 1909: 240; Alamo and

Valdivieso, 1997: 14.

Monodonta catenifera Potiez and Michaud, 1838: 315, pl

figs. 12-13.

Trochus torulosus Philippi, 1543,

bungen neuer oder wenig gekannter Conchylien, v. 1, pl

2. fig. 12.

Abbildungen und Beschrei-

Diagnosis: Low spire, white to cream-colored with

purple along edges of sculptural elements. Sculpture of

three to five prominent primary beaded vee cords.

Material Examined: UWBM 97S8S4, DV 13 e

cent, | (14.4), W = 23.9: UWBM 978585, D\

y= : ;

L = (13.3), W = (20.8); UWBM 97886, DV 1713-1, early

2-1, R

372-]

Page 170 THE NAUTILUS, Vol. 121, No. 4

Figures 42-45, 47-49. Cantallocostoma quadricostatum Wood, 1828. 42. UWBM 97884, DV 1372-1, Recent, oblique spire view,

width = 23.9 mm. 43. UWBM 97886, DV 1713-1, early Pleistocene, apertual view, width = 21.5 mm. 44. MUSM INV 137, DV

1355-1, late Pliocene, basal view, width = 16.0 mm. 45. UWBM 97854, oblique basal view. 47. UWBM 97885, DV 1372-1, oblique

lateral view, width = 20.8 mm. 48. UWBM 97887, DV 1355-1, oblique spire view, width = 18.1 mm. 49. UWBM 97886, oblique basal

view. Figures 46, 50-54. Cantallocostoma panistostum new species. 46. MUSM INV 141, DV 1635-2, early Pliocene, oblique

spire view, width = 14.3 mm. 50. UWBM 97891, DV 571-1, late Miocene, oblique spire view, width = 14.9 mm. 51. UWBM 97889,

DV 1598-1, syntype, oblique lateral view, width = 33.7 mm. 52. UWBM 97889, oblique basal view. 53. UWBM 97889, apertural

view. 54. MUSM INV 139, DV 1598-1, syntype, basal view, width = 24.9 mm. Figures 55-57. Intistoma pirqua new species.

UWBM 97892, DV 470-1, syntype, early Pliocene, length = 38.9 mm. 55. Lateral view. 56. Apertural view (lighting from upper right).

57. Oblique basal view. Figures 58,59. — Intistoma aureotinctum (Forbes, 1852). South of La Jolla, California, Recent. 58. UWBM

97596, lateral view, length = 20.0 mm. 59. UWBM 97897, oblique basal view, width = 17.8 mm. Figures 60-63. Agathistoma

patagonicum (VOrbigny, 1535). 60. UWBM 97893, Argentina, Recent, oblique spire view, width = 15.4 mm. 61. UWBM 97893,

oblique basal view. 62. UWBM 97895, DV 1032-2, late Pliocene, lateral view, width = 12.7 mm. 63. UWBM 97895, oblique spire

view,

T. DeVries, 2007

Pleistocene, L = (14.5), W = 21.5: haley 97SS7, DV

1355-1, late Pliocene, L = — V = 18.1; UWBM

OE? DV 1355-1], L, = (10.0), ae 1; MUSM INV

, DV 1355-1, L = (8.9), W = ie MUSM INV 138,

DV 1355-1, L = 14.1, W = (19).

Occurrence: Late Pliocene: southern Peru. Recent:

northern Peru to Chile.

Remarks: The number of primary beaded spiral cords

on the last whorl of Cantallocostoma quadricostatum var-

ies between three and five. Most modem specimens have

two widely spaced primary spiral cords on the anterior

half of the whorl and two closely spaced primary spiral

cords adjacent to the posterior suture (Figure 42). Some

specimens have an additional primary spiral cord be-

tween the two anterior spiral cords (Figure 47); other

specimens have one of the two posteriormost primary

spiral cords missing (Figure 43). The six known late

Pliocene specimens from southern Peru (Figure 45) and

single early Pleistocene specimen (Figure 43) have three

primary spiral cords.

Cantallocostoma panistostum new species

(Figures 46, 50-54)

Diagnosis: Exterior uniformly pale brown; spiral

sculpture of three beaded spiral cords. Basal spiral cords

bunched towards umbilical area.

i Shell nearly 35 mm in diameter. Spire

angle about 70 degrees. Periphery at base, sharply

rounded. Sutures impressed. Protoconch unknown. Te-

leoconch with five quadrate whorls. Axial sculpture ab-

sent except for intermittently rugose prosocline growth

lines. Spiral sculpture of * closely beaded spiral Sond at

base and two posterior spiral rows of more widely spaced

beads, about 25 per whorl, bracketing shoulder. Poste-

rior beads sometimes slightly protractively elongate. In-

terspaces rarely with beaded secondary i cords: usu-

ally with wavering tertiary spiral threads. Base weakly

convex, with six to ten closely spaced spiral cords adja-

cent to umbilical area. Outer shell layer pale brown,

without color pattern. Inner edge of outer lip smooth.

Umbilicus open, wide, with two adaxially situated spiral

cords. Umbilical veneer thin. Innermost umbilical cord

prominent, second cord thin; each terminates in col-

umellar tooth. Third tooth at base of columella nearly

obsolete. Columella thin; parietal flange absent.

Type Locality: DV 1595-1. eres along Panameri-

can Highway, shell beds exposed along descent from

north into Rio Yauca valley (Figure 64). 15°39'49"S,

74°31'50"W (Yauca 1:100,000 quadrangle). Lower

Pliocene.

Type Material: (DV 1595-1, all syntypes) UWBM

97889, DV 1598-1, L = (25.2), W = 33.7: UWBM 97890.

(e) Town or farm

0) 5km

SCALE

Contours in meters

Figure 64. Type locality (DV 1598-1) of Cantallocostoma

panistostum new species and other teguline localities between

Sacaco and Yauca.

DV 1598-1, L = 8.4, W = 11.2; MUSM INV 139, DV

1598-1, L = (13.7), W = 24.9: MUSM INV 140, Pan-

american roadcut at Yauca, early Pliocene, L = (17.9),

V = 33.3.

Other Material Examined: UWBM 97891, DV 571-

1, late Miocene, L = 10.4, W = 14.9; MUSM INV 141,

DV 1635-2, early Pliocene, L = 8.8, W = 14.3.

Occurrence: Pliocene: south-

ern Peru.

Late Miocene to early

Etymology: “Panis,” Latin noun meaning “bread,” and

“tostum,” Latin neuter past participle- adjective me aning

“toasted,” referring to the bread-crust color of this spe-

cles.

Remarks: The light brown color of the outer shell

layer on specimens ‘of Cantallocostoma panistostum re-

sembles that of specimens of Tegula hemphilli Oldroyd,

1921, a late Pliocene-to-Pleistocene species from Cali-

fornia (Grant and Gale, 1931). Specimens of T. hemprhilli

and numerous other Miocene and Pliocene Californian

teguline species with similar coloration lack the two um-

bilical spiral cords close to the axis and are covered by

numerous closely spaced primary spiral cords, none of

which are beaded.

Specimens of Cantallocostoma panistostum are found

in upper Miocene nearshore sandstones with Chloros-

toma quipua; Chorus frassinetti DeVries, 1997; Acan-

thina obesa; and Xanthochorus stephanicus DeVries,

2005; and in lower Pliocene cobbly bioclastic gravels as-

sociated with the mouth of the paleo-Rio Yauca with

disarticulated valves of an undescribed Anadara species,

venerid bivalves, Chlorostoma quipua, Xanthochorus

ochuroma, and Concholepas nodosa Moricke, 1896.

Genus Intistoma new genus

Type species: Trochus aureotinctus Forbes, 1852

Pleistocene to Recent, California.

j0e wd)

Page 2,

THE NAUTILUS, Vol. 121, No. 4

Diagnosis: Spiral sculpture of subsutural band of thick

protractive nodes and peripheral and sub-peripheral pri-

mary spiral cords, Base with three thick, primary spiral

cords. Umbilicus open.

Description: Shell up to 45 mm wide, spire angle

about 75 degrees. Whorls four to five in number, quad-

rate to carinate; sutures weakly impressed. Protoconch

unknown. Sculpture of thick rounded protractive axial

ribs intersecting with an equally thick spiral cords, pro-

ducing a broad subsutural spiral band of elongate pro-

tractive nodes, a near-basal peripheral band of stubby

protractive nodes more numerous than nodes in the sub-

sutural band, and a sub-peripheral primary spiral cord

with little axial modification. Tertiary threads sometimes

present; often corrugated by slightly raised strongly ob-

lique lamellar growth lines. Umbilicus open. Columella

thin; parietal flange erect to slightly overhanging umbi-

licus: parietal callus small. Umbilical veneer thick, cov-

ering all but wedge-shaped adapertural portion of um-

bilical wall. Umbilical spiral cord submerged in umbilical

wall, emergent terminally as prominent columellar tooth.

Smaller second columellar tooth sometimes present

abaxially adjacent to first tooth.

Etymology: “Inti,” the Inca sun god, with a nod to

California's sunshine and the sunset-orange color inside

the umbilicus of the type species, Intistoma aureotinc-

tum.

Occurrence: Late Miocene or early Pliocene: south-

ern Peru. Early Pleistocene to Recent: California.

Remarks: The new genus, Intistoma, is proposed for

two very similar species: the Pleistocene-to-Recent Cali-

ornian Intistoma aureotinctum and the early Pliocene

Peruvian I. pirqua new species. Specimens of both spe-

cies differ from those of nearly all other teguline taxa by

possessing three thick primary spiral cords on the b: ise,

rather than cords that are more numerous and thinner.

Specimens of Intistoma additionally differ from those

oroperly assigned to Agathistoma by ‘lac ‘king teeth on the

loor of the aperture aud the inner edge of the outer lip

and by lacking closely spaced be aded primary spiral

cords.

Intistoma aureotinctum has been considered the ex-

tant representative of a lineage of Californian Neogene

tegulines (Addicott, 1970) that includes the early Mi-

ocene Tegula dalli arnoldi Addicott, 1970, the late Mi-

ocene Tegula nashae Clark, 1915, and Pliocene Tegula

hemphilli Oldroyd, 1921. Specimens of fossil ‘aliforniz m

spec ies do have a subsutural band of e longate protrac tive

nodes, as do specimens of Intistoma, and some have

thick peripheral spiral cords, but none have the distinc-

tive intistomine combination of thick basal spiral cords

and tertiary spiral threads across the entire surface of the

W he rls

Intistoma pirqua new species

(Figures 55-57)

Diagnosis: Shell large, weakly bicarinate, with poste-

rior row of protractive nodes. Base of shell with three

broad spiral cords. Umbilicus open.

Description: Shell large, width more than 40 mm;

spire angle about 60 degre ees. Whorls ventricose, weakly

bicaraate: sutures deeply impressed. Protoconch un-

known; early spire whorls missing; teleoconch of at least

three whorls. Last whorl with sculpture of about 18 elon-

gate protractive ribs on posterior half of whorl; with

about 21 circular to slightly protractively elongate nodes

at periphery, two- ‘hinds the distance from suture to base:

and with a continuous low broad spiral cord at edge of

base. Base with three low broad spiral cords, innermost

cord bordering umbilical area and twice as wide as other

two cords. Faint traces of secondary spiral cords in in-

terspaces laterally and basally. Aperture oblique, outer

lip and colabral growth lines’ moderately prosocline (40

degrees). Wrbilieus open. Columella with at least one

tooth, directed basally; anterior portion par tly excavated,

partly missing. Floor of aperture partly missing; no teeth

evident.

Type Locality: DV 470-1, above rocky road from Ha-

cienda Tunca to Quebrada Huaricangana; hillside of

brown sandstone (Figure 65). 14°56" S, 75°09! W (Palpa

1:100,000 quadrangle). Upper Miocene or lower

Pliocene.

Type Material: (DV 470-1, syntypes, late Miocene or

@ DV locality-samples

0 5 10 km

—

SCALE

Contour interval is 500 m.

PUERTO’ <

9 CABALLAS

. £ " va

MearicahO Ze

\ on ‘Cerro Huar icangana —15°00'S

Figures 65.

new species.

Type locality (DV 470-1) of Intistoma pirqua

T. DeVries, 2007

Page |

early Pliocene) nae Aig L = (388.9), W

MUSM INV 142, L 2), W = (42).

Occurrence:

ern Peru.

Late Miocene to early Pliocene: south-

Etymology: “Pirqua,” Latinized version of “pirqa,”

Quechua word for “wall,” referring to the similarity of

this species’s sculpture and Incaic stone walls.

Remarks: The type specimens of Intistoma pirqua

closely resembles specimens of I. aureotinctum (Figures

58.59), diff ffering principally by being twice the size in all

dimensions and by having better dev veloped nodes on the

peripheral spiral cord. The specimens of I. pirqua were

found together with specimens of Chlamys simpsoni

(Philippi, 1887) and Panopea coquimbensis (dOrbigny,

1842), both species from lower Pliocene beds in Chile

(Herm, 1969) and southern Peru (Muizon and DeVries,

1985).

Genus Agathistoma Olsson and Harbison, 1953

Type Species: Trochus viridulus Gmelin, 1791 (by

original designation). Recent, Caribbean and northeast-

ern South America.

Remarks: One of two Recent specimens of Agathis-

toma patagonicum from Argentina (UWBM 97893) lacks

an open umbilicus (Figure 61) , as do some specimens of

the Pliocene Sierra Laziar outcrops in Argentina (Iher-

ing, 1907), suggesting either that the character is not

diagnostic for all species of Agathistoma (Olsson and

Harbison, 1953) or that the species in question might not

be a member of the Agathistoma group.

Agathistoma patagonicum (d’Orbigny, 1835)

(Figures 60-63)

Trochus (Monodonta) patagonicus “i Orbigny, 1535, vol. 3(4), p.

155; d’Orbigny, 1840, vol. 5(3), p. 408, pl. 55, figs. 1-4.

Neomphalius patagonicus ( (Orb.). ). —thering, 1907: 400.

Tegula patagonica Orbigny.—Dall, 1909: 240; Alamo and

Valdivieso, 1997: 14; Forcelli, 2000: 62, fig. 89.

Tegula (Agathistoma) patagonica (dOrbigny, 1835).—Rios,

1985: 20, pl. 9, fig. 77; Del Rio, 1998: 27, pl. 1, figs. 16-17.

Trochus corrugatus Philippi, 1844, Abbildungen und Beschire 2

bungen neuer oder wenig gekannter C onchylie n,v. 1, p.

67, pl. 2, fig. 7.

Trochus fuscesens Philippi, 1844, Abbildungen und Beschrei-

bungen neuer oder wenig gekannter Conchylien, v. 1, p.

92, pl. 3. fig. 8

Trochus ae gnyana Pilsbry, 1900: 110; Carcelles, 1945: 38, pl.

1, figs. 6, 7, 12-15.

Material Examined: UWBM 97893, Cabo dos

Bahias. Chubut Province, Argentina, Recent, L = 11.2,

W = 15.4: UWBM 97894. Cabo dos Bahfas, Chubut

Province. Argentina, Recent, L = 11.6, W = 14.6;

UWBM 97895, DV 1032-2. late Pliocene, L = (9), W =

12.7.

Diagnosis:

closely spaced beaded primary spiral cords with interca-

Shell under 20 mm wide. Sculpture of

lated secondary cords; beading sometimes obsolete. Col-

umella with three teeth: ee ae open or closed.

Occurrence: Late early to middle Miocene: Argentina

(Ihering, 1907). Late Pliocene: southern Peru. Recent:

northern Peru to Chile, southern Brazil to Argentina.

Remarks: A single incompletely preserved specimen

of an agathistomine was found between Yauca and Chala

in bioclastic deposits just below the highest marine ter-

race at 200 meters above sea level. Associated taxa that

are either locally or entirely extinct |Prisogaster valenciai

DeVries, 2006; Acanthina triangularis DeVries, 2003;

Chorus giganteus (Lesson, 1830); Concholepas camerata

DeVries, 2000; Xanthochorus xuster DeVries, 2005] are

indicative of a late Pliocene age (DeVries, 1997; 2000;

2003: 2005: 2006).

The closely spaced beaded spiral cords on the south-

ern Peruvian agathistomine resemble those on speci-

mens of Agathistoma verrucosum McLean, 1970, and A.

pictum McLean, 1970, Panamic species which pre ssently

range as far south as northern Peru (Alamo and

Valdivieso, 1997), but the base of the Peruvian Pliocene

specimen is not as flattened as it is on specimens of the

northern Peruvian species and the spiral cords are more

closely spaced, suggesting an assignment to A. patagoni-

cum.

Genus or Subgenus indeterminate

Tegula (s.1.) tridentata (Potiez and Michaud, 1838)

(Figures 66, 65)

Monodonta tridentata Potiez and Michaud, 1838, vol. 1, p. 321,

pl. 29, figs. 16-17.

Trochus tridentatus Potiez and Michaud.—Philippi, 1546, Die

Kreiselschnecken oder Trochoideen, p. 153, pl. 25, fig. 3.

Tegula tridentata (Potiez and Michaud).—Dall, 1909: 176; Car-

celles and Williamson, 1951: 262; Herm, 1969: 91; Aldea

and Valdovinos, 2005: fig, SG.

Tegula (Chlorostoma) tridentata (Potiez and Michaud,

1838).—Marincovich, 1973: 24, fig. 43; Véliz and Vasquez,

2000; p. 761, fig. 1C; Guzman et al., 1998: 36, fig. 24;

Forcelli, 2000: 90, fig. 60.

Trochus tridens Hupé, 1854: 145,

Material Examined: UWBM 97898, DV 1372-1, Re-

cent, L = 7.0, W = 9.9: UWBM 97899, DV es Re-

cent, lot of 5: UWBM 97900, DV 381-5, middle Pleis-

tocene, L = (11.7), W = 14.0.

Occurrence: Middle Pleistocene: southern Peru to

central Chile. Recent: northern Peru to southern Chile.

Remarks: Tegula tridentata is a small, high-spired,

purple-black species with an excavated white base. The

exterior is usually smooth, but some specimens have low

broad primary spiral cords on juvenile whorls. A narrow

open aanbilicis is bordered by a white columella with

three teeth. The two teeth closest to the axis lie at the

end of umbilical spiral cords, the outermost of which is

mostly submerged beneath a thick umbilical veneer. The

third tooth is situated adjacent to the floor of the aper-

Page 174

Figures 66, 65.

Oblique spiral view. 68. Oblique basal view. Figures 67, 69-72.

THE NAUTILUS, Vol. 121, No. 4

Tegula (s.1.) tridentata (Potiez and Michaud, 1838). UWBM 9789S, DV 1372-1, Recent, width = 9.9 mm. 66.

Tegula (s.1.) melaleucos (Jonas, 1$44). 67. UWBM 97901, Paracas

Hotel, Recent, oblique spire view, width = 23.9 mm. 69. OSU 37609, DV 211-3, late Pleistocene, lateral view, length = 28.2 mm.

70. UWBM 97901, oblique basal view. 71. OSU 37609, apertural view. 72. OSU 37609, oblique basal view, width = 29.0 mm. Figure

73. Tegula (s.1.)

(?) rubroflammulata (Koch in Philippi, 1843). OSU 3

7610, DV 341, early Pleistocene, oblique basal view, width

= 22.0 mm. Figures 74-78. Tegula (s.1.) masiasi new species. 74. UWBM 97904, DV 475-1, early Miocene, lateral view, length

= 16.6 mm. 75. UWBM 97903, DV 1019-1

. holotype, middle Miocene, oblique spire view, width = 15.9 mm. 76. UWBM 97903,

basal view. 77. UWBM 97905, DV 1648-1, early Miocene, lateral view, length = 14mm. 78. UWBM 97905, basal view showing edge

of umbilical area, width = 16.9 mm.

ture. The inside of the outer lip often has four to six short

elongate teeth. In the latter three characters the species

resembles Cantallocostoma quadricostatum, with which

it has been grouped using mitochondrial DNA sequences

by Hellberg (1998), who placed the species with Agathi-

stoma. In its color and obsolete spiral sculpture, how-

ever, “tridentata” specimens greatly resemble Peruvian

species of Chlorostoma, to hich they were assigned by

Marincovich (197: and Guzman et al, (1998). The only

fossil example of T. tridentata in Peru comes from a

middle neler. marine terrace bed near San Juan de

Marcona, southern Peru.

Tegula (s.1.) melaleucos (Jonas, 1544)

(Figures 67, 69-72)

Trochus melaleucos Jonas, 1544: 169; Philippi, 1846, Die kKrei-

selschnecken oder Trochoideen, p. 185, pl. 28, fig. 16.

Tegula melaleucos (Jonas). —Dall, 1909: 239.

Tegula ( Agathistoma) melaleucos (Jonas, 1S44).—Keen, 1971:

340, fig. 106; Alamo and Valdivieso, 1997: 13, fig. 22.

Not Tegula (Agathistoma) melaleucos (Jonas, 1S44).—DeVries

1986: 515, pl. 27, figs. 1, 2, 10, 12 [possibly Tegula (s.1.)

rubroflammulata ( (Koch in ayer 1843) ].

Material Examined: OSU 37609, DV 211-3, late

Pleistocene, L = 28.2, W = 29.0: UWBM 97901, Paracas

Hotel, Recent, L 17.3, W 23.9: UWBM 97902,

northern Peru, Recent, L = (11.5), W = 20.1. Specimens

assigned to Tegula melaleucos by DeVries (1986) but

more likely belonging to Tegula (s.1.) rubroflammulata;

OSU: 37610, DV 341, early Pleistocene: L = 22.0, W =

22.0; OSU 37611, DV 341, L = (14.8), W = (18.0).

Occurrence: Recent: northern Peru; rarely in south-

ern Peru.

T. DeVries, 2007

Remarks: Specimens of Tegula melaleucos are charac-

terized by a strongly bicarinate periphery, a line of pro-

tractive nodes between the senale ry and suture, and

protractive brown stripes laterally and basally, where

they spiral into an open umbilicus. A single spiral cord

emerges from the umbilicus and is truncated by a ar

umellar ridge that ends in a basally projecting tooth.

second prominent tooth protrudes at the juncture of :

columella with the floor of the aperture, and additional

small teeth may occur along the edge of a beveled ledge

that passes just inside the floor of the aperture.

Impertectly preserved specimens of Tegula from the

uppermost Pliocene / lower Pleistocene Mancora Ta-

blazo of northern Peru (Figure 73; DeVries, 1986; 1988)

have less impressed sutures and more convex profiles

than typical specimens of T. melaleucos and nodes near

the suture that are not protractive. These specimens are

better referred to T. rubroflammulata, a Recent species

that had been reported to range only as far south as

Colombia (Keen, 1971).

Tegula (s.1.) masiasi new species

(Figures 74-78)

Diagnosis: Spire whorls with broad spiral cords; um-

bilicus narrow, open; shell lacking purple-black outer

layer.

Description: Shell conical, up to 16 mm wide. Spire

angle about 70 degrees. Periphery at base; angular. Su-

tures appressed to impressed. Protoconch anion: te-

leoconch with at least four flat-sided to convex whorls.

Axial sculpture absent: colabral growth lines strongly

prosocline. Spiral sculpture absent or with several broad,

low tee cords on spire. Outer shell layer lacking

purple-black color. Base flattened to weakly convex,

without visible spiral sculpture. Inner side of outer lip

smooth. Umbilicus open. Columella thin, with at least

one tooth at end of umbilical cord. Parietal flange barely

overhanging umbilicus.

Type Locality:

km east of mouth of Quebrada Gramonal, on bluff over-

looking road to Funda $ Santa Rosa (Figure 79).

14°45'50"S, 75°30'22"/W (Lomitas 1:100,000 quad-

rangle). Lower middle Miocene.

Type Material: }UWBM 97903, DV 1019-1. holotype,

early middle Miocene, L = (7), W = 15.9.

Other Material Examined: U\WBM 97904, DV 47S8-

ie early Miocene, L = 16.6, W = (21.8); UWBM 97905,

DV 1648-1, early Miocene, L = 14.0, W = 16.9: MUSM

INV 143, DV 1648-1, L = 14.6, W = 19.8

Occurrence: Early to early middle Miocene: southern

Peru.

Etymology: Named in honor of Antonio Masias, pe-

troleum geologist from Arequipa, Peru, who has pro-

vided advice on Peruvian matters since we both attended

Oregon State University in 1977-1978.

DV 1019-1, Gramonal, about one-half

AS a de Ullujalla

@ Locality-sample

0 5 km

SCALE

Contour interval is

100 m.

600 }

eee

Figure 79. Type locality (DV 1019-1) of Tegula (s.1.) masiasi

new species.

Remarks: Specimens of Tegula (s.|.) masiasi are the

oldest known teguline »s from the Pisco Basin: they occur

near the base of a depositional sequence that unconform-

ably underlies the Pisco Formation near Cerros Colorado

(Figure 78), which implies the lower portion of the up-

permost Oligocene to lower middle Miocene Chilcatay

Formation (De Vries, 1998), as well as a few tens of

meters above the base of the Pisco Formation, which

indicates an early middle Miocene age (DeVries and

Schrader, 1997). Despite the poor state of preservation,

it does appear that these early and middle Miocene

specimens are not chlorostomines, which. first appear

with their purple-black outer shell layer in beds at Alto

Grande at about 9-10 Ma (Muizon and DeVries, 1955:

DeVries and Schrader, 1997).

DISCUSSION

Fossils found since 1990 offer a new perspective on the

origin of Tegulinae. Tegulines had been thought to have

arisen during the early or middle Miocene (Hickman and

McLean, 1990), based on knowledge of fossil taxa from

Japan and California [e.g., Tegula thea Nomland, 1917

Page 176

THE NAUTILUS, Vol. 121, No. 4

and T. varistriata Nomland, 1917, Santa ae beds,

California, upper Miocene (Nomland, 1917); T. dalli Ar-

nold, 1907, and T. dalli var. inornata ‘ciold: ae To-

panga Group, California, middle Miocene (Arnold, 1907;

Yerkes and Campbell, 2005); T. dalli arnoldi Addicott,

1970, Olcese Sand, California, uppermost lower Miocene

(Addicott, 1970; Sanchez and Prothero, 2003) ]. The dis-

covery of T. jeanae Squires and Saul, 2005, a late Cam-

panian species from the Chico Formation of California

with many attributes of Chlorostoma (ventricose whorls,

black-brown outer shell layer, absence of beaded spiral

cords), recasts discussions of teguline phylogeny (Squires

and Saul, 2005). A flat-sided, spirally bended Cretaceous

teguline, the late Maastrichtian T. ovallei Philippi, 1887,

from central Chile (Bandel and Stinnesbeck, 2000), fur-

ther demonstrates ri pre-Miocene diversity and. geo-

graphic distribution of tegulines.

As a consequence of these Cretaceous discoveries, an

evolutionary scenario rejected by Hickman and McLean

(1990), which placed the plesiomorphic Tegula near the

base of the trochid group, gains credence, whereas sce-

narios inspired by the molecular data of Hellberg (1998)

that eae a strictly late Neogene timeline for teg-

uline phylogeny are undermined by these Cretaceous

data, as sell as by early, middle, and late Miocene oc-

currences of tegulines in Peru.

The following observations provide some further con-

straints on phy logene tic hypotheses related to Tegulinae.

PERUVIAN CHLOROSTOMINE GROUP

The Peruvian chlorostomine group is comprised of four

modern species: Chlorostoma atrum, C. ignotum, C. eu-

ryomphalum, and C. luctuosum. Cradatoas of sculpture

indicate the latter two species may be one; C, luctuosum

would be the senior synonym. Adding C. quipua extends

the record of Peruvian ealonsctowines to 9 Ma (Muizon

and DeVries, 1985). This late Miocene occurrence pre-

cludes Hellberg’s (1998) molecularly based hypothesis

that these chlorostomines appeared in western South

America during the late Pliocene and constitute a sister

group to a North Atlantic Pliocene group of Agathistoma

Species.

The flattened base, open umbilicus, and spiral threads

of Chlorostoma quipua indicate that it is most closely

related to the modern C. luctuosum. Some specimens

of C. quipua from upper Pliocene beds above Playa

Huacllaco also have broad spiral cords like those on some

specimens of modern C. atrum. A single upper Pliocene

Hu: 1c lle ico spe cime n hi iS strongly convex spire al cords like

those on specimens of the modern Chilean C. ignotum.

This morphological variation suggests the onset of a ra-

diation of South American chlorostomines near the end

of the Pliocene. The late Pliocene was also a time of

species-level molluscan mass extinction throughout the

Peruvian Faunal Province (DeVries, 2001)

The origin of chlorostomines in Peru and Chile is not

known. Tegulines are found, rarely, in lower and middle

Miocene beds of southern Peru (T. (s.].) masiasi; Figures

74-78), but they do not resemble late Neogene Peruvian

Chlorostoma. Of Chilean Miocene species assigned to

Chlorostoma (Nielsen et al., 2004), none edu the

purple-black outer shell layer that characterizes the type

species of Chlorostoma and most moder Peruvian chlo-

rostomines. At three mm in length, Tegula austropacifica

Nielsen, Frassinetti, and Bandel, 2004, is extraordinarily

small for a chlorostomine. Tegula chilena Nielsen, Fras-

sinetti, and Bandel, 2004, lacks the flattened base of

chlorostomine species. Tegula matanzensis Nielsen,

Frassinetti, and Bandel, 2004, resembles Californian

specimens of C. funebralis, as the authors note, but the

specimen lacks critical portions of the columella from

which a better comparison can be made.

Chlorostomines appeared in southern Peru at the

same time as the turbinid genus, Prisogaster Mérch,

1850 (DeVries, 2006), and mollusks with Panamic affini-

ties (DeVries, 2006), an immigration pattern consistent

with either a western North American or boreal Asian

origin for the group. Molecular data (Hellberg, 1998)

‘acest Peruvian chlorostomines are more similar to

Californian than Asian taxa. Shell characters are equivo-

cal on this point. Protractive stripes, present on speci-

mens of the oldest Peruvian chlorostomine, C. quipua,

are also seen on some specimens of Californian C.

gallina, C. rugosum, and Asian C. nigerrimum and C.

rusticum. Protractive ribs, which occur in rare examples

of C. quipua, are found on some specimens of the Cali-

fornian “Tegula” brunnea and several Asian species. Im-

bricate subsutural spiral cords, which occur rarely on

juvenile whorls on specimens of C. atrum, are most char-

acteristic of the Californian C. fiumebralis (Figures 5, 6),

are weakly developed on specimens of the Califor nian C.

gallina and C. rugosum, and are not seen on specimens

of Asian species.

Peruvian chlorostomines are distinguished from all

Californian chlorostomines and all Asian taxa except a

few specimens of C. rusticum by possessing a very thin

and expansive umbilical veneer ‘that does not bury the

spiral umbilical cord. A thicker umbilical veneer di rapes

across the spiral umbilical cords in Californian and Asian

specimens, largely burying the spiral umbilical cord and

leaving visible only a blunt adaxial columellar tooth, a

tooth at the base of the columella, and an interve ning

depressed nacreous wedge (e.g., C hlorostoma funebralis;

Fie. 6). Peruvian Ghilbrestoutines also lack a second well

developed tooth at the base of the columella, a character

usually seen on Californian and Asian chlorostomine

specimens (Fig. 6). These two derived characters—thin

umbilical veneer, obsolete basal columellar tooth—-may

indicate that Peruvian chlorostomines are a sister group

to Californians Asian chlorostomines, with a common an-

cestor in the North Pacific Ocean older than late Mi-

ocene.

CANTALLOCOSTOMA GROUP

Cantallocostoma is an endemic western South American

genus characterized by beaded spiral cords and two

T. DeVries, 2007

Page 177

adaxially situated spiral umbilical cords. Cantallocostoma

panistostum appears first in upper Miocene beds of

southern Peru with other Panamic species (DeVries,

2002). It and the extant C. quadricostatum are unlike any

Neogene or Recent teguline from Peru or Chile. Speci-

mens of Cantallocostoma share with specimens of Inti-

stoma the presence of two adaxially situated spiral um-

bilical cords (mostly covered by a thick umbilical veneer

in specimens of Intistoma) and an erect columella with

little in the way of a parietal flange or callus. Genetic data

of Hellberg ( (1998), however, show no close affinity be-

tween C. quadricostatum and I. aureotinctum.

INTISTOMA GROUP

Intistoma has been created to include two very similar

species, the modern Californian I. aureotinctum and

early Pliocene Peruvian I. pirqua. Their distinctive spiral

sculpture ( (strong bicarinate periphery, protractive sub-

sutural nodes, thre -e broad basal spiral cords), absence of

a purple-black outer shell layer, absence of apertural

teeth, and the isolation of T. aureotinctum in mtDNA

phylogenies (Hellberg, 1998) indicate that neither Chlo-

rostoma nor Agathistoma properly encompass these taxa.

Two other groups, one consisting of the modern north-

ern Peruvian / Panamic Tegula (s.1.) melaleucos and pos-

sibly T. (s.1.) rubroflammulata, the other comprising Mi-

ocene and Pliocene species from California [e.g., T. (s.1.)

dalli\, exhibit the distinctive subsutural spiral row of pro-

tractive nodes of Intistoma, but both lack the three broad

basal spiral cords that characterize the new genus. The

Californian Neogene species also lack the differentiation

of spiral sculpture (coarse primary spiral cords, fine ter-

tiary spiral threads overrunning primary spiral cords and

interspaces) that is visible on well-preserved specimens

of I. aureotinctum.

Tegula (s.l.) tridentata (Potiez and Michaud, 1838)

Tegula (s.1.) tridentata is a small teguline with a record in

Peru and Chile extending no farther back than the

middle Pleistocene. The distinctive purple-black exterior

is shared with Peruvian chlorostomines, but the arrange-

ment and number of columellar teeth is like that of

Panamic species of Agathistoma. Mitochondrial DNA

data (Hellberg, 1998) are not helpful on this point of

phylogeny, as T. (s.1.) tridentata usually clusters with

Cantallocostoma quadricostatum, which it resembles in

only one significant shell character: two adaxial umbilical

spiral cords, both terminating in a columellar tooth. For

now, the proper phylogenetic assignment of T. (s.].) tri-

dentata remains elusive.

AUSTRAL AGATHISTOMA

Panamic agathistomines, which are so speciose in warm

waters of the Panamic Faunal Province, have been no-

tably unsuccessful in penetrating the cold waters of the

Peruvian Faunal Province. The only fossil agathostomine

from western South America is a specimen from upper

Pliocene beds of southern Peru assigned to Agathistoma

patagonicum. A Miocene agathistomine reported from

central Chile, Agathistoma antiquum Nie Isen, Fras-

sinetti, and Bandel, 2004, a Miocene occurrence of A.

patagonicum reported from Argentina (Ihering, 1907;

del Rio, 1998), and an extensive record of modern Aga-

thistoma patagonicum from southern Brazil to southern

Argentina, including the Magellanic waters of Argentina

(Carcelles and Williamson, 1951), if the synonymy of A.

fuscesens and A. orbignyana with A. patagonicum is ac-

acer (Forcelli, 2000), may point to a Miocene austral

origin for the subgenus ( Nielsen et al., 2004), rather than

the Pliocene western Atlantic origin suggested by Hick-

man and McLean (1990). ), Alternatively, the monophy ly

of Agathistoma might be suspect; compact, beaded,

rule toothed austral tegulines with or without open um-

bilici might constitute a long-lived sister group to a group

of Caribbean and Central American species.

CONCLUSIONS

Pre-late Miocene tegulines in southern Peru are exceed-

ingly rare, poorly preserved, and bear little resemblance

to late Neogene or extant taxa. During the early late

Miocene, at least two lineages of tegulines, Chlorostoma

and Cantallocostoma, entered Peruvian waters. [t is un-

clear whether these two genera originated in California

or Asia. Other trochids, namely Diloma Philippi, 1845,

are thought to have dispe srsed across the Equator and

across fie Pacific Ocean from Australia, rafted by buoy-

ant fragments of the brown kelp, Durvillaea Bory de

Saint-Vincent, 1826 (Donald et al., 2005). lies of Pe-

ruvian Tegula likewise live upon on brown kelp (Lessonia

Bory de Saint-Vincent, 1825) (Véliz and Vasquez, 2000;

V. Mogollon, pers. comm., 2006), and thus may have

been rafted to western South America from California or

Asia in the same manner as Diloma.

Species of Chlorostoma and Cantallocostoma re-

mained relatively unchanged until the end of the

Pliocene, when a mass extinction swept away 50 percent

of molluscan species in the Peruvian Faunal Province

(DeVries, 2001). At that time, chlorostomines experi-

enced a mini-radiation in southern Peru or Chile, with

one species, C. atrum, eventually spreading to southern

Argentina (Carcelles and Williamson, 1951). Cantallo-

costoma panistostum was replaced at the same time by

the modern C. quadricostatum.

Another lineage of tegulines, represented by the early

Pliocene Intistoma pirqua, appeared on Peruvian shores

by the early Pliocene. Although the genus is now extinct

in Peru, it persists in California in the guise of I. au-

reotinctum. It is likely that these species, with their br« vad

basal spiral cords, are not related to the “dall” lineage of

Californian tegulines that ranged from the early Miocene

to Pliocene (Addicott, 1970).

Page 178

THE NAUTILUS, Vol. 121, No. 4

ACKNOWLEDGMENTS

I would like to thank G. Collado (Facultad de Ciencias,

Universidad de Chile), L. Groves and J. McLean (Natu-

ral History Museum of Los Angeles County, USA), S.

Rugh (San Diego Museum of Natural History, USA), J.

Vasquez Umveradad Catolica del Norte, Coquimbo,

Chile), D. Véliz (Universidad de Chile, Santiago), G. Pas-

torino (Museo Argentino de Ciencias Naturales, Buenos

Aires), S. V. Mogollon ( (Universidad Nacional Federico

Villarreal, Lima, Peru), and W. J. Zinsmeister (Purdue

University, West Lafayette, Indiana, USA) for access to

museum collections, loans of specimens, or valuable ad-

vice. Helpful critiques were offered by D. Zelaya and M.

Griffin (Museo de Ciencias Naturales, La Plata, Ar gen-

tina). Support for field research was provided in part by

a Fulbright scholarship in 1999.

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APPENDIX

lusca: Gastropoda) en el norte de Chile: consideraciones

ecologicas y taxonémicas. Revista Chilena de Historia

Natural 73: 757-769.

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California. United States Geological Survey, Open-File

Report 2005-1019.

Locality-samples. GPS = latitude and longitude coordinates obtained by a hand-held GPS unit. GE = coordinates

obtained from satellite images available on-line from Google

DV 211-3

DV 341-1

DV 381-5

quad angle). Middle Pleistocene,

DV 382-1

75°05'26"W (

DV 3958-1

DV 401-1

Recent.

DV 463-1

DV 470-1

Lower terrace, five km north of Chala (Chala 1:100,000 quadrangle).

Above ee road from Hacienda Tunca to Quebrada Huaricangana; hillside of brown sandstone.

Earth™.,

Punta Lobitos, northern Peru, western point, terrace surface midway between sea cliff and inshore

edge of deposit. 04°27'12"S, S1°17'25"W (GE; Lobitos 1:100,000 quadrangle).

Quebrada Mogollon, northern Peru. Upper coquina of Mancora Tablazo. Lower Pleistocene.

San Juan- Lomas road, km 47.5. Uppermost coquina. 15°22'59"S, 75°03" 11"W (

Upper Pleistocene.

San Juan 1:100,000

San Juan / Lomas road, kilometer marker 50, flat-topped knoll south : highway. 15°22'02"S,

San Juan 1:100,000 quadrangle). Remnant of marine terrace.

Playa Canastones, Bahia de la Independencia, Peru (Punta Grande 1:100,000 quadrangle). Recent.

Hueco La Zorra, north end of beach. 14°02'31"S, 76°15'51"W (Punta Grande 1:100,000 quadrangle).

Upper Pleistocene.

14°56’ S. 75°09’ W (GE: Palpa 1:100,000 quadrangle). Lower Pliocene.

DV 478-2

Lomas : hileatay, northeast end of outcrop. 14°11'42"S 76°06'57"W (Punta Grande 1:100,000 quad-

rangle). Chile ‘atay Formation, lower Miocene.

DV 571-1

Alto Grande, about one km south of intersection with abandoned paved road to San Juan de Marcona,

on south-facing hillside west of Panamerican Highway; one of several shell banks. 15°26'57’S,

74°52'06"W (Acari 1: 100,000 quadrangle),

Yauca, roadcut on western side of Panamerican Highway as it descends to valley floor. 15°39'49"S,

74°31'50"W (Yauca 1:100,000 quadrangle).

Camu about one-half km east of canyon mouth. 14°45'50"S, 75°30'22"W

DV S09-1

DV 1019-1

quadrangle). Middle Miocene.

DV 1029-1]

quadrangle). Lower Pliocene.

DV 1032-2

). Middle upper Miocene.

Pisco formation. Lower Pliocene.

(Lomitas 1: 100,000

Yauca Depression, west of Pana merican Highway. 15°39'29"S, 75°35'08"W (GPS, Yauca 1:100,000

Morro Abra de los Chaparrinos, descending from highest terrace level, north and south of second

curve in Panamerican Highway. 15°52'59"S, 74°10'05"W (Chala 1:100,000 quadrangle). Upper

Pliocene.

DV 1252-1

above non-marine de posits. 15°45'56"S,

DV 1254-Bal 6

rangle). Upper lower Pliocene.

DV 1254-Bal 10 Section along Panamerican Highway,

meters above basement rocks in measured section. 15°53'25"S

quadrangle). Upper Pliocene.

Quebrada de la Vaca, roadcut along Pa inamerican Highway, south of south wall, uppermost terrace

74°18'50"W (GPS; Chala 1:100,000 quadrangle).

Section along Panamerican Highway, ten km southeast of Chala and above Playa Huaclla ico. 35 meters

above basement rocks in measured section.

15°53/25"S, 74°09'52"W (GPS; Chala 1:100,000 quad-

ten km southeast of Chala and above Playa Huacllaco, 47.5

ea 95"5

— 74°09'52"W (GPS: Chala 1:100,000

DV 1254-] Sacaco, shell banks southwest of north-south road to farmhouse (chacra). 15°33'03 S', 74°43'50"W

(GE; Yauca 1:100,000 quadrangle). Lower Pliocene.

DV 1355-1] Quebrada Pongo, one km upstream _ juncture with Quebrada Caracoles. 15°30'22"S, 74°45'40"W

(GPS; Yauca 1:100,000 qui adrangle) Upper Pliocene.

DV 1372-1] tocky beach on northwestern side ge Punta Lomas (Acari 1:L00,000 quadrangle). Recent.

DV 1418-1] East side of Acari Depression, 15°34'50"S, 74° 36'59°W (GPS; Yauca 1:100,000 quadrangle). Upper

Pliocene

T. DeVries, 2007

Page 15]

DV 1598-1

DV 1599-1

DV 1635-1

DV 1648-1

DV 1713-1

JM 82-19

JM 82-20

WIZ 345

Isla Ipun

Paracas Hotel

Roadcut oO Panamerican Highway, descent from north into Yauca. Shell beds. 15°39'49"S,

74°31'50"W (GPS: Yauca 1:100,000 quadrangle). Lower Pliocene.

ae La Zorra, north end of beach (see DV 401-1). Recent.

Yauca re aes west of Panamerican Highway. 15°39'33"S, 75°34/54"W (GPS; Yauca 1:100,000

quadrangle). Lower Plioce ne.

Westward- facing side of valley, southwest of Cerros Colorado. 14°22'25"S, 75°53'52”/W (GPS; Punta

Grande 1:100,000 quadrangle).

Marine terrace on east side of Panamerican Highway north of road to Acari. 15°36'09"S, 74°41'08’"W

(GPS; Yauca 1:100,000 quadrangle). Lower Pleistocene.

Cerro El Huevo, northeast of San Juan de Marcona. 15°18’ S, 75°09’ W (San Juan 1:100,000

quadrangle). Upper Pleistocene.

Cerro E] Huevo, northeast of San Juan de Marcona. 15°18’ S, 75°09" W (San Juan 1:100,000

quadrangle). Upper Pleistocene.

Coquimbo, Chile. Pleistocene. Approximately 29°58" S, 71°20' W (GE).

Isla Ipun, Chile, shores of eastern embayments. 44°38’ S, 74°44’ W (GE). Recent.

Beach south of Hotel Paracas, facing Bahia Paracas, southern Peru. 13°50'09"S, 76°15’ 19”"W (GE,

Pisco 1:100,000 quadrangle).

THE NAUTILUS 121(4):182-190, 2007

Page 182

Three new species of Paryphantopsis (Gastropoda: Pulmonata:

Charopidae) from the Nakanai Mountains, New Britain,

Papua New Guinea

John Slapcinsky

Robert Lasley

Florida Museum of Natural History

and Department of Zoology

University of Florida

Gainesville, FL 32611 USA

[email protected]

ABSTRACT

Recent surveys (February-March, 2005) of the terrestrial snail

fauna of the Nakanai Mountains, central New Britain, Bismarck

Archipelago have uncovered several undescribed species, in-

cluding three new species of Paryphantopsis, a diverse genus of

charopid snails, previously believed to be endemic to mainland

New Guinea and adjacent islands of the Louisiade Archipelago.

The three species are described using shell, genital, and udu ww

morphology. Although the land snail fauna of New Britain is

arguably the best sampled in Papua New Guinea, there has

been little sz unpling in the interior mountains of the Nakanai,

Whiteman, Baining, and Willaumez ranges, which harbor pre-

viously undetected species of terrestrial snails.

Additional Keywords: Pulmonata, Charopidae, Paryphantopsis,

New Britain, Papua New Guinea

INTRODUCTION

This is the third in a series of reports on the results of

recent field surveys for terrestrial mollusks from Papua

New Guinea. The previous two reports focused on spe-

cies of Paryphantopsis from the

mainland New Guinea (Sk apcinsky, 2005) and nearby is-

lands of the Louisiade Archipelago (Slapcinsky, 2006).

This report reviews Paryj shantopsis species collected

during three weeks of field surveys in February and

March 2005 from the N

Pomio, East New Britain Province, Papua New Guinea

Figure 1). Paryphantopsis, a diverse genus of charopid

snails endemic to New Guinea and surrounding islands,

is comprised of 23 described species | (Solem, 1970:

Slape insky, 2005: Sl ape ‘insky, 2006) that are distributed

from Papua (Irian Jaya) to the Louisiade Archipe lago

and that, prior to this ‘study, were not known from New

Britain

Nearly all species of Paryphantopsis have distributions

stricted to single mountain ranges or islands where

eastern peninsula of

Nakanai Mountains northwest of

they are found in hilly or mountainous terrain from 60 to

4000 meters elevation. In upland habitats they can be the

most abundant snail species reaching densities of more

than 10 individuals per square meter on Sudest and Ros-

sel Islands in the Louisiade Archipelago (personal obser-

vation). Despite their small size (4-12 mm), their bright

yellow body coloration and ia ul activity in exposed

locations on tree trunks, vegetation and rotting wood

make them among the most hanes snail species. These

showy snails are under- sampled and recent surveys have

near ly doubled the number of recognized species. Fur-

ther sampling in New Guinea and surrounding islands

will certainly uncover additional species of Pary) phantop-

sis as well as many other more cryptic snail species.

New Britain, a large volcanic island, 35145 km?, ex-

tending from 148° to 152° E longitude and from 4° to 7°

S latitude, emerged in the late Miocene ( S—LO mya). The

island’s basement rocks were deposited by volcanic ac-

tivity, between the upper Eocene and the middle Oli-

gocene. Volcanic activity ceased in the early Miocene

le ading to regional subsidence and the de “position of ex-

tensive reef comple xes. At the end of the Miocene, sub-

duction of the Solomon plate under the Bismarck plate

led to renewed volcanism, resulting in a chain of young

arc volcanoes along the northwestern coast of New Brit-

ain (Woodhead et al. 1998). At the same time, reef com-

plexes along the south and eastern coasts of the island

were apidly uplifted, creating extensive karst mountains.

One of these ranges, the Nakanai Mountains, contains

the Southern He ‘misphe re’s deepest caves, some nearly

1200 meters in depth (Audra et al. 2001). New Britain's

isolation, complex geology, and extensive raised lime-

stone have allowed the radiation of a diverse and 1: urgely

endemic land snail fauna (Rensch, 1934; 1937 ) that is

among the best surveye din Papua New Guinea. How-

ever, rough karst terrain and lack of roads have limited

nearly all surveys to coastal lowlands, and most taxa from

Nakanai, Whiteman, Baining,

and Willaumez ranges are still poorly surveyed (Beehler,

interior mountains of the

J. Slapcinsky and R. Lasley, 2007

Page 183

151.4°E 151.6°E

<~....

Nakanai 2

Mountains | PRS) ) Sas Ss

} / A ge TRS

\ / LASSE

Jacquinot Bay on

Figure 1. Distribution of Paryphantopsis in New Britain,

Papua New Guinea; 0 = other sites sampled.

1993), even for macrofauna such as mammals (Flannary,

pilasters, PR = penial retractor muscle, SD = spermathecal

duct, SP = spermatheca, VA = vagina, VD = vas deferens.

Terminology of vegetation types follows Paijmans (1976).

Specimens are deposited in the following institu-

tions: Bernice P. Bishop Museum, Honolulu (BPBM);

Florida Museum of Natural History, Gainesville (UF);

Natur-Museum Senckenberg, Frankfurt (SMF): Papua

New Guinea National Museum, Port Moresby

(PNGNM); Wroclaw University Museum of Natural His-

tory (MNHW),.

SYSTEMATICS

Family Charopidae Hutton, 1884

Genus Paryphantopsis Thiele, 1928

Type Species: Flammulina (Paryphantopsis) lamel-

ligera Thiele, 1928, by original designation.

Paryphantopsis corolla new species

(Figures 2-7, Table 1)

7 ; : ; ae ; Holotype: UF 366508, J. Slapcinsky, 25 February

1995: 12) and birds (Orenstein, 1976). The invertebrate : YP . J I : 3 /

: icon 2 Negi . elie 2005.

fauna of New Britain’s interior mountains is almost en-

tirely unknown. This is disturbing because intensifying Paratype: UF 366453 (1 specimen), J. Slapcinsky, 25

land usage, including logging and the establishment of oil

palm plantations (McAlpine and Fryne, 2001), threatens

to deforest extensive areas of New Britain before they

can be adequately inventoried, potentially leading to

largely undocumented losses in biodiversity.

MATERIALS AND METHODS

Specimens were hand-collected, drowned overnight, and

then preserved in 75% ethanol. Gross anatomical dissec-

tions were made under 75% ethanol using a dissecting

microscope. Radulae were isolated from dissected buccal

masses using 5% sodium hypochlorite solution. Scanning

electron micrographs of radulae were made using a Field

Emission SEM. Line drawings of the genital anatomy

were made from digital images, and measurements were

taken using an ocular micrometer. Shell measurements

were made as figured in Slapcinsky (2005). 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

February 2005.

Type Locality: Papua New Guinea, East New Britain

Province, New Britain, 12 km northwest of Marmar

Village on the trail to Pakia Village, 5.432° S, 151.460° E,

900 meters altitude (Figure 1).

Habitat: Collected in mixed hill forest on vegetation

within 1 m of the ground.

Description: The adult shell is depressed-globose,

small for the genus, 3.5-3.6 mm (mean = 3.55, see

Table | for sample size and standard deviation) in diam-

eter and 2.2-2.3 mm (mean = 2.25) in height, with 2.5-

2.6 (mean = 2.55) rapidly expanding whorls (Fig-

ures 24). The apical surface of the whorls flattens be-

tween the deep suture and the shell periphery which is

slightly angular a little above its mid-point. The basal

surface of the whorl is evenly rounded from the angular

periphery to the umbilicus. The spire is slightly elevated,

0.1—0.2 mm (mean = 0.15). Teleoconch whorls descend

slowly and regularly and shell height/diameter ratio is

0.61-0.66 (mean = 0.63). Approximately 1.3 rounded

protoconch whorls are sculptured with L3 rows of spiral

Table 1. Measurements in mm of undamaged adult shells of three species of Paryphantopsis.

Species N H D SH SD AH AW W

P. corolla 2 Mean+=SD 2.25+0.07 355+0.07 0.15+0.07 1604000 160+000 215+007 2.55 +0.07

Range 2.9-2.3 3.5-3.6 0.1-0.2 1.6-1.6 1.6-1.6 2.1-2.2 2.5-2.6

P. fragilicosta 6 Mean+SD 3.43+018 5.00+020 0.134005 2334012 248+016 292+0.16 3.00+0.09

: Range 3.2-3.7 4,8-5.3 0.1-0.2 2.9-9.4 2.3-2.7 2.83.2 2.9-3.]

P. nucella 10 Mean+SD 3.75+0.32 5.22+0.23 0.18+0.04 2.06+020 2.784027 316+0.22 281+0.12

Range 3.34.1 4.9-5.7 0.1-0.2 1.8-2.5 2.4-3.3 2.9-3.6 2.6-3.0

N = number of specimens, H = height, D = diameter, SH = spire height, SD = spire diameter, AH = aperture height

AW = aperture width. W = number of whorls

Page 184 THE NAUTILUS, Vol. 121, No. 4

Figures 2-7. Paryphantopsis corolla, 2-4. Shell, Holotype UF 366508, Scale bar = 1 mm; Figure 5, Genitalia, UF 366453.

Scale bar = 1 mm. 6-7. SEMs of radula, UF 366453. 6. Central and lateral teeth. 7. Marginal teeth. Scale bars = 10 xm,

mss

J. Slapcinsky and R. Lasley, 2007

Page 185

pits, which continue on the teleoconch where the pits

elongate nearly fusing to form incised spiral lines. These

apical pits are typical of nearly all species of Paryphan-

topsis as, for example, in Pary) shantopsis louisiadarum

(see Solem, 1959: plate 13, fig 6). Approximately every

third growth line is accentuated with a pe sriostracal ex-

tension that bears a sharply pointed triangular process at

the shell periphery. These processes are approximately

0.2 mm in length and 0.3 as wide at the base. The pro-

toconch and teleoconch whorls are uni formly brown ex-

cept for the columellar edge of the peristome and um-

bilicus, which are slightly darker brown. A reflection of

the peristome covers approximately 0.4 of the umbilicus.

The aperture is large and ovate with an aperture:width to

aperture:height ratio of 1.31-1.35 mm (mean = 1.34).

The external body color is bright yellow in life and

there are no bands or other color patterns. Specimens

preserved in ethanol fade to uniform cream. The head is

short similar to other species of Paryphantopsis and the

posterior of the foot is slightly shorter than average for

the genus.

The vas deferens is 0.2 the diameter of the head of the

epiphallus which bears a short subapical diverticulum

approximately 0.1 the length, and 0.6 the diameter of the

epiphallus where they join (Figure 5

the epiphallus is uniform in diame the basal 0.3 is

twice as wide and ovoid. The penial retractor muscle is

long and originates at the diaphragm, inserting 0.6 the

way to the base of the epiphallus. The penis is approxi-

mately the same length as the epiphallus and the same

diameter as the apical 0.7 of the epiphallus. The penis,

narrow apically, widens slightly below the apex, and

tapers basally to 0.2 the diame of the atrium where

they join. The atrium, widest at the insertion of the penis,

narrows abruptly by 0.6 at the gonopore. The vagina

narrows slightly at the junction with the free oviduct and

spermathecal duct. The S-sh iaped free oviduct is 1.2

times the width of the spermathecal duct at their junc-

tion with the vagina. The diameter of the spermathecal

sea doubles from its junction with the vagina distally to

0.4 the length, then narrows to 0.4 its greatest diameter

at 0.7 the length, and remains narrow to the junction

with the ovate spermatheca.

The central teeth of the radula (Figure 6, second from

the right) are tricuspid, 7-S zm wide and 11-12 pm

long, of similar shape and length but slightly narrower

and shorter than the first laterals, 8-9 fxm wide and 12—

13 pm long (Figure 6). The mesocones of both the cen-

trals and first laterals are tall and slender, and project

beyond their basal plates. The ectocones of the central

teeth are trigonal and symmetric. Ectocones and en-

docones of the laterals are trigonal and about 0.5 the

height of the mesocones. The endocones of the laterals

are slightly larger but otherwise of similar shape to their

ectocones. The first 5 teeth to the left and right of the

central row are similar to the first lateral teeth, the next

3 teeth on either side grade in shape and are difficult to

classify as either lateral or marginal teeth. The last 4

marginal teeth are wider, 10-11 pm wide, and shorter

). The apical 0.7 of

7-S pm long (Figure 7). Both the endocones and ecto-

cones of the marginal teeth are irregularly multicuspid.

The endocones are 0.9-0.8 as tall as the mesocones

and usually bear 3 cusps, although these are sometimes

divided into additional cusps. The ectocones sit on a

rectangular base and are divided into 5 or more cusps

that are 0.7 to 0.8 the height of the mesocones.

Remarks: The only other Paryphantopsis species with

shells bearing periostracal processes are: P. abstrusa

Slapcinsky, 2005; P. fultoni (Coen, 1922); P. lebasii

Slapcinsky, 2005; P. lame llizera (Thiele, 1928): P. striata

(Fulton, 1902); P. yawii Slapeinsky, 2005; and P. yelensis

Slapcinsky, 2006. However, the periostracal processes in

P. corolla do not overlap unlike P. fultoni and P. yawii,

and are sharply pointed unlike P. lebasii. Also, the pro-

cesses are prominent unlike P. abstrusa and P. yelensis.

Finally, the shells of P. lamelligera and P. striata are

more than twice the size of P. corolla. Paryphantopsis

corolla has an epiphallus that is basally robust unlike all

other Paryphantopsis species for which the genital

anatomy is known e xcept for P. nucella and P. misimensis

Slapcinsky, 2006. In contrast to P. corolla, P. nucella has

an apically inflated penis in addition to a large dark glan-

dular area on the spermathecal duct, and P. misimensis

lacks an apical diverticulum. The ectocones of the mar-

ginal teeth sit on a rectangular extension of the tooth,

unlike all other Paryphantopsis tor which the radular

morphology is known other than P. fragilicosta and P.

nucella.

Etymology: The species name is from the Latin femi-

nine noun corolla and describes the shells resemblance

to a small garland or crown.

Paryphantopsis. aa new species

(Figures 8-13, Table 1

Holotype: UF 353995, J. Slapcinsky, 2S February

2005.

Paratypes: Papua New Guinea, East New Britain

Province, type locality, J. ae 28 February 2005,

BPBM 282461 (2 specimens), PNGNM (1 spe cimen),

UF 366505 (4 specimens), UF 366507 (9 specimens).

Type Locality: Papua New Guinea, East New Britain

Province, New Britain, Saukale, 13 km northwest of

Marmar Village on the trail to Pakia Village, 5.426° S,

151.453° E, 910 meters altitude (Figure 1).

Habitat: Collected in mixed hill forest on vegetation

within | meter of the ground.

Description: The adult shell is globose to depressed-

globose, average size for the genus, 4.5-5.3 mm (mean =

5.00, see Table 1 for sample size and standard deviation)

in diameter and 3.2-3.7 mm (mean = 3.43) in height,

with 2.9-3.1 (mean = 3.00) rapidly expanding whorls

(Figures 8-10). The suture is deeply impressed and the

shell periphery is evenly rounded. The spire is slightly

elevated, 0.1—-0.2 mm (mean = 0.13). Teleoconch whorls

descend slowly and regularly until the end of the body

whorl, which desce its. slightly more rapidly. The shell

Page 186 THE NAUTILUS, Vol. 121, No. 4

aes

Figures 8-13. Paryphantopsis fragilicosta. 8-10. Shell, Holotype UF 383995. Scale bar = 1 mm. 11. Genitalia, UF 366507.

Scale bar = | mm. 12-13. SEMs of radula, UF 366507. 12. Central and lateral teeth. 13. Marginal teeth, Scale bars = 10 jm.

J. Slapcinsky and R. Lasley, 2007

Page 187

height:diameter ratio is 0.65-0.71 (mean = 0.69). There

are eee 1.4 evenly rounded protoconch whorls

sculptured with 13 spiral rows of small pits which con-

tinue on the teleoconch where the pits elongate nearly

fusing to form discontinuous spiral striae. Short peri-

ostracal extensions that do not bear periostracal pro-

cesses are present approximately every four growth lines,

These extensions are often worn or absent in older adult

shells. The protoconch and teleoconch whorls are usually

uniformly brown, shiny, and translucent, although older

worn shells can be dull brown, opaque, with a white

protoconch. The umbilicus is closed or nearly closed by

a reflection of the peristome. The aperture is large and

ovate, with an aperture:width to aperture:height ratio of

1.0S—1.28 (mean = 1.18).

The external body color is bright yellow in life and

there are no bands or other color patterns. Specimens

preserved in ethanol fade to uniform cream. The head is

short similar to other species of Paryphantopsis and the

posterior of the foot is of average length for the genus.

The vas deferens is 0.2 the diameter of the slightly

inflated head of the epiphallus which does not bear a

diverticulum (Figure 11). The epiphallus is approxi-

mately 1.5 the length and 0.5 the diameter of the penis

and widens only slightly at the junction with the penis.

The penial retractor muscle is moderate in length, origi-

nating from the diaphragm and inserting at the basal 0.3

of the epiphallus. The penis is robust and of uniform

width apically, narrowing basally to 1.2 the width of the

atrium just before their junction. The atrium narrows

slightly and then broadens between the vagina and gono-

pore. The vagina broadens slightly at its junction with the

free oviduct and spermathecal duct. The straight free

oviduct is 0.5 the width of the spermathecal duct at their

junction with the vagina. The sper mathecal duct is rela-

tively wide basally, tapering to 0.3 its basal diameter at its

midpoint, and remaining narrow until the junction with

the ovate spermatheca.

The central teeth of the radula (Figure 12, middle

row) are — S-9 pm wide, and 12-13 pm long, of

similar shape and length but slightly narrower and

shorter than the first lateral teeth, 10-11 xm wide and

14-15 pm long (Figure 12). The mesocones of the cen-

tral and first later sal teeth are tall, slender, and project

slightly beyond their basal plates. The ectocones of the

central teeth are trigonal and symmetric. Ectocones and

endocones of the lateral teeth are trigonal and about 0.5

the height of the mesocones. The endocones of the lat-

eral teeth are slightly larger but otherwise of similar

shape to their ectocones. The first 10 teeth to the left and

right of the central row are similar to the first laterals.

The next 3 on either side grade in shape and are difficult

to classify as either lateral or marginal teeth. The last 4

marginal teeth are slightly wider, alot 11-12 pm wide,

and shorter, 7-10 ym long (Figure 13). The base of the

unicuspid or bicuspid endocones of the first and second

marginal teeth is reduced, and they originate from the

side of their mesocones. The ectocones are reduced, and

their rectangular bases are usually devoid of cusps. The

third and fourth marginal teeth often lack all cusps and

are reduced to rectangular bases.

Remarks: ~The only other Paryphantopsis species with

shells that have periostracal extensions on the growth

lines and no processes at the margin are: P. arcuata Jut-

ting, 1964; P. dauloensis Solem, 1970; P. filosa Jutting,

1964: P. koragae Slapcinsky, 2005; P. latior Jutting, 1964;

P. matawanensis Slapcinsky, 2005; P. platycephala Jut-

ting, 1964; and P. pygmaea (Bavay, 1908). Paryphantop-

sis fragilicosta is larger than P. filosa, is less depressed

than P. dauloensis, has a higher spire than P. latior and P.

platycephala, and has stronger spiral sculpture than P.

pygiaca and P. arcuata. Paryphantopsis fragilicosta

lacks an apical diverticulum, unlike all Paryphantopsis

species for which the genital anatomy is known, except

for P. louisiadarum, P. misimensis, and P. vanatinensis,

which are much larger, as well as P. lebasii and P. yawii,

which have long periostracal processes. The ectocones of

the marginal teeth of P. fragilicosta sit on a rectangular

extension of the tooth, unlike all other Paryphantopsis

for which the radular morphology is known other than P.

corolla and P. nucella. It differs from these species by

having very few or no cusps on the marginal teeth.

Etymology: The species name derives from the com-

bination of the Latin adjective fragilis meaning crackling

or easily a and the feminine noun costa meaning

rib, and refers to the shells fragile periostracal exten-

sions.

Paryphantopsis nucella new species

(Figures 14-19, Table 1)

Holotype: UF 3583996, J. Slapcinsky, 258 February

2005.

Paratypes: Papua New Guinea, East New Britain

Province, New Britain: BPBM 282462 (2 specimens),

MNHW MP LOOL (2 ease PNGNM (2 speci-

mens), SME 329401 (2 2 specime ns) UF 383993 (54 speci-

mens), UF 383994 (3 specimens), type locality, J.

clea 28 February, 2005; UF 366504 (5 speci-

mens), UF 366506 (7 specimens), 12 km northwest of

Marmar Village on the trail to Pakia Village, 5.432° S,

151.460° E, 900 meters altitude, jes lapcinsky, 25 Feb-

ruary 2005

Type Loc: ality: Papua New Guinea, East New Britain

Province, New Britain, Saukale, 13 km northwest of

Marmar Village on trail to Pakia Village, 5.426° S,

151.453° E, 910 meters altitude (Figure 1).

Habitat: Collected in mixed hill forest on vegetation

within 1 meter of the ground.

Description: The adult shell is globose to depressed-

globose, average for the genus, 4.9-5.7 mm (mean =

5.22, see Table | for sample size and standard deviation)

in diameter and 3.1—-4.1 mm (mean = 3.75) in height

with 2.6-2.9 (mean = 2.81) rapidly expanding whorls

(Figures 14-16).

The apical surface of the shell whorls

Page 185 THE NAUTILUS, Vol. 121, No. 4

Figures 14-19. Paryphantopsis nucella. 14-16. Shell, Holotype UF 353996, Scale bar = 1 mm. 17. Genitalia, UF 383993

Scale bar | mm. 18-19. SEMS of radula, UF 383993. 18. Central and lateral teeth. 19. Marginal teeth. Scale bars = 10 wm

J. Slapeinsky and R. Lasley, 2007

Page 189

are flattened between the deeply impressed sutures

and the submedian shell periphery, which gives the shell

a somewhat angular appearance. The spire is slightly

elevated, 0.1-0.2 mm (mean = 0.18). The teleoconch

whorls descend slowly and regularly until the end of the

body whorl and more rapidly near the aperture. Shell

height: diameter ratio is 0.62—0.78 (mean = 0.72). There

are 1.5 evenly rounded protoconch whorls sculptured

with spiral rows of small pits. Approximately 14 rows can

be seen on the apex of shells. These pits become elon-

gated on the teleoconch whorls nearly fusing to form

weak incised spiral striae. Spiral striae w veaken, becoming

obsolete on the final 0.30 of the body whorl where shell

sculpture becomes predominated by growth lines all of

which are accentuated with short periostracal extensions

that do not bear peripheral processes. The protoconch is

white, and teleoconch whorls gradually darken from yel-

low brown to brown. The umbilicus is closed bya reflec-

tion of the peristome. The aperture is ovate except for

the flattened apical surface and has an aperture:width to

aperture:height ratio of 0.94-1.25 (mean = 1.14).

The external body color is bright yellow in life and

there are no bands or other color patterns. Specimens

preserved in ethanol fade to uniform cream. The head is

short similar to other species of Paryphantopsis and the

posterior of the foot is slightly shorter than average for

the genus.

The vas deferens is 0.2 the diameter of the head of the

epiphallus, which bears an apical diverticulum that is

approximately 0.3 the length and 0.7 the diameter of the

midpoint of the epiphallus (Figure 17). The apical 0.7 of

the epiphallus is uniform in diameter: the basal 0.3 is

ovoid and twice as wide. The penial retractor muscle is

moderate in length originating from the diaphragm and

inserting 0.6 the way to the base of the epiphallus. The

penis is 0.7 the length of the epiphallus. Its apical 0.3 is

as robust as the base of the epiphallus and narrows rap-

idly basally. The basal 0.7 of the penis is 0.3 narrower

than the apex and uniform in diameter to the junction

with the atrium. The atrium is 1.5 times wider than the

penis at their junction and remains constant in diameter

to the gonopore. The vagina expands slightly at the junc-

tion with the free oviduct and spermathecal duct. The

free oviduct is broad, not folded, and approximately the

same diameter as the base of the spermathecal duct. The

basal 0.3 of the spermathecal duct is wide and sur-

rounded by darkly pigmented glandular tissue. The basal

0.5 of the spermathecal duct tapers apically to 0.2 of its

original diameter and remains narrow until the junction

mali the ovate spermatheca.

The central teeth of the radula (Figure 18, middle

row) are tricuspid, 10-11 zm wide and 14-15 pm long,

and of similar shape and length, but are slightly narrower

and shorter than the first lateral teeth, 11-12 wm wide

and 16-17 pm long (Figure 18). The mesocones of the

central and first lateral teeth are tall and slender, and

project beyond their basal plates. The ectocones of the

central teeth are trigonal and symmetrical. Ectocones

and endocones of the lateral teeth are trigonal and about

0.5 the height of the mesocones. The endocones of the

lateral teeth are slightly larger but otherwise of similar

shape to their ectocones. The first 12 teeth on either side

of the central row are similar to the first laterals. The next

three on either side grade in shape and are difficult to

classify as either iaeieale or marginals. The last four teeth

are clearly marginals and are irregularly multicuspid,

wider, about 13-14 pm wide, and shorter, 8—9 wm long

(Figure 19). The endocones are as tall as the mesocones

and are usually unicuspid, but sometimes bear 2 or 3

cusps. The ectocones sit on a rectangular base and are

usually divided into 2-6 cusps that are 0.6 to 0.7 the

height of the mesocones.

Remarks: = Paryphantopsis nucella is similar to only P.

koragae in having short periostracal extensions on all

growth lines; ese growth lines do not bear processes at

the shell periphery. The shell of P. koragae, however,

differs in having the shell periphery above, rather than

below, the midpoint of the whorl. Paryphantopsis nucella

is unique among Paryphantopsis, for which the genital

anatomy is known, in having a large glandular area on the

base of the spermatheca and havi ing both an apically

robust penis and a basally robust epiphallus. The ecto-

cones of the marginal teeth sit on a rectangular extension

of the tooth, unlike all other Paryphantopsis for which

the radular morphology is known other than P. corolla

and P. fragilicosta.

Etymology: The species name derives from the Latin

feminine noun nucella and describes the shells resem-

blance to a small nut.

Discussion and Conclusions: Despite previous in-

tensive surveys for terrestrial snails in coastal New

Britain, much of the diversity of the island’s interior

mountains may remain to be described. More explora-

tion is clearly needed in the Nakanai Mountains, where

uplifted karst terrain has promoted the development of a

unique and diverse snail fauna. Our cursory surveys,

which did not penetrate deeply into, or reach high el-

evations in the Nakanai Mountains, uncovered several

previously unreported species, including the three Pary-

phantopsis species described here. Most species of Pary-

phantopsis appear to be restricted to single mountain

ranges or islands. The absence of these three species

from relatively intensively surveyed coastal areas suggest

they are endemic to the Nakanai Mountains. It is likely

that other endemic species will be found in the W hite-

man, Baining, and Willaumez ranges of New Britain. The

Charopidae were until recently, considered a minor com-

ponent of the terrestrial mollusk fauna of New Guinea

(Solem, 1983). Our surveys (Slapcinsky, 2005; 2006) sug-

gest charopid species radiations on New Guinea and sur-

rounding islands rival the spectacular radiations exhib-

ited by this family in the oceanic Pacific (Solem, 1983).

Based on shell and genital morphology, the relation-

ships among the three Paryphantopsis species on New

Britain are unclear. Paryphantopsis corolla and P. nu-

Page 190

THE NAUTILUS, Vol. 121, No. 4

cella share several unusual traits that may indicate a com-

mon origin. In both species, the marginal ectocones are

divided into numerous irregular cusps, the base of the

epiphallus is robust, and an apical diverticulum is present

on the epiphallus. Paryphantopsis fragilicosta, on the

other hand, lacks these characters but has a large, glo-

bose shell that can be difficult to differentiate from that

of P. nucella. However, all three New Britain species

share unique rectangular bases to their marginal teeth, a

possible syne spomorphy. If so, characters that have been

historically used to determine monophyly, i.e. genital and

shell morphology, appear to be rapidly evolving and vari-

able in this group. Additional morphologic and genetic

characters are necessary before a clear picture of the

relationships between Paryphantopsis species within

New Britain and the adjacent New Guinea mainland can

be resolved.

ACKNOWLEDGMENTS

We thank the landowners of Marmar and Pomio for per-

mission to work on their land and for field assistance:

Anton Buntapeko, Fred Kraus, Esrum Lonpulpagetuna

and Damian Matalo for additional field assistance; Gai-

kovina Kula and Maureen Ewai of Conservation Inter-

national for providing logistical support and advice;

Florence Paisparea, Environmental Officer for East New

Britain Province for facilitating our visit to Marmar;

Papua New Guinea National Museum and Art Gallery

for providing in-country collaborative assistance; Papua

New Guinea Department of Environment and Conser-

vation, Papua New Guinea National Research Institute,

and East New Britain Provincial Government for per-

mission to work in East New Britain Province; Karen

Kelley, Electron Microscopy Core Laboratory, Univer-

sity of Florida for imaging radulae; G. Paulay for lending

photographic equipment; and Jack Worsfold for sharing

bibliographic information. Fieldwork for this ee

Was supporte od by National Science Foundation grant

DEB 0103794 and the Unive rsity of Florida Foundation,

McGinty Endowment.

LITERATURE CITED

Audra, P., S. E. Lauritzen, and P. Rochette. 2001. L’hyperkarst

des montagnes Nakanai. Modéle d’évolution dun réseau

juvénile (gouffre Muruk) basé sur des datations U/Th et

paléomagnétiques des sediments. 93-99 in Audra, P.,

P. DeConinck, and J.-P. Sounier, (eds.) Nakanai 197S—

1998: 20 ans d’exploration, Association Hémisphére Sud,

Antibes, 224 p

Bavay, A, 1908. Mollusques terrestres et fluviatiles. Nova

Sar Zoology 5: 269-292, pl. 14.

Beehler, M. (ed.) ) 1993. A biodiversity analysis for Papua

New Ge Papua New Guinea conservation needs as-

sessment, Vol. 2. Biodiversity Support Program, Washing-

ton, DC & Department of Environment and Conserva-

tion, Boroko, Papua New Guinea, 1-433 p.

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.

Flannery, T. F. 1995. Mammals of New Guinea. Cornell Uni-

versity Press, Ithaca, 568 pp.

Fulton, H. C. 1902. Descriptions of new species of land Mol-

lusca from New Guinea. Annals and Magazine of Natural

History 7(9): 182-154.

Jutting, W. S. S. v. B. 1964. Non-marine Mollusca of West New

Guinea. Part i Pulmonata, I. Nova Guinea, Zoology 26:

1-74, pls. 1-2

McAlpine, 1 R. fe Fryne, D, E. 2001. Land use change and

intensification in Papua New Guinea 1975-1996. Asia Pa-

cific Viewpoint 42(2/3): 209-215.

Orenstein, R. I. 1976. Birds Ple esyumi Area, Central New

Britain. The Condor, Vol. 78(3): 370-374.

Paijmans, kK. 1976 Vegetation. ms Pajminas K. (ed.) New

Guinea Vegetation, Australia National University Press,

paper 212 pp.

1934, Systematische und tiergeographische Unter-

ce iiber die Landschneckenfauna des Bismarck-

Archipels. I. Archiv fiir Naturgeschichte 3(3): 445-488.

Rensch, I, 1937. Systematische und ey ee Unter-

suchungen iiber die Landschneckenfauna a Bismarck-

Archipe els. IT. Archiv fiir Naturgeschichte 6(3); 526-644.

Slapcinsky, J. 2005. Six new species of ere (Gas-

tropoda: Pulmonata: Charopidae) from the Papuan Pen-

insula of New Guinea. The Nautilus 119: 27-42.

Slapcinsky, ]. 2006. Paryphantopsis (Gastropoda: Pulmonata:

Charopidae) from the Louisiade Archipelago of New

Guinea. The Nautilus 120; 119-130.

Solem, A. G, 1959. On the family position of some Palau, New

Guinea, and Queensland Jand snails. Archiv fiir Mol-

luskenkunde $8: 151-155 pls. 12, 13, 2 figs.

Solem, A. G. 1970. The Enc eee 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 H, Fami-

lies Punctidae and Charopidae, Zoogeography. Field

Museum of Natural History, Chicago, ix + 336 pp.

Thiele, J. 1925. Mollusken vom Bismark-Archipel, von Neu

Guinea und Nachbar-Inseln. Zoologische Jahrbiicher 55:

119-146.

Woodhead, J. D.. S. M. Eggins, and R. W. Johnson. 1998.

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Rensch,

THE NAUTILUS 121(4):191-200, 2007

Page 191

Mysella gregaria new species, a bivalve (Galeommatoidea:

Montacutidae) commensal with an intertidal burrowing sea

anemone from North Carolina, USA

Lennie Rotvit Thomas Fox

Jorgen Littzen

Ase Jespersen!

Department of Cell Biology and

Comparative Zoology

Institute of Biology

University of Copenhagen

edie rsitetsparken 15

Dk- 2100 Copenhagen 0, DENMARK

North Georgia College and State University

Dahlonega, GA 30957 USA

ABSTRACT

Mysella gregaria new species (Bivalvia: Galeommatoidea: Mon-

tacutidae) is described from Wrightsville Beach, North Caro-

lina, USA. Several individuals were collected from the body

column of an intertidal burrowing unidentified actinian. The

morphology of the shell and soft parts is described and com-

pared with other species of the genus from the W. Atlantic

Ocean and with other bivalves associated with solitary antho-

zoans. Mysella gregaria is a ctenidial brooder and specimens

are either males or females; no true hermaphrodites were

found. Contrary to many less social commensal bivalves, repro-

duction in M. gregaria does not involve sperm storage. We

suggest that this may be correlated with the species’ gregari-

ousness.

Additional Keywords: Mollusca, reproduction, anatomy, sperm

morphology, gregariousness.

INTRODUCTION

During intertidal collecting on a muddy sand flat at

Wrightsville Beach, North Carolina, USA, one of us (TF)

collected a number of galeommatoidean bivalves from

S—10 cm long specimens of an unidentified burrowing

actinian. The host species was rare, as not more than an

estimated 5-S specimens were found during occasional

visits to the locality in the period from October 1970

through July 1975. Unfortunately, none of them were

preserved for later identification. Only two of the col-

lected actinians had commensal bivalves attached.

Many bivalves of the superfamily Galeommatoidea

have a commensal life style as they live together with

species of bottom- dwelling marine invertebrate hosts

such as other bivalves, polychaete s, sipunculans, ech-

' Corresponding author, [email protected]

iurans, crustaceans, and echinoderms. Commensal asso-

ciations between bivalves and solitary anthozoans are re-

stricted to three known cases (Yamamoto and Habe,

1961; Ponder, 1971; Oliver, 1993) and it was therefore

judged to be of interest to examine the present associa-

tion more closely. The study has disclosed that the bi-

valves represent a new species of Mysella Angas, 1877, a

genus that comprises both commensal and free- living

mivalwes es and species that are commensal on certain con-

ditions, free-living on other conditions (Ockelmann and

Muus, 1978).

MATERIALS AND METHODS

Thirty-two bivalves were retrieved from the skin of a host

specimen collected on 9 July 1975. The site of collection

was a sandy mud tidal flat in Banks Channel, Wrightsville

Beach, North Carolina, and the approximate coordinates

are longitude 77.8° W and latitude 34.2° N. The anemo-

nes were dug from the lower intertidal zone during a

spring low tide. The number associated with the second

host and the date of collection was not noted. The be-

havior of the bivalves after removal from the host was not

studied, but it was observed that they detached easily.

Seventeen of the bivalves were preserved in Heiden-

hain‘s Susa for several hours. This procedure decalcifies

the shells and the sizes given based on measurements of

the mantle are therefore approximate. Three other bi-

valves were fixed in 70% ethanol and used for the de-

scription of the shell and for type material. Six specimens

were embedded in Araldite and cut into 2-jzm thick se-

rial sections that were stained with toludine blue. Seven

other specimens were embedded in Paraplast, sectioned,

and the 8-jm thick serial sections stained with hema-

toxylin and eosin (H+E). Ultrathin sections were per-

formed on the testis of one of the males. These sections

Page 192

THE NAUTILUS, Vol. 121, No. 4

were contrasted with uranyl acetate and lead citrate and

examined in a JEOL 1LOOSX electron microscope. Scan-

ning electron micrographs were made using a JEOL

JSM-6335F SEM. Photos of the type specimens of My-

sella casta (Verrill and Bush, 1897) (USNM 77632) and

M. barbadensis Dall, 1899 (USNM 95703) were used for

comparison with M. gregaria. Shell length (SL) and

height (SH) are given to the nearest 0.1 mm.

SYSTEMATICS

Family Montacutidae Clark, 1855

Genus Mysella Angas, 1877

Mysella gregaria new species

(Figures 1-20)

(Figures 1-10): Observations

The SL of twenty measured

Description: SHELL

were made on three shells.

specimens varied from 3.0 to 6.0 mm. The measure-

ments SLxSH in the type material are 4.5x3.6 mm,

5.1x3.8 mm, and 5.5x4.3 mm. The outline is almost per-

fectly oval, slightly higher in the anterior part, all margins

being evenly rounded. The valves are relatively flattened,

very thin, semitransparent, and with a light- brown. to

grey-brown periostracum, which is darkest in the dorsal

part. There are no coatings of ferruginous deposits. The

surface is smooth, with very fine commarginal lines and

even finer radial striae. No growth checks were ever

visible. The interior surfaces of the valves are polished.

The umbos are not very prominent and placed slightly

toward the posterior region. In the right valve there are

two diverging teeth; the anterior tooth is elongate-

subtriangular and more prominent than the narrower,

posterior one. They are separated by a stout ligament

placed immediately below umbo in a triangul: uv senliee

The left valve is edentulous but has a produced dorsal

margin that fits into the teeth of the right valve. The

anterior adductor scar is subtriangular, the posterior

more oval, both fused with the respective foot retractor

scars. The pallial line is relatively broad and lacks a pallial

sinus.

MANTLE (Figure 11): The mantle folds are fused far

behind and for a short distance to separate the mantle

opening into an inhalant-pedal aperture and an exhalant

aperture. While the first forms a long slit ma the four-

fifths of the length of the ventral side, the exhalant ap-

erture is very small and located far poste pai Since live

animals were not observed, we do not know whether, or

to which extent, the mid mantle fold may cover the out-

side of the shell or whether there are any siphons. The

mantle edges bear minute papillae that are most distinct

Figures 1-4.

Vysella gregaria. Cleaned shell of holotype l.

rom inside. 4. Left shell seen from outside. Shell length

Right shell seen from inside. 2. Right shell seen from outside. 3. Left

5.1. mm

Rotvit et al., 2007 Page 193

Figures 5, 6. Mysella gregaria. Paratype, outside view of left (5) and right sides (6). Shell length 4.5 mm.

in the dorso-anterior sector. A typical, ciliated rejection outline, whereas the smaller posterior adductor is more

fold is located just anterior to the end of the inhalant- oval. A small pedal protractor muscle is located ventrally

pedal opening. and clearly outside the anterior adductor muscle. The

MUsCULATURE (Figures 11, 13): The anterior adductor two pedal retractor muscles are equally large and of

muscle is the larger of the two and is subtriangular in moderate size. They terminate in the base of the foot,

Figures 7-10. Mysella gregaria. Holotype, SL.5.1 mm. Scanning electron micrographs of right hinge seen in direct view (7) and

slightly tilted from below (8) and of left hinge seen in direct view (9) and slightly tilted from below (10). Scale bar represent

200 ym

Page 194

THE NAUTILUS, Vol. 121, No. 4

Figures 11,12. Mysella gregaria. 11. Anatomy of male, right valve, mantle and gill removed, heart and pericardium omitted. 12.

Light microscopic representation of sperm cell. Abbreviations: aa, anterior adductor muscle; ae, acrosome; ar, anterior pedal

retractor muscle; bg, byssus gland; eg, cerebral ganglion; dg, digestive gland; f, flagellum; fg, foot groove; fu, fusion of left and right

demibranch behind visceral mass; ga, gill axis: in, intestine; k, kidney; Ip, labial palps: mp, middlepiece; n, nucleus; pa, posterior

adductor muscle: pg, pedal ganglion and statocysts; pp, protractor pedis muscle; pr, posterior pedal retractor muscle; so, sexual

opening: ss, style sac; st, stomach; te, testis; vg, visceral ganglion. Arrowheads indicate water entering along the inhalant-pedal

opening and exiting through the exhalant opening.

while only a few fibers spread further into the foot. There

are almost no muscle fibers serving the byssus gland.

FOOT AND Byssus GLAND (Figures 11, 14-16): In pre-

served specimens, the foot extends forward and a little

beyond the shell margin. It is bluntly rounded in front.

The ventral side is distinctly ciliated and has a furrow

that extends from in front of the heel to tip of the foot.

In some specimens a blackish pigment spot can be seen

at the very tip of the foot. The very small byssus cavity

appears Y-shaped in transversal sections as it consist of a

right and left slit which both discharge into the median

furrow by way of a single duct. The lining epithelium is

composed of a mixture of ciliated and of mMUCOUS-pro-

ducing goblet cells. A group of more distant glandular

cells terminate between the cells of the byssus cavity

epithelium through long, slender ducts, Although such

cells are normally involved in the formation of byssal

threads, none were ever seen either in sections or on

vhole animals. The nature of the glandular epithelium

indicates that slime threads, rather than normal byssal

threads are produced by the byssus gland.

GILLs (Figures 11, 17): The gill axis, which runs from

near the umbo, has an oblique course backward. Each

gill is triangular. An outer demibranch is entirely absent

and only the inner one is present. The ascending lamel-

lae of the left and right demibranch are fused behind the

foot, more anteriorly to the lateral sides of the visceral

mass. Both gills are fused to the mantle edges immedi-

ately in front of the exhalant opening. A food groove is

only present along the ventral edges of each gill. Inter-

lamellar junctions are present, but they are very few,

whereas interfilamentary bridges are numerous and oc-

cur regularly, Right and left hypobranchial glands of nor-

mal size occur well inside the exhalant aperture. A pair of

relatively large labial palps lies on either side of the

mouth, Their opposing surfaces are provided with 9-10

ciliated ridges.

ALIMENTARY CANAL (Figure 11): The esophagus is a

Rotvit et al., 2007 Page 195

Figures 13-16. Mysella gregaria. 13. Sagittal section of a male (SL 3.0 mm) showing the course of the protractor pedis muscle

(pp). 14. Transverse section of a male (SL 4.3 mm) through the foot and the opening of the byssus cavity (be). 15. Byssus gland cells

(bg) opening into byssus cavity 16. Transverse section through byssus cavity of the male shown in Fig. 14. Abbreviations: aa, anterior

adductor muscle; ar, anterior pedal retractor muscle; be, byssus cavity; bd, byssus duct; bg, byssus glandular cells; eg, cerebral

ganglion: ep, epithelium of byssus cavity: fg, foot groove: me, mucous cells. 2-4m thick Araldite sections stained with toluidine blue

(13, 14 and 16) and $-ym thick paraplast sections stained with H+E (15). Scale bars represent 100 zm (13-15) and 30 zm (16).

Page 196 THE NAUTILUS, Vol. 121, No. 4

2 eK a

Deena rein OF POY hae a

Ota a

pr Qo ag

[Yee a es Sad

Figures 17-20. Mysclla gregaria. Transversal section of male (SL 4.3 mm) (17) and of female (18, 19), 20. Testis with abortive

oocyte (oo). Abbreviations: au, auricles; id, inner demibranch; in, intestine; k, kidney; ne, nerve; np, nephridiopore; ov, posterior

wall of ovary; pe, pericardial cavity; pr, posterior pedal retractor muscle; rp, renopericardial canal; so, sexual opening; te, testis. 2-zm

thick Araldite sections stained with toluidine blue. Scale bars represent 200 wim (17), LOOpm (18, 19) and 50 zm (20).

Rotvit et al., 2007

Page 197

short curved tube. The stomach is relatively capacious

and heavily cuticularized. The style sac forms a wide

posterior continuation from the stomach, is elongated

conical, nearly as long as the stomach, and directe d pos-

tero- ventrally. It is placed within the right side of the

visceral mass. The intestine leaves from the underside of

the stomach near its connection to the stvle sac, runs

alongside the style sac and forms a loop around its tip,

then sie dorsally between left and right parts of the

gonad to loop backwards to the rectum The bound: wy

between the ciliated intestine and the unciliated rectum

is marked by a sharp transition zone dorsal to the poste-

rior adductor.

The digestive gland is mainly located ventral to the

stomach and around the posterior part of the esophagus.

In addition to a large ventral communication with the

stomach, there are also smaller openings into it.

REPRODUCTIVE SysTEM (Figures 11, 17, 1S, 20): Eight

sectioned bivalves (SL = 3.0-5.6 mm) were males, wile

three other sectioned bivalves (SL = 4.1-6.0 mm) were

females. No truly hermaphroditic specimens were found,

but a few mature oocytes were present in the testis of

one of the males (Figure 20).

All males were sexually mature. The very large testis

occupies the posterior half of the visceral mass being

replaced more anteriorly by the style sac, stomach, ond

digestive gland. The ge meral she ape as seen in decalcified

specimens is relative ly constant. A large undivided pos-

terior portion gives rise to right and left halves that sur-

round the intestine and extend ventrally to send a few

short branches forward. Two other forwardly directed

finger-shaped branches embrace the stomach. The

paired aaa issue from the most posterior, undi-

vided part of the testis. They are short but have promi-

nent funnel-shaped and heav ‘ly ciliated openings into the

suprabranchial chamber. Spermatogenesis proceeds ev-

erywhere along the walls of the different portions of the

testis. There is no special chamber for storing the mature

sperm which, in a non-orientated manner accumulate in

huge numbers in the central parts of the different por-

tions of the testis. Many spherical to ovoid cells, 5-7 ym

in diameter, lie scattered among the mature sperm.

These cells have a centrally place a nucleus plus one or

two inclusions. We were unable to discover how these

cells arise and if they are in the any way associated with

the spermatogenesis.

Because of the insufficient fixation the TEM micro-

graphs were of low quality. They nevertheless show

enough details to illustrate the general ultrastructure of

the flagellate sperm cells (Figure 11, B). The acrosome is

1.4-1.5 pm long and basally (near the nucleus) 0.6 xm

broad. It is divided into a terminal subspherical body

(0.4x0.6 2m) and a slightly tapering acrosomal vesicle,

which is basally deeply invaginated to accommodate a

filamentous subacrosomal material. The nucleus is 0.7—

0.S xm across and 1.2—1.3 zm in length. The 1.4-1.5 wm

long middle piece was extremely ill- -preserved without

any identifiable mitochondria but seems from a broad

base near the nucleus to taper backwards.

The female sexual openings (Figure 18) are identical

in size and structure to those of the male. Two females

were obviously spent, but one of them had retained a few

abortive oocytes (diameter ca. 60 jzm) within the ovary

and a few embryos in the suprabranchial chamber. Ex-

cept for this, none of six decalcified but unsexed speci-

mens examined in transparent light (SL = 4.0-5.6 mm)

were brooding ova or larvae. No structures for storing

foreign sperm were found in any of the sectioned ts

valves.

EXCRETORY SYSTEM (Figures 11, 18, 19): Left and right

halves of the kidney communicate in the median plane

for a short distance. Each one is composed of several

smaller and larger sacs with glandular walls. Antero-

laterally the kidney opens directly to the suprabrancial

chamber through two ciliated nephroducts, which are

unchez wracteristically short and have porous nephrid-

eae placed not ‘far from the genital opening. Rather

far behind within the kidney, two long straight and

heavily ciliated renopericardial canals run Ponand to

open into the forward and ventral part of the pericar-

dium.

Host ReLations: All 32 individuals were attached

onto the body column of a single sea anemone, whereas

the number attached to a second host was not noted.

None were found on any of the other host specimens

from the same site, which suggests that the species is

gregarious. It was observed that the attachment to the

Rost was very loose as several of the bivalves had fallen

off during the collection. This corresponds with the con-

clusion that no true byssus threads are produced and that

attachment rather takes place by means of slime threads.

Holotype: BIV-445 (Zoological Museum, University

of Copenhagen), a cleaned shell, SL = 5.1 mm.

Paratypes: BIV-446, a shelled animal, SL = 4.5 mm;

USNM _ 1107828 (National Museum of Natural History,

Washington, DC), a shelled animal, SL = 5.5 mm; all

from type locality, 9 July 1975.

y: Banks Channel, Wrightsville Beach,

North Carolina, USA. (ea. 77.8° W, 34.2° N), lower in-

tertidal zone during spring low tide, sandy mud substra-

tum. 9 July 1975. Thirty-two specimens were attached to

a single host.

Type Locality:

Etymology: the species name is derived from the

Latin grex, flock, and refers to the gregarious life style of

the new species.

DISCUSSION

Identification: The details of the shell and hinge al-

locate the species into the genus Mysella (family Mon-

tacutidae) and indicate that it is close to the North At-

lantic M. bidentata (Montagu, 1803). In M. gregaria the

protractor pedis muscle is inserted immediately ventral

to the anterior adductor as in M. bidentata (Montagu,

1803) and in Montacutona compacta (Gould, 1S61), in

contrast to the condition in another group of mon-

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THE NAUTILUS, Vol. 121, No. 4

tacutids, in which it splits the adductor in dorsal and

ventral portions (Jespersen et al., in press). Mysella has

often been combined with Rochefortia Velain, 1877, but

the two differ with respect to the dimensions of the teeth

of the right valve (Coan et al., 2000; Holmes et al., 2006).

In species of Mysella the right valve has a stout trans-

verse tooth anterior to the resilifer and a posterior tooth

is either small or absent. In Rochefortia the right valve

has two subequal diverging teeth, which smenuld place

M. gregaria in Roc hefortia. The distinction ignores the

av ailable soft anatomical characters which, especially in

species with a Rochefortia type of hinge, vary consider-

ably: A small outer demibranch is present in some (M.

tumida (Carpenter, 1864), M. verrilli (Dall, 1899), and

M. sovaliki MeGinitie 1959), absent in other (M. biden-

tata and M. gregaria). The esophagus is a simple tube in

all the species except for M. verrilli, in which it forms a

suctorial proboscis. Foreign sperm are either not stored

(M. gregaria and M. verrilli) ), attached directly to the gill

surface (M. tumida), or stored in an unpaired pouch-

shaped seminal receptacle within the visceral mass (M.

bidentata) or in paired receptacles in the outer demi-

branch (M. sovaliki). A new definition of the genus My-

sella will probably be called for as additional data on the

anatomy of a wider range of species will bring about a

complete rearrangement of the genus and its included

species.

From the described species of Mysella from the East

American waters, M. gregaria may be distinguished by

the following chinacters: M. planulata (Stimpson, 1857)

has a very prominent hinge and the umbo is placed more

poste sriorly, It attaches to buoys and wharf pilings or oc-

curs in muddy sand of the Zostera community (Abbott,

1974: Franz, 1973). In Mysella ovata (Jeffreys, 1881) the

umbo is extremely protruding and - hoa far posteriorly.

It occurs in de spths of 183 to 287 m. The shell of M.

triquetra (Verrill and Bush, 1898) is ‘eualatee| and the

poste ror part of the shell is distinctly rostrate, not ey venly

rounded. Mysella verrilli, a deep water species, has the

umbo placed far posteriorly and the esophagus is devel-

oped as a suctorial proboscis (Allen, 2000). In M. stria-

tula (Verrill and Bush, 1898), both teeth are delicate and

very narrow, the anterior tooth is shorter, and the very

small umbo is located more posteriorly than in M. gre-

garia. The shell of the following two species were stuchier |

from photos provided by the National Museum of Natu-

ral History, Smithsonian Institution (USNM): The ante-

rior part of a left valve of M. barbadensis Dall, 1899

(USNM 95703) is almost three times as long as the pos-

terior part (in contrast to the claim that it is shorter as

described by Dall, who obviously mistook a left valve for

a right one). The left valve of the type of M. casta Verrill

and Bush, 1898 from North Carolina (USNM 77632) is

more elongate (SH:SL = 7:10) than the valves of M.

gregaria (SH:SL = 7.8:10) and the anterior end relatively

longer. Nevertheless, among the

the shell of

gregaria.

ast American species,

casta is the one most similar to M

Comparison with North Atlantic/Arctic species of

Mysella: Conchologically M. gregaria resembles M.

bidentata (Montagu, 1803) but differs anatomically in

that the latter has an unpaired seminal receptacle and

dimorphic sperm (Jespersen and Liitzen, 2001). The

shell of M. cuneata (Verrill and Bush, 1898) is distinctly

asymmetrical, as the right valve shows a slight concavity

with a consequent ait waren along the ventral margin

(Gage, 1968). In M. tumidula ( (Jeffreys, 1866), the pos-

terior shell margin is distinctly angular, not evenly

rounded. Mysella moelleri (Morch, 187" 7) and M. sovaliki

both differ from M. gregaria in the hinge structure and,

more significantly, in having preserv ed a small outer

demibranch (Petersen and Liitzen, in press). Mysella

planata (Dall in Krause, 1885) has a thick shell, in which

the left valve has the dorsal line modified into two teeth,

and the right valve shows only one, anterior tooth (van

Aartsen, 1996).

Comparison with Other Bivalves Associated with

Anthozoans: Commensalism between bivalves and

anthozoans is rare. Nipponomontacuta actinariophila

Yamamoto and Habe, 1961, is small Japanese bivalve that

has been found attached immediately outside the ring of

tentacles of Halcampella maxima Hertwig (Actiniaria:

Halcampoididae). Details of the relationship are not

known, except that three specimens sitting close together

ren the ring of tentacles were ‘lhuatrated by "Hobe

(1973). ). Although the outline of the shell and the position

of the umbo in N. actinariophila are somewhat similar to

the studied species, the hinges are clearly different, as

teeth are only present in the left valve in N. actiniari-

ophila, not, as in M. gregaria, on the right valve. Mon-

tacutona ceriantha Ponder 1971, from "Cenanties sp.

(Ceriantharia) in Moreton Bay, E Australia, is easily

separated from M. gregaria in ‘that each valve has four

cardinal teeth and a small outer demibranch is present.

Furthermore, M. ceriantha is attached to the interior of

the tube of the host, not to the body, and in a small

numbers (Table 1) (Ponder, 1971). A third bivalve, Hal-

campicola tenacis Oliver, 1993, from Rottnest Island off

Perth, SW Australia has a hinge similar to Montacuta, the

right valve with a strong anterior cardinal merging with a

ee ginal lateral ridge and an obsolete posterior car-

dinal. The left valve has anterior and posterior strongly

projecting marginal extensions which fit into the right

valve (Oliver, 1993). Besides, the ligament has a iitho-

desma and there is a small outer demibranch. A single

bivalve was found on each of six hosts (Halcampoides sp..,

Actiniaria: Halcampoididae) among 20 collected.

seems fairly obvious that all four anthozoan-associated

bivalve species are not specifically interrelated |

Reproduction: Eight of the sectioned bivalves (SL =

3.05.6 mm) were sexually mature males and three other

(SL = 4.1-6.0) were females. The females were spent but

one had retained a few embryos in the suprabranchial

chamber which shows that this species, like other gale-

ommiatoideans, is a ctenidial brooder. A few mature 0o0-

cytes, left over from a previous ovulation, were present in

Rotvit et al., 2007

Page 199

Table 1. Mean number of commensal bivalves per host specimen and reproductive specializations in montacutid bivalves. Abbre-

viations: dw, dwart males; sp, spermatophores; sr, seminal receptables; ss, sperm sacs; tt, testis transplantation.

Mean no. per

Bivalve species host specimen

Host species

Repre rch ictive

specializations References

Montacuta percompressa ] Holothurioidea tt Fox et al., 2007

Peregrinamor ohshimai 1 Crustacea sr, dw Liitzen et al., 2001a

Salpocola philippinensis ] Sipuncula tt? Liitzen et al., in press

Halcampicola tenacis ] Anthozoa > pers. comm. (Graham Oliver)

Litigiella pacifica 1-3 Sipuncula sr Liitzen and Kosuge, 2006

Montacutona ceriantha LS Anthozoa sr Ponder, 1971

Entovalva lessonothuriae <2 Holothurioidea sp Kato, 1998; Liitzen et al., 2005

Pythina arcuata ca. 2 Brachiopoda sr pers. comm. (JL)

Anisodevonia ohshimai 1.8-2.7 Holoturioidea sp Kosuge, 2001; Liitzen et al., 2005

Nipponomysella subtruncata 2.5 Sipuncula sr, SS Liitzen et al., 2001b

Tellimya ferruginosa <3 Echinoidea tt ob et al., 2007

Mioerycina coarctata 3.8 Sipuncula sr Gage, 1979

Mysella cuneata 5 Sipuncula sr Gage, 1968

one of the males, which could indicate that females may

change sex and that the species shows her maphroditic

tendencies. The species in all probability reproduces by

outcrossing, since none of the sectioned bivalves showed

truly simultaneous hermaphroditic characteristics.

All galeommatoideans brood the ova in a ctenidial

brooding chamber consisting of the inner and, if present,

the outer demibranch. This is also where fertilization

takes place. In a normally-filtering bivalve the gills and

the currents they generate do not favor a casual jaiake of

sperm suspended i in the water. The ciliary activity of the

gills probably functions as a barrier for penetration of

most sperm cells and the only other access to the brood-

ing chamber is against the flow of filtered water expelled

through the exhalant aperture. Many an have over-

come these difficulties by bulk transfer of sperm in con-

tainers of different nature to the female’s suprabranchial

chamber (see O Foighil, 1985a, for a review). Exactly

how they do this is not understood except in a single case

(O Foighil, 1955b). Some species have minute dwarf

males that are permanently and intimately associated

with the much larger female and still other produce

sperm of two types th at form spermatozeugmata. Some

of that spermatozeugmata is probably capable of inde-

pendent mobility. None of these methods of a precise

transfer of sperm occur in M. gregaria. The testis is ex-

ceptionally large in the species, mu we speculate that the

resulting high production of sperm cells may compensate

for the inevitable loss suffered during the transfer be-

tween the opposite sexes of sperm cells that are simply

broadcast into the water.

Sperm transferred to a female (or hermaphrodite) are

often stored for a considerable period either on the gills

or in seminal receptacles of various types and location.

e think it is likely that there may be some correlation

between the presence or absence of storing devices and

the chance of encountering bivalves of the opposite sex.

The commensal galeommatoid bivalves are sequestered

along with their host into a microhabitat that they prob-

ably never leave. A few examples show that the number

of montacutid bivalves present per host (or host burrow)

is usually small (Table 1). Except for H. tenacis, which

has not been anatomically studied, all these species have

evolved various measures that allow long-time storage of

sperm in the female (or hermaphrodite), which effec-

tively enhance the chances that spawned ova can be fer-

tilized even if no sexual partners are around. Conversely,

in a highly social species, like M. gregaria, such mecha-

nisms are evidently much less weeded: It would be in-

teresting to study the relation between the sizes of the

micro- populations i in other Montacutidae that neither ex-

hibit sperm transfer nor sperm storage. Unfortunately,

besides the present species, the known cases are limited

to two species with unknown spatial distribution, Tel-

limya tenella (Lovén, 1846) and Mysella moelleri (Fox et

al. 2007; Petersen and Liitzen, in press).

The present study has shown the need for further in-

formation on the anatomy as a tool of a better under-

standing of the taxonomy, which, to a much too large

extent, has been based only on shell characters. More

data are also wanted on the numerical relations between

hosts and commensals and, above all, analyses on the

sexual behavior of commensal bivalves are in very high

demand.

ACKNOWLEDGMENTS

The study was supported by a grant to JL and AJ from

the Danish National Science Found: ition (no. 51-00-

0278). We are grateful to Mr. Bjarne Bisballe, Zoological

Museum, Cope snhagen, for operating the Scanning elec

tron microscope aid to Mr. Gert Brovad, same institu-

tion, for photographic assistance. We further thank Dr.

Graham Oliver, National Museum of Wales, Cardiff, Uk

for providing information about the occurrence of Hal-

campicola tenacis on its host.

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THE NAUTILUS 121(4):201-209, 2007

Page 20]

Corbula tarasconii, a new species of Corbulidae (Bivalvia) from

offshore Brazil

Eliane P. Arruda

Osmar Domaneschi

Departamento de Zoologia

Instituto de Biociéncias

Universidade de Sao Paulo

Caixa Postal 11461

05422-970, Sado Paulo, BRAZIL

[email protected]

Jonata de A. Francisco

José Carlos N. de Barros

Laboratorio de Malacologia

Departamento de Pesca e Aqiticultura

Universidade Federal Rural de Pernambuco

Avenida Dom Manuel de Medeiros, S/N

52171-900, Recife, BRAZIL

jonatal 98 [email protected]

[email protected]

ABSTRACT

Corbula tarasconii, a new species of Corbulidae is described

from material collected along the Brazilian coast. No living

specimens are known and only shell characters were compared

with the most similar Corbula species from the western Atlantic

and eastern Pacific oceans. Short, ventrally curving rostrum and

sculpture of low, rounded commarginal ribs he wing their bases

about three times broader than the interspaces are the most

important diagnostic characteristics of C. tarasconii. These

characters distinguish the new species from other western At-

lantic and eastern Pacific Corbulidae.

Additional Keywords: Corbula tarasconii, Corbulidae, Bivalvia,

taxonomy, Brazilian littoral, new species.

INTRODUCTION

During a research project carried out by the two senior

authors on Corbula Bruguiére, 1797, occurring in Bra-

zilian sea waters, specimens of an unknown species were

found in samples made at four different localities be-

tween southern Bahia state and northern Rio de Janeiro

state. Working independently, the last two authors found

the same unknown bivalve among the benthic fauna col-

lected during field research on the continental shelf of

northern Bahia state. Careful examination of all speci-

mens gathered by the four authors, plus comparison with

the nominal species hitherto published in the literature,

led to us to conclude that the specimens represent a new

species.

Corbulids are readily recognized by their small size

usually less than 20 mm in she ll length) and their in-

equivalve condition, with the right valve larger, more

convex, and overlapping the — one. All corbulids are

shallow-burrowing suspension feeders inhabiting sandy,

sandy-mud, or muddy substrata, usually at depths greater

than 4 m Lamprell et al., 1998).

Two living subfamilies of Corbulidae, Corbulinae

Lamarck, 118, and Lentidiinae Vokes, 1945, encompass

all extant (~S5) species; a few tropical representatives live

in brackish rivers and streams (Coan et al., 2000). Len-

tidiinae is represented by oan of the single genus

Lentidium Cristofori and Jan, 1832: Corbulinae is by far

more specious and includes 25 genus-level taxa (Coan et

al., 2000), among which Corbula is the largest. Keen

(1969) previously rubdaaded Corbula into 18 subgenera,

several of which were rather poorly defined and in need

of revision, whereas authors such as Warmke and Abbott

(1961), Abbott (1974), Bernard et al. (1993), Coan et al.

(2000), Mikkelsen and Bieler (2001) and Anderson and

Roopnarine (2003) elevated some subgenera to generic

status, a decision not shared by Coan (2002). In this latter

paper, Coan (2002), considered elevating subgenera to

genera premature, because the arrangement of these

taxonomic categories is still fraught with inconsistencies

and additional characters need to be better defined.

The genus Corbula has long been a source of nomen-

clatural confusion and many authors have been at-

tempte -d to resolve it. It is beyond the s scope of this paper

to discuss the systematics of the entire group once Coan

(2002) has already presented a consensus based on the

current rules of the Intemational Code of Zoological No-

menclature (1999),

According to Mikkelsen (2004), there are 13 species of

Corbulidae in the western Atlantic, eight of which were

previously cited by Rios (1994) as occurring on the Bra-

zilian ap Corbula (Corbula) caribaea @ Orbigny,

1853, C. (C.) lyoni Pilsbry, 1897, C. ) patagonica

dOrbigny, oe C. (C.) tryoni E. A. Smith, 1880, C.

(Caryocorbula) sypmella Dall, hae: C. (Caryocorbula)

dietziana C. B, Adams, 1852, C. (Juliacorbula) cubani-

ana dOrbigny, 1853, and C. inate operculata

Philippi, 1848. Revising Varicorbula from the western

Atlantic, Mikkelsen and Bieler (2001) considered C. (V.)

operculata cite od by Rios (1975; 1985; 1994) as a synony-

mous with V. dis) yarilis (VOrb igny, 1842) or misidenti-

fied specimens of V. philippit ( (E. A. Smith, 1885)

Page 202

THE NAUTILUS, Vol. 121, No. 4

In this contribution, we describe a new species of Cor-

bula from Brazilian waters, based on shell characters

only, because no living specimens were obtained, and we

compare this new species with its most closely related

species from Atlantic and Pacific waters.

MATERIALS AND METHODS

Twenty whole shells and 56 disarticulated valves (28

right and 28 left) of the new species were collected on

the Brazilian continental shelf, between northeastern Ba-

hia state (11°58.7' S, 36°49.2' W), and northeastern Rio

de Janeiro state (21°20'28” S, 40°16'09" W). Shell mor-

phology was compared with the most closely related spe-

cies known from the western Atlantic and eastern Pacific,

borrowed from the Departamento de Zoologia da Uni-

versidade de Sao Paulo, Brazil (one lot of C orbula aequi-

valvis Philippi, 1836, and one of C. caribaea dOrbigny,

1853, both without catalog number), Museu de Zoologia

a de Sao Paulo, Brazil ( C. bicarinata G. B.

Sowerby, 1833, lot MZSP 67964), and Santa Barbara

Museum of Natural History, USA (C. marmorata Hinds,

1843, lots SBMNH 83076, SBMNH 131640, and SB-

MNH 141610). Shell characters and illustrations of C. ira

Dall, 1908, provided by Coan (2002), were the basis for

comparison with those in the new spe cies.

The holotype and 14 paratypes were deposited in the

malacological collection of the Museu de Zoologia da

nixersidede de Sao Paulo (MZSP), 15 paratypes in the

Museu Nacional do Rio de Janeiro (MNRJ), and 46

paratypes were deposited in the Museu Oceanografico

Prof. Eliézer de Carvalho Rios (MORG).

SYSTEMATICS

Order Myoida

Family Corbulidae Lamarck, 1818

Subfamily Corbulinae Lamarck, 1818

Genus Corbula Bruguiére, 1797

Subgenus Caryocorbula Gardner, 1926

Corbula tarasconii new species

(Figures 1-15)

Type Locality:

Santo state, 20°45’

Off Guar: Municipality, Espirito

S, 40°25 . Brazil, 60-65 m depth.

Holotype: Museu de Zoologia, Universidade de Sao

Paulo, MZSP 84452 (Figures 1-5, 8, 11).

Measurements: 7 mm length, 5 mm height, 4 mm

width

Paratypes: MZSP 84453 to $4461, northeast of Bahia

state, 11°58.7' S, 36°49.2' W., O1 Nov. 2000, 100 m

(MZSP 84453: |] complete shell; MZSP 84454 to 84461:

6 right, 2 left valves); MZSP 84462 to 84464, southeast of

Bahia state to - ar Best Municipality, Espirito Santo

state, 15°33' S W to 20°45’ S, 40°25’ W (MZSP

84462-84464: 3 ee shells, 1 left valve); MZSP

86026, northeast of Rio de Janeiro state, 21°20'28" S,

40°16'09" W, Feb.—Mar. 1992, 139 m (1 complete shell);

MNBRJ 11146-11157, northeast of Bahia state, 11°58.7

S, 36°49.2' W, OL ne 2000, 100 m (MNRJ 11146,

MNBJ 11147: 2 complete shells; MNRJ 11148-11157: 5

right, 5 left valves); ); MNRJ 11040, southeast of Bahia

state, 15°53'S2” S, 38°31'09" W, 30 Apr. 1996, 66 m (1

right, 1 left valve); ): MNRJ 11812, off Guarapari Munici-

pality, Espirito Santo state, Oct. 1992, 60-70 m (1 com-

plete shell); MORG 50792, off Boipeba Municipality,

Bahia state, 13°35'18.33" S, 38°54'48.27" W, Feb. 2003,

41-53 m (4 complete shells); MORG 50789, off Camamu

Municipality, Bahia state, 13°55'58.79" S, 38°05'28.13”

W, 11 Dec. 2002, 52 m (7 complete shells, 16 right, 19

left valves).

Diagnosis: Shell trigonal to trigonal-elongate, small

(maximum length ~S mm), thin to moderately thick, in-

flated, subequilateral, a a short rostrum, gently curv-

ing ventrally. Left valve smaller than right. Sculpture

sinailar in both shell valves, comprised of lowe regularly

spaced rounded commarginal ribs crossed by minute,

radially arranged pustules: base of commarginal ribs

about three times broader than the anteroogll’ spaces.

Description: Prodissoconch I and II markedly sub-

orbicular. Prodissoconch I with a coarse and irregular

surface texture under the SEM (length: 66.6 to 84.4 wm,

n = 5); prodisocenct IT with subtle growth lines (length:

189.5 to 233.2 um, n = 5) and separated from the dis-

soconch by a sharp transitional line and change in sculp-

ture on the latter. Dissoconch small ( (length: min. = 2.4

mm, max. = 8.11 mm, mean = 5.92 + 1.36 SD [standard

deviation]; height: min. = 1.7 mm, max. = 6.8 mm, mean

= 4.48 + 1.06 SD; n = 47; measurement taken from right

valve), trigonal to trigonal-elongate, moderately diisle in

gerontic specimens, inflated, subequivalve, subequilat-

eral with short rostrum.

Free margin of the right valve completely overlapping

the entire free margin of the left valve. Rostrum acutely

rounded, gently curved ventrally. Posterior slope of each

valve narrow, slightly concave, forming an acute angle

(~20°) between posterior dorsal margin and the lew:

rounded radial keel. Radial keel an inverted, gentle sig-

moid line from umbo to the posterior limit of ventral

margin; plane tangential to posterior slope forming a

slightly obtuse angle with the plane tangential to cenrnl

slope. Valve surface, excluding the posterior slope, regu-

larly convex, except for a slight concavity in ee

ventral area, just anterior to the keels. Umibos prosogy-

rous, at about 36% of shell length from anterior end,

aligned with cardinal tooth on right valve and cardinal

socket on left valve. Anterior dorsal margin straight, ven-

trally directed, continuous with evenly convex anterior

margin, the latter situated below median longitudinal

shell axis; posterior dorsal margin slightly convex, as long

as, but less steep than the anterior dorsal margin; poste-

rior margin long, obliquely truncated, and forming a

short rostrum with posterior end of ventral margin; ven-

tral margin evenly convex, except for a straight to slightly

concave portion just anterior to the radial keel. Juvenile

Arruda et al., 2007 Page 20

Ite mgnht va 3. Internal view of the left valve. 4. Internal view of the right valve. 5. Dorsal view to show the po

keels (broad irrow) and escutcheon (narro irrow). Scale baz 2 mm 6. Paratype MNRJ 11154 external view of a transluce

Page 204

THE NAUTILUS, Vol. 121, No. 4

Figures 7-S.

a thin-shelled specimen (7) and a thick, gerontic specimen (8),

showing differences in the development of the hinge plate at

the resilial socket region (arrow). 7. Paratype MNRJ 11154. 8.

holotype MZSP $4452. Scale bar = 0.5 mm.

shells thin, whitish-translucent, turning moderately thick,

and whitish-opaque as the specimens grow older; peri-

ostracum partially preserved on posterior slope, espe-

cially on left valve, missing on remaining shell area.

External sculpture similar in both shell valves, com-

prised of commarginal ribs crossed by minute pustules

~28.1 wm in basal diameter), the latter showing a ten-

dency to align radially. ( onimarginal ribs regularly

spaced, very low, rounded. with bases about three times

broader than the intercostal spaces; commarginal ribs

bec omimng moderate ‘ly elevated lamellae on poste rior

slope of left valve only. Radial lines of pustules present all

over shell surface, close »r to each other on poste nor slope:

pustule lines visible through translucent shells.

mediately

\rea im-

front of umbos sunken; lunule absent. Es-

cutcheon lanceolate (~1/3 of shell leng h), delimited on

right valve by a low, rounded elevation on the posterior

slope, and on left valve by a slender radial rib; this slen-

der radial rib formed by confluence and abrupt decrease

in height of adjacent commarginal ribs, the last extending

onto umbo almost parallel to free margin of escutcheon,

Inner surface whitish porcelaneous, crowded with ran-

domly scattered submicroscopic pustules (~16.5 jzm in

basal diameter), distinguished under SEM only

Hinge axis almost parallel to anterior dorsal margin.

Right hing “sa with a cardinal tooth aligned with

umbo, and a resilial socket sunken under umbonal

lon: ¢ pe ae pik: ena stout with its apex curled

clors: lly ISOSC( lc trian le shaped when viewed from its

plat narrow deeply retracted at re

Corbula tarasconii. Hinge of the right valve of

Corbula tarasconii. Hinge of the left valve of

Figures 9-11.

thin- shelled specimens (9-10) and a thick, gerontic specimen

(11), showing differences in the development of the hinge plate

at the cardinal socket region (arrow). 9. Paratype MNRJ 11150;

10. Paratype MNBJ 11149; 11. Holotype MZSP $4452. Scale

bar = 0.5 mm

silial socket region in thin-shelled specimens, becoming

expanded, thick, and more evident as specimens grow

older. Left hinge plate with a deep, trigonal cardinal

socket just posterior to umbo, and a thick, short chon-

drophore projecting almost perpendicular to plane of

hinge plate when viewed from its dorsal side; ee plate

narrow, deeply retracted at cardinal socket region in

thin-shelled specimens, becoming expanded, thick, and

more evident as specimens grow older. Dorsal face of the

chondrophore shallowly excavated and divided into an

anterior and a posterior trigonal area by a radially placed,

shallow, narrow groove; posterior margin of posterior

trigonal area becoming thicker and projecting as a stout,

rounded, tooth-like knob as specimens STOW older.

Inner face of the right shell valve bearing a well-

impressed commarginal groove for reception of entire

free margin of the left valve. Anterior adductor muscle

scar ovate, slightly to well-impressed; posterior adductor

scar rounded in frontal view, on top of a slightly to well-

elevated callosity. Anterior and posterior pe ‘dal muscle

scars conspicuous and fused dorsally with corresponding

adductor muscle scar. Pallial line narrow, glazed, far

Arruda et al., 2007 Page 205

Figures 12-17. Scanning electron micrographs of Corbula tarasconii. 12-14. Paratype MZSP 84454. 12. External view of the left

vi ei e showing regularly spe peed: low commarginal ribs (scale bar = 1 mm). 13. Frontal view of the posterior slope to show pustules

radially arr inged and commarginal ribs turning into moderately elevate lamellae. Scale bar = 200 jum. 14. Detail of central slope

(external view) showing the tendency of the pustules to arrange themselves radially. Scale bar = 200 jzm. 15-16. Paratype MZSP

84456. 15. Detail of the umbonal region and the hinge plate region of the right valve showing the low, rounded border of the

escutcheon (arrow). Scale bar = 100 xm. 16. Internal view to show the presence of randomly seattered pustules. Scale bar = 50 jum.

17. Paratype MORG 50789, detail of the umbo to show the limit of both prodissoconch I ‘and II (arrows). Scale bar = 62 wm.

from free border in both shell valves (farther in the right

valve), especially in its anterior two thirds. Siphonal re-

tractor muscle scar straight.

Etymology: This species is named after Dr. José Car-

los Tarasconi, a physician interested in collection and

molluscan studies, who kindly donated the specimen

from his collection, herein designated as the holotype.

Observation: The holotype is the best preserved

specimen among all complete ones; nevertheless, both

shell valves hae. the postero-ventral portion of the ven-

tral margin slightly broken. The brownish-red color

viewed in the internal side of the holotype (Figures 3-4)

was not observed among the paratypes.

Remarks: The gay i in Corbula tarasconii of a

small prodissoconch I (length: 66.6 to 84.4 jum), dis-

Figure 18. Corbula tarasconii. Paratype MZSP $4462. Cam- tinctly separated from the larger prodissoconch I

. : G 9339 5

era lucida drawing of the inner surface of the right valve show- (length: 189.5 to 233.2 pm), the latter devour of surface

ing shell outlines, hinge, muscle scars and well impresse -d com- ornamentation, except for growth lines. 1 his suggests

marginal groove for reception of the free margin of the oppo- that the species has planktotrophie development. accord-

site valve (arrow). Scale bar = 1 mm. ing to the discussions in Jablonski and Lutz (1980) and

THE NAUTILUS, Vol. 121, No. 4

Page 206

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Page 207

Hain and Arnold (1992) on the relationships between

prodissoconch morphology and modes of development.

This new species encompasses all diagnostic charac-

teristics presented by Keen (1969) and Coan et al. (2000)

both for the family Corbulidae and genus em Its

subequivalve, trigonal to trigonal- elongate shell with

moderately coarse commarginal ribs, similar on both

shell valves allow the inclusion of this species in the sub-

genus Caryocorbula Gardner, 1926, as established by

Anderson (1996) and Coan (2002). Beside these charac-

teristics, C. tarasconii shares with Caryocorbula species

a short chondrophore th that projects almost perpendicular

to the plane of the hinge plate when viewed from its

dorsal side.

The allocation of C. tarasconii in Caryocorbula based

in qualitative shell characters is an initial attempt to al-

locate the new species to one of the named subgenera.

As observed by Anderson and Roopnarine (2005),

Caryocorbula “is relatively conservative in its morphol-

ogy, making qualitative methods for alpha-level tax-

onomy difficult.” Subgenera of Corbula have been poorly

defined and fraught with inconsistencies, and a full-scale

revision of the family is long overdue (Coan, 2002).

Much more studies are needed to gather new taxonomic

characters, which could better define the subgenus-level

categories of Corbula and corroborate whether C, taras-

conii is correctly allocated to Caryocorbula.

Eighteen living species of Corbula are currently refer-

able to the eastern Pacific (Coan, 2002) and thirteen to

the western Atlantic (Mikkelsen, 2004). The western At-

lantic species more closely related to C. tarasconii are C.

aequivalvis and C. caribaea, and those of eastern Pacific

water are C. bicarinata, C. marmorata, and C. ira.

Corbula tarasconii is distinguished from C. aequiva-

lvis by being approximately 50% smaller in length, con-

spicuously inequivalve, with the posterior slope set off

from the central slope by a low, rounded radial keel,

sharp and stout in the latter species. The commarginal

ribs in C. tarasconii are low, rounded, with the base

about three times broader than the intercostal spaces and

becoming lamellate on the posterior slope of the left

shell valve, while in C. ae quivalvis they are low to mod-

erately high, each with a quite acute apex and basal width

equivalent to the intercostal spaces. The narrow, lan-

ceolate escutcheon. better demarcated in the left shell

valve of C. tarasconii, is another remarkable difference

distinguishing this new species from C. aequivalvis,

which has a wide, lanceolate escutcheon, wider in the

right valve, and well-demarcated in both valves by el-

evate ribs. The species can also be differentiated by the

form and development of both the chondrophore and the

right cardinal tooth. In C. tarasconii, the chondrophore

has an inconspicuous to small tooth-like knob, and is

shallowly excavated, with the dorsal face divided into two

areas by a low, slender ridge; in C. aequivalvis, the tooth-

like knob is lar ger and higher and the chondrophore is

more aie from the free border of the hinge ai

with the dorsal face divided into two areas by a high

ridge, with the anterior area deeply excavated. Viewed

from its convex face, the right cardinal tooth is right-

triangle-shaped in C. aequiv alvis and isosceles-triangle-

shaped in C. tarasconii.

Corbula tarasconii greatly differs from C. caribaea by

its short, ventrally curving rostrum that in C. caribaea is

moderately to well produced and aligned with the an-

tero- -posterior shell axis. Viewed from its inner surface,

the posterior margin of the rostrum, in most individuals

of the latter species, has a sinuous outline; the rostrum is

frequently extended farther posterior by lateral, siphonal

plates made of calcified periostracum. The rostrum of C.

tarasconii neither has sinuous outline nor siphonal

plates.

The eastern Pacific C. bicarinata, compared to C,

tarasconii, has an oval-subquadrate to trigonal outline, a

shallow depression on the disc area farther anterior to

the radial keel and aligned with the umbo-ventral axis,

and the posterior slope set off from the central slope by

a sharp, stout radial keel. The right cardinal tooth with a

right-triangle shape and the wider, almost fan- shaped

escutcheon, set off from the posterior slope by two stout,

lateral ribs in C. bicarinata also differentiate it from C.

tarasconii.

Based on the figures and description given by Coan

(2002), the eastern Pacific Corbula ira is similar to C.

tarasconii in outline, configuration of the retractor sipho-

nal muscle scar, umbos position, but greatly differs in its

larger size and disc area sculptured vault strong, less nu-

merous, rounded commarginal and fine radial ribs.

Corbula marmorata, the third eastern Pacific species

closely related to C. tarasconii, has a more elongate,

trigonal-ovate shell sculptured with strong, high, and

acute commarginal ribs anterior to the radial keel. Both

species share a shallow depression just anterior to the

posterior radial keel and a short, ventrally curved ros-

trum, but the former species is also differentiated by the

presence of a second shallow depression aligned with the

umbo-ventral axis.

ACKNOWLEDGMENTS

The authors are greatly indebted to Paul Valentich-Scott,

Santa Barbara Museum of Natural History, Santa Bar-

bara, Luis R. L. de Simone, Museu de Zoologia da Uni-

versidade de Sa0 Paulo (Brazil), Norma C. Salgado, Mu-

seu Nacional, Rio de Janeiro (Brazil); and Paula Spo-

torno de Oliveira, Museu Oceanografico Prof. Eliézer de

Carvalho Rios (Brazil) for the loan of specimens of Cor-

bula spp. Our thanks are due to Enio Matos and Eduardo

Matos who provided assistance in the techniques of

SEM, Departamento de Zoologia do Instituto de Bio-

ciéncias da Universidade de Sao Paulo for scientific and

infrastructural support, and FAPESP - Fundagao de Am-

paro a Pesquisa do Estado de Sao Paulo that provided

funding for this work. Special thanks to José H. Leal,

Paula M. Mikkelsen and to the anonymous reviewer for

their valuable comments on the manuscript.

Page 208

THE NAUTILUS, Vol. 121, No. 4

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THE NAUTILUS 121(4):210-213, 2007

Page 210

Two new gastropod species (Neogastropoda: Drilliidae,

Turridae) from the western Atlantic Ocean

Donn L. Tippett

10281 Gainsborough Rd.

Potomac, MD 20854 USA

ABSTRACT

Two new deep-water species from the western Atlantic are

proposed: Drillia (Clathrodrillia) blakensis and Hindsiclava

rosensticlanus. Drillia blakensis is nearest Drillia (Clathrodril-

lia) petuchi Tippett, 1995, and Hindsiclava rosensticlanus re-

calls Hindsiclava polytorta (Dall, 1881). Animal anatomy, es-

pecially foregut anatomy, is described for H. rosenstic anus.

Additional Keywords: New species, Brazil

INTRODUCTION

The species proposed here are examples of the richness

of the turrid fauna that continues to be discovered in the

deep waters of the western Atlantic. Although collected

in the 1960s, neither has been prey ously reporte sd. The

type material of Drillia blakensis was part of the Bullis

collection, secured as by-catch from the R/V OREGON.

Hindsiclava rosenstielanus was dredged by the Univer-

sity of Miami's R/V PILLsBury, but was only recently

discovered during a review of the ane unsorted

portion of the mollusk collection at the

Miami's Rosenstiel School of Marine and pune

Science. It is evident that further exploration and re-

search will continue to reveal new material.

MATERIALS AND METHODS

Empty shells and shells with preserved animals were ex-

amined. Preserved material was dissected. Radulae were

mounted on microscopic slides and stained with Pronto-

cil + CMCP 10, Type specimens were deposited at the

National Museum of Natural History and other institu-

tions. The classification used is that proposed by Taylor,

Kantor, and Sysoev, 1993, which involved a rearrange-

ment of the tre adition: I classification of the Turridae. Ab-

breviations are: ANSP, Academy of Natural Sciences,

Philadelphia Pe nnsylvania; MCZ, — um of Com-

parative Zoology, Harvard Unive rsity,

sachusetts: UMML

School of Marine and Atmospheric Science, University

of Miami, Florida; USNM, National Museum of Natural

History, Smithsonian Institution, Washington, DC

Cambridge, Mas-

Invertebrate Museum, Rosenstiel

Iniversity of

SYSTEMATICS

Drilliidae Olsson, 1964

Genus Drillia Gray, 1838

Type Species: Drillia umbilicata Gray,

sequent designation, Gray, 1847.

1838, by sub-

Subgenus Clathrodrillia Dall, 1918

Type Species: Murex gibbosus Born, 1778, by original

designation.

Drillia (Clathrodrillia) blakensis new species

(Figures 1-3, $)

? Turridae sp.—Lamy and Pointier, 2001: 22, number

73, list, p. 26, photo.

Description: Shell of medium-size (to approximately

45 mm), elongate, fusiform, with tall spire and large body

whorl measuring about 0.5 shell length. Shell tapering

gradually with moderate basal constriction to moderately

elongate, open, slightly notched, slightly bent right ante-

rior canal. Protosanch decollated, teleoconch w hols ten.

Whorls well-rounded, shoulder sulcus on upper third

concave, suture prominent. Sculpture of rounded, regu-

larly spaced axial ribs with equal interspaces, e xtending

from shoulder to followi ing whorl on spire and to base on

body whorl. Ribs increase in number with shell growth,

seven on early whorls, 12, narrower and closer spaced, on

penultimate, ten or 11 on body whorl leading to small

varix 0.25 whorl back of lip e dee, two or theese possibly

abortive ribs following varix. Fine spiral threads overall,

weaker on sulcus. Ape rture parallel-sided with apically

directed, U-shi ape vd sinus poste poke bordered on body

whorl by flat parietal tubercle. Lip sloping roundly for-

ward below sinus, upper edge directed upward, narrow-

ing sinus somewhat, producing spout-like appearance.

Stromboid notch distinct. Color dirty-white overall, faint,

pale brown peripheral band, blotch of same color on

varix and spots preceding tops of axial ribs on later

whorls. Operculum (Figure S)

with roundly pointed anterior end and terminal nucleus.

of chestnut color, ovate

Type Material: Holotype, USNM 900034, 400450. mm,

May 1965; three paratypes, USNM_ 1096708, data same

Tippett, 2007

Figures 1-7.

44.5 x 13.6 mm, apertural, lateral, dorsal views.

mm, off Cape San Antonio, Cuba. Scale bar = 25 mm.

as for holotype, 45.7 x 16.6 mm, 44.1 x 14.7 mm, 42.5 »

16.0 mm (ex-José and Marcus Coltro collection., ex-

author's collection); all dredged by R/V OREGON on type

locality.

Other Material Examined:

elas Wolfe collection , 41.2 x

OREGON on type locality.

one specimen, Dr. Dou-

15.5 min, dredged by R/V

Type Locality: Blake Plateau; precise location un-

known, data presumably not retained.

Distribution: Blake Plateau and possibly off Saba Is-

land, Netherlands Antilles (Lamy and Pointier).

Discussion: Drillia blakensis is most similar to Drillia

Clathrodrillia) petuchi Tippett, 1995, from which it dif-

fers in being narrower, having less robust ribs, finer spi-

rals. a broader, non-tabulate, more sloping sulcus, and

fainter color pattern. The author has not seen the shell

figured in Lamy and Pointier, stated to be 48 mm in

ength and from 150 m depth, however it appears from

he illustration to be D. blakensis, differing in being

slightly broader, having a slightly shorter anterior canal

and stronger peripheral color banding, features within

intraspecific variation limits.

Etymology: Named after the Blake Plateau, the type

ocality.

Turridae H. Adams and A. Adams, 1853 (1838)

Crassispirinae Morrison, 1966

Genus Hindsiclava Hertlein and Strong, 1955

Type Species: Clavatula militaris Hinds, 1543, by

original designation.

Hindsiclava rosenstielanus new species

Figures 4-6, 9, 10

7. Pleurotoma (Drillia) polytorta Dall, 1881, holotype,

Drilliids and turrids. 1-3. Drillia (Clathrodrillia) blakensis new species, holotype, USNM 900034, 44.8 «16.0 mm,

apertural, lateral, dorsal views. 4-6. Hindsiclava rosensticlanus new species, holotype, USNM 1086746, off Riohacha, off Colombia,

USNM 412171, 32.6 x 9.6

Crassispira polytorta (Dall, 1881): Okutani, 1983: 304,

description and figures (apertural and lateral views plus

radular teeth).

?Turridae sp.—Lamy and Pointier, 2001: 22, number 71,

list and photo.

rece iis

48 mm), elongate, fusiform, turreted, with tall spire,

body whorl about 0.4 of shell length, whorls rounded

below concave shoulder slope, gently constricted at base

to moderately long, anterior canal open, notch lacking.

Protoconch with 2.5 smooth whorls, tip central, first 1.5

whorls glossy, remainder dull-white, terminating in short

portion of whorl with 4-5 axial riblets that quickly en-

large to axial ribs in adult shell. Teleoconch whorls 9-10.

Ape rture of parallel sides and with moderately deep,

U-shaped posterior sinus on shoulder slope. Pé irietal tu-

be rcle absent. Suture distinct, almost channeled, slightly

Wavy. Subsutural cord of double threads. Axial ribs ro-

bust (11-12 on body whorl, 11 on penultimate), extend-

ing faintly across sulcus above and to next suture on

Shell of medium-size (to approximately

spire, disappearing on base; interspaces half again as

wide. Uniform, regularly spaced spiral threads (5-6 on

whorl periphery) cross ribs, producing modest, laterally

elongate nodules, then continue forward to anterior ca-

nal. Microsculpture of fine spiral threads, including sul-

cus, irregular in strength and distribution, Notch and

varix lacking. Color dirty-white overall, traces of dark

periostracum.,

Gross Anatomy: Animal ee foot with upturned

propodium operculum locate d posteriorly on foot

Head bearing two tentacles, e me with eye dorsally on an

co gee midway from base. Penis behind right ten

tacle. reflected back under mantle Respiratory siphon

on left a

head, with moderate anal sinus on right. Gills and os

bearing a fold, mantle edge extending across

Page 212

THE NAUTILUS, Vol. 121, No. 4

Figures 8,9. Opercula, inner (left) and outer (right) views.

8. Drillia (Clathrodrillia) blakensis, 7.5 min length. 9. Hindsi-

clava rosensticlanus, 11 mm length. Seale bar = 5 mm

phradium visible through mantle. Rectum on right,

Puckered rhynchostome between and slightly below ten-

tacles. R hyne hostomal sphine ter present. R hynchocoel

with strong linear folds internally. Rhynchodeal wall cir-

cularly folded ef to retraction. Proboscis long, circularly

folded along its length in preserved animal due to con-

traction, linear folds posteriorly Large buecal mass and

ivity posterio1 to rhynchodeum. No sphincter seen at

beginning of esophagus. Large, highly coiled poison

cland and bulb

‘lands present Radular ribbon ventral and posterior to

body cavity. Right and left salivary

Figure 10. Hindsiclava rosenstieclanus, radula, teeth are 300

um length.

buccal mass. Gland ducts and radular ribbon enter at

beginning of esophagus ventrallly and posteriorly to buc-

cal mass. Incomplete radula an approximately 40 pairs

of wishbone marginal teeth (Figure LO) measuring ap-

proximately 300 jum. Operculum (Figure 9) medium am-

ber, ovate with flat proximal side, rounded peripheral

side, ends rather sharply rounded, terminal nucleus at

anterior end,

Type Material: Holotype, USNM 1086746, west of

Riohacha, off Colombia, 11°32’ N 073°23' W, 549

R/V Pillsbury 781, 30 July 1968, 3 m, otter trawl, ex-

UMML.30.10758; paratypes (ex-UMML,): one specimen,

USNM 1107006; one specimen, MCZ 359155; one speci-

men, ANSP 416320; seven specimens UMML 30.10758,

four with animal preserved, three shells only. All from

type locality.

Other Material Examined: USNM 902064, three

specimens, off Cartagena , Colombia, (ex-José and Mar-

cus Coltro collection, ex-author’s collection); Pleuwrotoma

(Drillia) polytorta Dall, 1SS1, USNM 412171, holotype.

Type Locality: West of Riohacha, off Colombia.

Discussion: Hindsiclava rosensticlanus is most similar

to Hindsiclava polytorta (Dall, 1881) (Figure 7) reported

from off Cuba, Hindsiclava rosensticlanus differs by its

wider sulcus, doubled subsutural cord, fewer ribs (e.¢.,

nine on sixth spire whorl versus 12 on equivalent whorl of

polytorta), presence of fine secondary spiral threads

overall, and absence of parietal tube rcle. The specimen

reported by Okutani, measuring 58 x 19 mm, from 328—

470 m off Surinam, appears to be this species despite the

larger size and the radular teeth which are dissimilar at

the location of their basal attachment (drawings appear

stylized). The specimen reported by Lamy and Pointier

(2001) from 450 in off Point Noire, Guade loupe, is 63

mm in length and, appears to be this species, again de-

spite its larger size, and also despite the geogr: phic dlis-

tance of Guadalupe from the type loc ality of rosenstiela-

nus. The specimens in USNM 902064 are identical with

the type mate rial exce pt that the axial ribs are slightly

broader.

Tippett, 2007

Page 213

Etymology: The species is named for the Rosenstiel

School of Marine and Atmospheric Science, University

of Miami, from which the eal was obtained.

ACKNOWLEDGMENTS

The author thanks the Department of Invertebrate Zo-

ology, National Museum of Natural History, Smithsonian

Institution for the opportunity of wor king with the mol-

lusk collection and use of the equipment and facilities.

Nancy Voss donated specimens of Hindisclava rosen-

stielanus from the UMML. Yolanda Villacampa prepared

the SEMs. Dr. Jerry Harasewych was constantly helpful

in providing use of equipment, advice and support. The

author thanks these people.

LITERATURE CITED

Dall, W. H. 1SS1. Reports on the results of dredging under the

supervision of Alexander Agassiz, in the Gulf of Mexico

and in the Caribbean Sea, 1S77—79, by the U.S. coast

steamer “Blake,” Lieutenant-commander C. D. Sigsbee,

U.S.N., and Commander J. R. Bartlett, U.S.N.,

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Lamy, D, and J-P. Pointier. 2001. Les molluques profonds des

Antilles Frangaises. Xenophora 95: 21-27.

Okutani, T. 1983. Mollusks. In: Masatsune, T. and T. Okutani

(eds.) Crustaceans and mollusks trawled off Suriname and

French Guiana. Japan Marine Fishery Resource Research

Center, pp. 187-354.

Taylor, J. D., Yu. I. Kantor, and A. V. Sysoev. 1993. Foregut

anatomy, feeding mechanisms, relationships and daseitt.

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170.

Tippett, D. L. 1995. Taxonomic notes on the Western Atlantic

Turridae (Gastropoda: Conoidea). The Nautilus 109: 127—

135.

;

&, =

;

THE@MNAUTILUS

Volume 121

2007

AUTHOR INDEX

ARRUIDA. Fi. IPs cote acdatmeactnin deed tratingunadseuankentiaenacehin 201 IIMAs Si Fe Bicwascseatecetancen arn dtwadagatdins aasutabiccsduh aan’ 99

BANDYOPADAYAY: Po Ks ccscscisveenesarsatetereaahe ovcuewesorataes 131 LOTZEN,, ewe inkses nenctaan tededdeanaa rontaesdsensatas scencatvanas 191

BARROS: [Guise ecsassneacsalsensseeisas tematssynaantesne 95, 99, 201 MARIOUIING, Py ote sitsctetaanssanitecesinie rangi eeselas srenteantes 159

BEU As Concpapetatonzatsutsaknbenuisannshddanaeeoasaaernns Scatenensn 90 IVECUINTAING Gz ne hcnkronnexsaaaniorranaaane noiathenwhocaunesnummaacete 139

BUYNEVICH Te Visca ccadisaastnccessactaliisessndinaiaradecercesnnes 37 INEKOILAS: [oacinedtinatactier dues aarinan sipinedssodmemmarenetnds 17

GOTEES alae He pita ty ieccheideta este eset tera decate cine ncatitadnuunsartatte eit eySars 17 OTIVERAS. Bist MMs siehose sa cweacacivsesesnss-aensarnsaitidoacdravns efetdle dean aR ncdlvetni a bu 13

GONCEPCION: (GPs asca Se aves Sanee sn eniosannanccneertesatoreestes 131 OWNBY), iePoicats hnensssegtwecetiae te cuou.ancs nettseta ricedihuawhde 13

POE VIRMES: Miva des ce aeteden cars hcmtete mia eonseaca taut agauamecanatatnaee 163 PEARCE,“ Ve fusauseiauessactenediirons aun teueteauewnteteeniaens 66

IDOMANESGHI, 'O% occ cinsccdexd ceteaenvasenh des thie cove seiabohenban 201 PENGHASZADEH: (Pie ss.4ccccetcnacoanactaantiecisagaidecavasasess 139

PRED Sy Mi Gi scccnosiianstana narakguaa adiscmeneainnonanuae Masten ceive 66 PETG, Ré Bi asnavoucsaviasendesstndnatcigestccans anadecaanenes 95, 99

NOs Wa ree cae Ae tea sce densi diese turecunamsniacdadanheatacd tena 191 PORTIS Ric Ws.53 vocaGanee gones-onigidtennalanssteiadatdacandeiaaens 105

IRANGISCO,, |). Asi. 2s.ctneneoametecesee vie doszetaie dnnmuatentesiae 201 ROTVIT:, Das iceland Co ncrua ne bed tenant oetonnd ade saakuuudeyaiteoeeoes 191

WREST D. Ye pcscceacsshteadhe2 teearianseideatais Moanenan tanner 43 ROY, Miviissesssicicisissusvorscctsvassaeristesiwnssasacsiesasieoeness 20

GAWEARDO, AG. Sizccdacacidcctaanavicassmeriaciesgwinnaagenaanenceeat 139 SANTOS SA DDS ss scsctaaesaincrancnaaaduinancad sent naewntaatincnd mi me 131

IARASEWYCH;, Mi (G..d.d esa ccneessassanctasanngadesave san 76, 90, 146 SAWE; Lisaas cvesacatnoussaantadgeesanida de odteah a concaamnadereaa avons 1

FIARDING, IM. oiiet oncedesenadsnsaan redisatecmacteatt aaageanssietons 146 SCHMELZ, G. W. 2... cc ccc cece cence eee e scenes eeenenesneeseenseees 105

EIERAEDE, Py Mi MT cenoncsesantnccoa vate unennmue oSeawead Wiecades 131 SLARGINSKY;. Joo. detawnssaacs dacs ihe senoneiasctee us tsaar cect nan ene 182

JESPERSEN, Ava ccsssecaccdsstaiiiadsensancarpncatensaateuhavas sabnnes 191 SQUIRES. Re Discs. diwasai shicwaricbidboiieas dest anercmdesieaances 1

INTEia 0: se anaes aabit weemacaseeoseenaah gaataeie heacecaminsaasenkaecans 76 STRONG 3 Eis: los dsasanainacasneacinhantradteniae tin manasncnsoecdet tinted 43

INURITA:: Kix foseucaSes toavawandeincd stuse cade wanendicaaceadieeaasawenes 66 PPE TAD Te Soon ii detehernt eae tee sciet sanewseae Reoeeee 210

TASTE: RS sole duciztsindemearsersittsts ie atssniernde seeadaednamatuenanenetelle 182 WY ATI =. ME secccc2 ccaateeete rade setae eine teins fied Gite awl aiaalatiarbiarae es 131

Pe MG Ss vec cat wn atest oaicuta ne apecauadsesaaaatnnnisaninanncoa te 104 WAEIAMSS WD TD ecincnspadsaawegeSedana socacd adn stwhcadsnah adeno 0

NEW TAXA PROPOSED IN VOLUME 121]

GASTROPODA

Bathrotomaria annejoffeae Harasewych and Kiel, 2007, new species (fossil, Pleurotomariidae).. 2... ee S4

Bathrotomaria bedetteae Harasewych and Kiel, 2007, new species (fossil, Pleurotomariidae) . 2... 82

Brachycythara beatriceae Mariottini, 2007, new species (Conidae) 0... ee 159

Cantallocostoma DeVries, 2007, new genus (fossil, Trochidae) .. 0... 169

Cantallocostoma panistostum DeVries, 2007, new species (fossil, Trochidae) .. 2... 0. ee 171

Chlorostoma quipua DeVries, 2007, new species (fossil, Trochidae). .. 0... 167

Cirsotrema chipolanum Schmelz and Portell, 2007, new species (Epitoniidae) .. 2... ee 110

Drillia blakensis Tippett, 2007, new species (Drilliidae) 2... 210

Epitonium conwaiae Schmelz and Portell, 2007, new species (Epitoniidae) .. 2... ee 122

Epitonium hoerleae Schmelz and Portell, 2007, new species (Epitoniidae). 2... eee 124

Epitonium incomitatum Schmelz and Portell, 2007, new species (Epitoniidae) Stata, Saas £0 ak Ue oes ee Bae es 120

Epitonium kalistos Schmelz and Portell, 2007, new species (Epitoniidae) . 2... 126

Epitonium regina Schmelz and Portell, 2007, new species (Epitoniidae). 0... 120

Epitonium vokesae Schmelz and Portell, 2007, new species (Epitoniidae) . 6... ee 126

Gerdiella alvesi Lima, Barros, and Petit, 2007, new species (Cancellariidae).. 2... . 100

Hindsiclava rosenstielanus Tippett, 2007, new species (Turridae) .. 2... 0 ee teats . 211

Intistoma DeVries, 2007, new genus (fossil, Trochidae) .. 2... ee eee 17]

Intistoma pirqua DeVries, 2007, new species (fossil, Trochidae). 2... 0. ee ee a E72

Microcancilla jonasi Barros and Petit, 2007, new species (Cancellariidae)... 2... 02... boa) A emdvesr i BAL UE asitactated . 96

Paryphantopsis corolla Slapcinsky and Lasley, 2007, new species (Charopidae)..... 2.0... . Bees By & dine, Gos Bee are 183

Paryphantopsis fragilicosta Slapcinsky and Lasley, 2007, new species (Charopidae) . . . . . SG jun & Aone Gee pier _ 185

Paryphantopsis nucella Slapcinsky and Lasley, 2007, new species (Charopidae).. 6... 0. ee ee . 187

Obornella thompsonorum Harasewych and Kiel, 2007, new species (fossil, Pleurotomariidae)..... 6.2.0... Ais tits a ey MOO

Opalia mica Schmelz and Portell, 2007, new species (Epitoniidae) ....... . eed GP URE ae aad pret teed Gate 2 3. da

TTD Ee? IN A A WS

Volume 121

2007

Opalia politesae Schmelz and Portell, 2007, new species (Epitoniidae).. 2... ee 115

Sassia melpangi Harasewych and Beu, 2007, new species (Ranellidae). 2... ee 90

Tegula masiasi DeVries, 2007, new species (fossil, Trochidae) .. 2.0... ee 175

Vertigo malleata Coles and Nekola, 2007, new species (Vertiginidae). 2... 19

BIVALVIA

Corbula tarasconii Arruda, Domaneschi, Francisco, and Barros, new species (Corbulidae). .. 0... 0. ee ee 202

Mysella gregaria Rotvit, Liitzen, Jespersen, and Fox, new species (Montacutidae). 2. ee 192

Warren D. Allmon

Rafael Araujo

Jon Bryan

Lyle Campbell

Eugene V. Coan

Robert H. Cowie

Robert T. Dillon, Jr.

Marien Faber

Diarmaid O Foighil

Matthias Glaubrecht

Jochen Gerber

M. G. Harasewych

Kenneth Hayes

REVIEWERS FOR VOLUME 121

Gregory Herbert

Robert Hershler

Steffen Kiel

Frank Koehler

Paula M. Mikkelsen

Patricia Miloslavich

Jeff Nekola

Sven Nielsen

Anton Oleynik

P. Graham Oliver

Marco Oliverio

Guido Pastorino

Kathryn E. Perez

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

ENDOWMENT

FOR THE ARTS

Tra Richling

Barry Roth

John Slapcinsky

Richard L. Squires

Ellen E. Strong

Jouni Taskinen

Donn L. Tippett

Paul Valentich-Scott

David Véliz

Geerat |. Vermeij

Janice Voltzow

Diego Zelaya

iu a

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