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k'HE NAUTILUS

Volume 123, Number 1

March 31, 2009

ISSN 0028-1344

A qua rterhj devoted

to malacologii.

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THE€7NAUTILUS

CONTENTS

Volume 123, Nu))ihcr 1

March 31, 2009

ISSN 0028-1344

Diego G. Zelaya A redefinition of Pseudokelh/a Pelseneer, 1903 (Bivalvia: Cyainiidae) and tlie

Cristian Ituarte description of a new species from the Sontfiern Ocean 1

John D. Taylor Phylogenetic position ol the liivalve family Cvrenoididae — remo\ al Irom (and

Eniily^ A. Glover further dismantling ol’) the siiperfamily Lncinoidea 9

Suzanne T. Williams

Donald F. MeAlpine Establishment and persistence of the copse snail, Arionfd arbusfonnn (Linnaeus,

Frederick W. Sehueler 1758) (Gastropoda; Helicidae) in Canada 14

John E. Maunder

Ronald G. Noseworthy

Man C. Sollows

Research Note

Juan M. Diaz

Fernando Gasl

Diana C. Torres

Research Note

Fred G. Thompson

Michael \\^. Hevn

Drew N. Camphell

Rediscoveiy ol a Caribbean living fossil: Plioladomi/a Candida G.B. Sow^erbv I,

1823 (Biv'alvda: Anomalodesmata: Pholadomyoidea) 19

Thiara scabra (O. F. Aliiller, 1774): Tlie introduction ol another Asian Ireshw'ater

snail into the United States 21

Erratum

Roland Hoiiart

Carole M. Hertz

A review ol Ti/phisopsis jousseaume, 1880, and Ti/phisala |ousseanme, 1881

(Gastropoda: Muricidae) ol the eastern Pacific (2006) 23

Notice

24

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3

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THE NAUTILUS 123(1): 1-8, 2009

Page 1

A redefinition of Pseiidokellija Pelseneer, 1903 (Bivalvia:

Cyamiidae) and the description of a new species from the

Southern Ocean

Diego G. Zelaya

Division Zoologia Im'ertebrailos

Mnseo cle la Plata, Paseo del Bosque s/n

1900 La Plata, Buenos Aires,

ARGENTINA

[email protected]

Cristian Ituarle

Division Zoologia Invertehrados

Mnseo Argentine de Cieneias Naturales

Av. Angel Gallardo 470

C1405DJR, Buenos Aires, ARGENTINA

ABSTRACT

A new species oi Pseitclokelli/a, Pseiiclokclh/a fninki Ironi Soutli

Shetland Islands, is described. P. fninki is characterized by its

subcircular shell outline, the periostracum forming low lamel-

late commarginal folds, radial sculpture absent, and by the

presence of a single posterior siphonal opening, a fact that is

in contrast with that reported for the t\pe species of Pseiulo-

kelh/a, Kcllia cardifonnis Smith, 1SS5. The presence of a com-

plete follicle surrounding each developing ooc)4e, persisting

throughout vitellogenesis, a condition not currently knowi for

other bivalves, is confirmed as a generic diagnostic character.

The generic redefinition, based on shell morphology and ana-

tomical features described for Pseudokclhja fninki. is given.

Additional Kci/icords: Cyamioidea, Ciiamiocanlimn. Kcllia,

Perrierina. reproduction

INTRODUCTION

Pseiidokelh/a was proposed by Pelseneer (1903) to real-

locate Kcllia canlifonnis Smith, 1885, a species

described from Kerguelen Islands. The diagnosis for the

new genus was given in association wdtli description

of the anatomy oi Kcllia carclifonnis. Species oi Pscudo-

kellija were originally distinguished from those of Kcllia

Tnrton, 1822, by having two, brancliial and anal, siphonal

openings and for being dioecious, Pelseneer (1903) also

reported a peculiar reproductive trait for Pseitdokelh/a

carclifonnis: the presence of a complete follicular epithe-

lium snrroimdiirg each developing oocyte, a condition

not otheiwidse known for bivalves. Snbsecpiently, four

nominal species of Pseudokelhja were described: Psetido-

kelhja gradata Thiele, 1912, from Gauss Station, Pseudo-

kellija stillwelli Hedley, 1916, from Adelie Land and

Davis Sea (a synonym of F, carclifonnis, according to Dell

(1990)), Pseudokelhja georgiana Dell, 1964, from South

Georgia, and Pseudokelhja inexpectata Dell, 1964, from

South Georgia and South Orkneys. The descriptions

of these four species were based exclusively on shell

characters; after tliat not a single study provided infor-

mation on their anatomy. Consequently, to date, it is not

possible to confirm if the anatomical characters reported

by Pelseneer (1903) are diagnostic for Pseudokelhja car-

difonnis or shared by other species of the genus.

In the present paper, a new species of Pseudokelhja

from South Shetland Islands is desciibed; details on

anatomy, reproductive traits and shell morphology

provide additional information for a better definition of

the genus.

MATERIALS AND METHODS

The specimens studied were originally deposited at the

Zoologisches Museum (ZMB), Germany. They were col-

lected during the 1982, 1985, and 1986 Soviet Antarctic

Expeditions to King George Island, South Shetland

Archipelago. Voucher specimens are deposited at the

ZMB, Mnseo de La Plata (MLP), and Mnseo Argentine)

de Cieneias Naturales (MACN), Argentina.

We studied the holoRqre of F gradata (ZMB 63109),

specimens of F. cardiforniis (ZMB 63136), F inexpectata

(ZMB 114683), and F. georgiana (MLP 12999) from

t\qre localities, and specimens of F. gradata from Zoo-

logische Staatssarnmlnng Miinich (ZSM), Germany

(ZSM 20012865: 63° 01.10' S, 61° 09.10' W'; ZSM

20041320: 62° 00.09' S, 60° 19.31' W). Specimens cur-

rently assigned to Kcllia suhorbicularis (Montagu, 1803)

(MLP 11563); Kcllia niagellanica Smith, 1881 (MLP

13000); and Kcllia sp. (AILP 13001) from Argentine

waters were also used for comparative purposes.

The anatomical description of Pseudokelhja franki was

based on dissections under stereoscopic microscope;

seven specimens were pi'ocessed for histology, inclusion

was performed either in Paraplast® or 1 listoresin®;

specimens were completely sectioned at 5 pm thick,

using a Leica RM 2355 microtome.

Shell morphology was studied throngli scanning

electron microscopy (SLM). Shell measurements were

Page 2

THE NAUTILUS, Vol. 123, No. 1

obtained according to the following criteria: L: niaxininin

antero-posterior distance; H: inaxinmm dorsoventral dis-

tance perpendicular to L; W: maximnin distance across

\ al\ es. Mean value and standard deviation for the ratios

H/L and VV/H are given (n = 16 specimens). Hinge teeth

nomenclature is indicated in figures 10 and 11.

SYSTEM ATICS

PseiiclokeUi/a Pelseneer, 1903

T>pe Species: KcUi/a cardiformis Smith, 18S5 (hy

monohpy)

Pscudokelhja franki new species

(Figures 1-26)

Diagnosis: Shell suhcircnlar, inflated, only sculptured

with marked growth lines; periostracum forming lamel-

late commarginal folds. Posterior portion of the right

cardinal tooth (C3b) well developed. A single posterior

siphonal opening, the anal, present.

Description: Shell small, maximnm ohsei'ved L = 4.2

mm, shell outline subcircular, slightly longer than

high (H/L = 0.94±0.04), inflated (W/H = 0.74T0.04)

(Figures 2-5, 12). Anterior margin short and round,

imperceptibly connected w4th dorsal margin, forming

a \\4de cun^e with the anterior part of ventral

margin (Figures 2-5). Ventral margin markedly cun'ed.

Posterior margin rounded, nearly vertical in larger speci-

mens following the posteroventral cun'e (Figures 2-4).

Posterior part of dorsal margin snherect or slightly

cmved. Beaks prosogyrous, inflated, globose at tip,

strongly discernible above dorsal margin, slightly dis-

placed anteriorly (Figure 2-5, 7). Prodissoconch ovate,

about 400 pm in diameter, surface sculptured with

minute granules (Figures 6, 13). Shell surface whitish,

shiny, with veiy low and I'onnded commarginal grow'th

lines, irregularly distrilnited (Figure 15). Periostracum

translucent, forming low lamellate periostracal folds

(Figure 14). Inner shell surface whitish, dull. Hinge

plate narrow, somewdiat enlarged anterior to beaks, just

at the point of insertion of cardinal teeth (Figures 8-11).

Hinge: left valve (Figures 8, 10): cardinal tooth 2 (C2)

solid, triangular, cusp subceutral; cardinal tooth 4 (C4)

relatively short, straiglit, and solid, with cusp displaced

posteriorly; lateral posterior tooth (LIl) long, narrow,

and low, well separated from posterioi' margin. Right

valve (Figures 9, 11): cardinal tootli (C3) hook-like,

formed by large, solid anterior portion (C3a), bifid at

base, and sliort and narrow posterior portion (C3h), In

larger specimens, C3a and C3b form a nearly right

angle (Figure 11). whereas in smaller ones the angle

between C3a and C3b is more acute and C3b longer

(Figure 9). Right inner posterior lateral tooth (LI) long,

moderately solid, with centrally located cusp; outer pos-

terior lateral tooth (LIII) merged with dorsal shell mar-

gin. Internal ligament set in a small, short, shallow

resilifer posterior to cardinal teeth; e.xternal ligament

short, posterior to beaks. Scars of anterior and posterior

adductor muscles ovate, the anterior, slightly longer

(Figure 7). Pallial line entire.

Anatomy: Mantle margin largely unfused, forming a

long pedal gape (Figure 20), fused at the posterior quar-

ter, delimiting the anal opening; below this point, an

enlarged portion of the middle mantle fold corresponds

to the position of the absent branchial mantle opening

(Figure 19). The anal opening and the enlarged portion

of the mantle margin are flanked Iry a row of 12 to 16

micropapillate tentacles on each side, placed in an alter-

nating pattern (Figures 18, 19). Auteilor and posterior

adductor muscles almost equal in size, the posterior one

ovate in section, the anterior more elongated and nar-

rower (Figure 16). Foot long, w4th a well differentiated

heel (Figure 16); a small byssal gland, functional

Figures 2-15. Fficiidokclhia franki new' species. Specimens from Maxwell Bay, King George Island, 100 m (station 30/49). 3, 4.

Holop pe. 2, 5-15. Otlier specimens. 2-5. Outer view. 2, 3. Riglit \ alve. 4. Left valve. 5. juvenile. 6. Prodissoconch. 7. Inner view,

lelt valve. 8-11. Details of hinge plate. 8, 9. Specimen 4.2 mm L. 10, 1 1. Specimen 5.5 mm L. 8, 10. Left valve. 9, 11. Bight valve.

12. Dorsal v'iew. 1.3. Detail of prodissoconch .sculpture. 14. Periostracal folds. 1.5. Periostracal folds and grow'th lines. Scale bars:

2-4, 7 = 1 mm; 5 = 250 pm; 6 = 100 pm; 8-11 = 200 pm; 12 = 500 pm; 13 = 25 pm; 14 = 100 pm; 15 = 5()^pm. Abbreviations: C2,

C.3a, C3b, C4 = cardinal teeth; 14-1411 = lateral teeth I-llI; ilig = internal ligament; elig = external ligament.

D. C;. Zelava and C. Itiiartc', 2009

Patic

Page 4

THE NAUTILUS, Vol. 123, No. 1

Figures 16-20. Psciidokelh/a franki: anatomy. 16. Gross anatomy from the right side. 17-20. Transverse sections showing details of

demihrancli Insion and mantle border. 17. Posterior fusion of tlie inner demiliranchs. 18. Detail o( mantle folds and tentacle. 19. Fusion

ol posterior portion of the mantle border. 20. Anterior portion of tire mantle border. Scale bars: 16=1 mm; 17, 19, 20 = 300 pm; 18 =

1.50 pm. Abbreviations: aain = anterior adductor muscle; al = ascending lamella of inner demibraiich; bf = branchial fusion; dl =

descending lamella of inner demihrancli; e = embiyo; f = foot; id = inner demibraiich; imf = inner mantle fold; h = heel; mb = mantle

liorder; nimf = middle mantle fold; od = outer demihrancli; omf = outer mantle fold; paiii = posterior adductor muscle; I = tentacle.

in adults, present; byssns comprising a single, long fila-

ment. Outer and inner demibranelis present, witli

well -developed ascending and descending lamellae (Fig-

ure 16). Height of outer demibraiich representing one-

third the height ol inner one; posterior end ol outer

demibraiich (used to the mantle; left and right inner

demibranelis, also fused at posterior end, determining

defining a siiprabranchial chamber eontinuons with the

anal opening (Figure 17). Length ol descending lamella

of outer demibraiich about a half of ascending one;

filaments of ascending lamella of inner demibraiich

decreasing in length toward the posterior end.

D. (t. Zelaya and C. Itnarte, 2009

Page 5

Figures 21-26. Pseudokelhja franki: oocytes ami enibry'os. 21-24. Histological sections. 25, 26. SEM photomicrographs. 21. Pre-

vitellogenic and early vitellogenic oocytes. 22. Vitellogenic oocvte. 23-26. Einhi-vos attached to the inner deinihranch. Scale bars:

21, 24-26 = 100 pm; 22 = 50 pm; 23 = 200 pm. Abbre\’iations: aw = acinar wall; bf = brancliial lilaments ol the inner demibranch;

dg = digestive gland; e = embiyo; evo = early pre-vitellogenic oocvte; fe = follicle cell; g = gonad; h = heel: ifj = interfilamental

junction: nu = nnclens; pg = pedal gap; pvo = pre-\itellogenic ooc\te; st = stalk.

Reproductive Traits: Pseudokelhja frauki is dioe-

cious, brooding its einbmts within the inner denii-

Itranchs attached to the ascending Filaments by short

stalks (Figures 16, 23-26). The architecture ol oogenesis

sliows a peculiar Feature, cousistiiig of tire Formation oFa

complete oue-cell-thick Follicle surrounding each devel-

oping oocyte, whicii persi.sts to the end of vitellogenesis

(Figiu-es 21, 22, 24).

Tyiie Locality: Maxwell Bay, 62° 10-19' S, 58°35-5.S'

VV', King George Island, South Shetland Islands, 100 m

(station 30/49)'.

Tx'jjc Material: Holotvpe (ZMB 114680-a) and 10

paratxpes from the tvpe locality' (five at ZVIB 114680-b;

three at ML? 12997;' two at MACN-ln 37535).

Other Material Examined: 36 specimens. Maxwell

Bay, King George Island (station 30/43), 50 m (ZMB

114679); 84 .specimens. Maxwell Bay, King George

Island, 100 m (station 30/49) (ZMB 1 14680-0); 1 speci-

men, VlaxAvell Bay, King George* Island, 10-15 m

(station 11/134) (ZMB 114682); 6 specimens, Ma,xwell

Bay, King George Island, 40-50 m (station B 301)

(ZMB 114681).

Page 6

THE NAUTILUS, Vol. 123, No. 1

I'igures 27-38. Psciidoki'Uiia spcde.s. 27-29. P. rardifonitis (ZMB 63136). 30-32. P. inexpeciaiu (ZMB 114683). .3.3-.3I5.

P Amdahl. 33. .Spcdiiicii from 62° OO.OT S 60° 19.31' W' (ZSM 2()()41.320). 34, 3.5. Svnhpe (ZMB 63109). 36-38. P georoianu

(,M1,B 12999). 27, .30, 33, 36. Oiilcr viow Irit valve, and eletail ol .sliell .sciipOire at the n'glit .side. 28, 31, 34, 37. Hinge plate left

valve. 29, 32, .3.5, 38. I linge plate right valve. Seale Bars: 27, 30, 33, 36 = I nun; 28, 29, 31, 32, .34, 35, 37, 38 = 500 pin.

D. G. Zelaya and C. Ituarte, 2009

l^age 7

Distribution: Only known from South Slietland

Islands (Figure 1).

Etymology: The species is named alter Frank Kdhler,

Australian Museum, Sydney, and associated \rith the

Museum fiir Naturkunde, Berlin.

Remarks: In general shell outline, PseiidokeUi/a franki

is most similar to Pseuclokelli/a cardifonnis (Figures

27-29), from which it differs in having a relatively longer

and more straight posterior part of dorsal margin and a

ventral margin comparatively more strikingly cuiwetl. The

shell outline of T georgiana (Figures 36-38) differs from

that of Psciidokelhja franki in being markedly ti'apezoidal;

P. grodata (Figures 33-35) and P. inexpectata (Figures

30-32) have snbtrapezoidal shell outlines.

The absence of radial scidpture on the outer shell

surface is a distinctive character of Pseud okelh/a franki

(Figures 2-5). Also distinctive in P. franki is the pres-

ence of widely separated and low lamellate commarginal

periostracal folds (Figure 14); in other species of Pscu-

dokeJhja, the periostracum shows densely packed and

fine commarginal threads (Figures 27, 30, 33, 36). Pseu-

dokellija franki shows a weak commarginal sculpture

represented by low and rounded irregular ridges,

which seems to originate through growth disruptions

(Figure 15); in the remaining species this sculpture is

less evident, being represented only by shallow growth

lines (Figures 27, 30, 33, 36). In the case ok PseudokeUi/a

gradata, 3-4 shaqr growth disruptions, described as

“grades”, appear (Figure 33).

The hinge of the largest specimens of Pseiidokelh/a

franki is similar to that of the other species of the genus,

mainly differing in haring a larger cardinal tooth 3b and

a more solid cardinal tooth 2 with a triangular base. In

the smaller specimens the cardinal 3 is arched, with C3b

more developed.

Anatomically, Psciidokelhja franki differs from P. car-

difonnis (the only other species in the genus for whicli

anatomy is known) iii having only one defined mantle

opening, the anal, and a differentiated portion of middle

mantle fold below the anal opening that seems to repre-

sent the inhalant branchial aperture (lacking only a ven-

tral point of fusion delimiting the opening). Pelseneer

(1903) reported two posterior siphonal openings in

P. cardiformis. As it was described by Pelseneei' (1903)

for P. cardiformis, P. franki showed to be dioecious. Out

of the seven specimens histologically studied, four were

males and three females, with no signals of a possible

consecutive sexualitv detected.

TOWARD A BETTER DEFINITION OF

PSEUDOKELLYA

The generic definition of PseudokeUtja given by Pelse-

neer (1903) when describing P. cardiformis was based on

three characters: the presence of two posterior (branchi-

al and anal) siphonal openings, the dioecious condition,

and a peculiar mode of oogenesis comprising the

formation of a complete follicle surrounding each devel-

oping ooc)4e (a condition not known, at that times, lor

any other bix alve).

According to Thiele (1934: 858) the diagnostic char-

acters of Pseudokelli/a are: “shell roundish or somewhat

angular, nnifonuly bulging, with weak radial sculpture;

umbo moderately elevated, situated in the center: hinge

margin posterior to the ligament prolonged somewhat

ridge-shaped; anterior hinge teeth of the left \ alve fairly

long, diverging in an acute angle".

After the new infoi'iuation coming from the new spe-

cies described here and the species described after tlie

diagnosis by Thiele, an e.xpanded redescription of the

genus is needed.

Redescriplion of PseudokeUija: Shell small, shell

outline subcircular to snbtrapezoidal, ventral margin

uniformly cun'ed or more sharply cuiwed at pcisterior

half; beaks prosog)'rous, snbcentrally located. Prodisso-

conch sculptured witli microscopic granules; teleo-

conch usually sculptured \rith a \ariable number of

more or less marked radial cords, sometimes absent.

Periostracum usually elevated in fine threads or low

lamellate folds. Growth lines variably marked, some-

times looking like commarginal sculpture. Hinge plate

narrow, enlarged anterior to beaks, just at the cardinal

teeth insertion. Right valve \rith a hook-like cardinal

tooth (C3), formed by an anterior part (C3a) \ amng

from short and stout to long and slender, witli a trian-

gular base, bifid to a variable degree; and a smaller

posterior portion (C3b), sometimes extremely reduced

in size; a well-developed, elongated, narrow, and low

inner posterior lateral tooth (PI), and an outer poste-

rior lateral tooth (PHI) not well-separated from dorsal

margin. Left valve: two cardinal teeth, the anterior

(C4) running parallel to the anterior part of dorsal

margin, and the posterior (C2), usually elongated and

smaller, parallel or forming an acute angle with C4; a

single and elongated posterior lateral tootli (PII),

present. Resilifer small and shallow, located below

beaks. Internal and e.xternal ligaments, present. Man-

tle witli one or two posterior siphonal openings. Gills

each composed of two demibranchs. Foot witli a w'ell-

differentiated heel, having a small byssal gland. Ani-

mals dioecious, retaining the embiyos attached by

short stalks to the inner demibranch filaments; a

complete follicle surrounds each developing oocyte

throughout \itellogenesis.

O O

Coiiipari,son with Other Genera: When describing

PseudokeUija, Pelseneer (1903) focused in the pres-

ence of two siphonal (branchial and anal) openings

and the dioecious condition, in oposition to Kellia

wliich is a hermaphrodite and has only one posterior

siphonal opening, the anal. It is to be noted that, in

ccmtrast to that described by Pelseneer (1903) Pseudo-

keUija franki show's a single posterior mantle opening.

Additional characters differentiating Kellia from Pseu-

dokeUija are found in the hinge morphology: the

THE NAUTILUS, Vol. 123, No. 1

Page 8

former has a simple and triangular right cardinal tooth

C3 and two left cardinal teeth (C2 and C4) arranged

in a chevron pattern. Moreover, in Kellia both right

and left posterior lateral teeth are consistently stron-

ger than in Pseiiclokelh/a. In addition, KeUia only has

the internal ligament. Lastly, the radial sculpture and

periostracal folds present in some species of Pseudo-

kelh/a are absent in Kellia.

The hinge of Pseudokelh/a closely resembles that of

Ci/amiocarditnn Soot-Ryen, 1951, and Penieiina Ber-

nard, 1897, two genera also occurring in the Southern

Ocean. However, in the last two an additional tooth

behind the cardinal C2 (referred to as cardinal tooth

4b) appears (Lamy, 1917; Zelaya, 2008; pers.obs.). Adult

specimens of Ci/omiocardiiim and Penieiina have an

always well-developed C3b, which is reduced in size in

larger specimens of PseiidokeUi/a . Penieiina also has

tnlrercles anterior and posterior to the beaks resembling

a taxondont hinge, a character absent in members of

PsendokeUi/a (see Zelaya, 2008).

The Geographic Di.strihiition of Pseiidokellya:

Currently known species of PseudokeUt/a are restricted

to Sub-Antarctic and Antarctic waters. According to Dell

(1990), P. cardifonnis and P. gradata are probably ciren-

mantarctic, e.xtending to the Scotia Arc Islands and the

former, reaching Mahdnas and Kerguelen Islands. The

remaining three species are restricted to the Scotia Arc

islands: P. inexpectata known from South Georgia and

South Orkneys Islands (Dell, 1964), P. georgiana from

South Georgia (Dell, 1964), and P. franki from South

Shetland Islands (present study).

AGKNOWLEDGMENTS

DZ would like to express his gratitude to Matthias

Glaubrecht for his kind invitation for a short term stay at

ZAIB and all the staff of the museum for the facilities

and help provided during that stay; Enriko Schwabe and

Michael Schrodl kindly provided assistance during the

\4sit to the ZSM. The visits to the ZMB and ZSM

were possible thanks to a DAAD - GONIGET grant.

P. Mikkelsen and G. Goan provided valuable criticisms,

here acknowledged. The authors are members of the

Gonsejo Nacioual de Investigacion Gientihca y Tecno-

logica (GONIGET), Argentina. This paper was partly

funded by PIGT2005 38015 from ANPGyT.

LITERATURE GITED

Dell, R. K, 1964. Antarctic and sub-Antarctic Mollusca:

Amphineura, Scaphopoda and Bi\alvia. Discovery

Reports 33: 93-250, pis. 2-7.

Dell, R. K. 1990. Antarctic Mollusca wdth special reference to

the fauna of the Ross Sea. Bulletin of the Royal Society of

New Zealand 27: 1-311.

Lainy, E. 1917. ReHsion des Crassatellidae vivants du Museum

d’Histoire Naturelle de Paris. Journal de Conchyliologie

62(4); 197-270, pi. 6.

Pelseneer, P. 1903. Mollusques (Ainplrineures, Gastropodes et

Lamellihranches). Resultats du Voyage du S.Y. 'Belgica'. Expe-

dition Antarctique Beige 1897-1899, zoologie 3: 1-85, 9 pis.

Thiele, J. 1934. Handbuch der systeinatischen Weichtier-

kunde. Gustav Eischer Verlag, Jena. Bd 3, Dritter Teil,

pp. 119.3-1528.

Zelaya, D. G. 2008. Reallocation of Cyamiacardium crassilab-

niin Dell, 1964, into Penieiina Bernard, 1897 (Bivalvia:

Gyamiidae). The Nautilus 122: 52-55.

THE NAUTILUS 123(1):9-13, 2009

Page 9

Phylogenetic position of the bivalve family Cyrenoididae —

removal from (and further dismantling of) the superfamily

Lncinoidea

John D. Taylor

Emily A. Glover

Suzanne T. Williams

Department of Zoology

The Natural History Miiseuiir

London, SWT 5BD UNITF,D KINGDOM

[email protected]

[email protected]

s.\villiams@nhm. ac.uk

ABSTRACT

A molecular analysis using sequences from I8S and 28S rKNA

genes of the brackisli and freshwater bivalve Cijrenoida flori-

dana, in conjunction wdth a wide range of other heterodont

biwilves, demonstrated a close I'elationship with the families

Corbiculidae and Glauconomidae and distant from the laici-

noidea, wliere the Cyi'enoididae liad been usually classified.

Based on this result it is proposed that the Cyrenoididae be

removed from the Lucinoidea, which, for lixing taxa, now

includes only tire family Lucinidae.

Additional Kci/words: Bixalvia, fleterodonta,

Cijrenoida floridana, 18S rKNA, 28S rRNA

INTRODUCTION

Until recently, most ' classifications of bivalve mollusks

included within the snperfamily Lucinoidea several

component families (Lucinidae, Fimbriidae, Thyasiri-

dae, Ungulinidae Cyrenoididae, and fossil iVIactrom)-!-

dae) (e.g. Dali, 1901; Chavan, 1969; Boss, 1982, Vaught,

1989; Amler, 1999). The Lucinidae and some Thyasiri-

dae are notable for the chemosvmbiosis wdth sulphide-

oxidizing bacteria housetl in the ctenidia (Southward,

1986; Taylor and Glover, 2006). Molecular analyses of

tlie Lucinoidea, compared wdth a wdde range of other

heterodont bivalves, demonstrated that superfamily

was not monophyletic, wdth the Thyasiridae and Uugiili-

nidae not closely related to the Lucinidae (Williams, Tay-

lor, and Glover, 2004; Taylor, Williams, and Glover, 2007;

Taylor et ah, 2007). The Ungulinidae group near families

such as the Veueridae, Corbiculidae, and Mactridae,

while the Thyasiridae form a basal clade within the

Euheterodonta and are considered as a distinct

snperfamily — Thyasiroidea. Fimbria fimhriata, one of

the two living species of Fimbriidae, nested together

wdth Lucinidae species, wdth no support for separate

lamilial status, and the nominal family w'as syuonymized

accordingly. Apart from the Lucinidae, the only other

family wdth living species still classified wdthin Luciuoi-

dea is the Cvrenoididae, but lack of suitably presen’ed

material has precluded inclusion in molecular analyses.

From morphological exidence, Williams et ah (2004) and

Taylor and Glover (2006) thought a relationship to the

Lucinidae unlikely, Cijrenoida havdng medium to long

fused siphons, well developed labial palps and ctenidia

wdth two demibrauchs. These statements led Bieler and

Alikkelsen (2006) to place Cyrenoididae as incciiae scdis.

The Cyrenoididae Adams and Adams, 1857 (= Cyre-

nellidae Gray, 1853) comprise a small group of around

ten nominal species of little-studied bixalves inhabiting

brackish to freshwaters, classified into a single genus,

Cijrenoida (txq^e species C. diiponfia joannis, 1835)

(Figures 1-3), distributed in western Africa, eastern

and w^esteru Americas and some islands of the Garibbe-

an. The West African species inhabit brackish mangrove

habitats (Pilsbiy and Bequaert, 1927). In the eastern

USA, Cijrenoida floridana Dali, 1901 (Figures 4-7)

ranges from Dehuvare to the coast of the Gulf of

Mexico, maybe as far west as Yucatan (\"okes and Wkes,

1983), w'here it inhabits fresh and brackish water habi-

tats (Leathern, Kinuer, and Maurer, 1976; Kat, 1982;

Bishop and Hackney, 1987).

Dali (1895) was the first to place the Gyrenoididae

within the Lucinoidea, .stating (p. 545) “These are estua-

rine Lucinacea.” Later (Dali, 1901: 817) stated “. . .shells

ol this group with a Luciuoid animal and Diplodonta-

like shell, exhibit a hinge .structure w'hich is w'holly

distinct from any other of the Lucinacea.” Many later

classifications, inciudiug the iulluential Treatise of Inver-

tebrate Paleontology (Ghavau, 1969), lollow^ed Dali in

placing the Gyrenoididae within the Lucinoidea (e.g.

Yokes, 1980; Boss, 1982, Vaught, 1989, Skelton and Ben-

ton, 1993; Amler, 1999).

Page 10

THE NAUTILUS, Vol. 123, No. 1

Figures 1-7. Ci/reuoida (hipcnitid (1-3) and C. flohdana (4-7). 1. Cijrenoida dnpoiitia Joannis, left valve, Senegal (BMNH

2008 10.5.5). Scale bar = 10 nun. 2-3. Hinge teeth ol Ci/rcnoida dupontia. right \-alve (2) and left valve (3). Scale bar = 2 nun.

4. Cipvnoida floiidaiui Dali, left valve. Blue Hole, Big Pine Key, Florida (BMNH 200S1054). Scale bar = 2 niin, .5-6. Hinge teeth

(SEM images) ol Ci/rcnoida floridtma, right \alve (.5) and left \'alve (6) (BMNH 20081054). Scale Irar = 500 [.nn. 7. Ciirenoida

lloiidnna. liv ing specimen with short, fused sijrhons. Blue Hole, Big Pine Key, Florida. (Photo R. Bieler, September 2007).

Nevertheless, dillerent opinions were expressed hy

other authors, I'ischer (1887: 1096), for exainple, placed

Cyrenoididae (as Cyrenellidae) into a suborder Concha-

cea, near [oCorhicuIn and Ungnlinidae hut apart from the

Lncinoidea. W'hile Thiele (1934) included Cyrenoididae

with other Iresh and brackish water bivalves in the.s7/rp,s'

Sphaeriacea hut not positioned closely to laicinoidea. The

lainily was elevated to siiperlaniily status by Olsson ( 1961 :

227) hut placed near to Lncinoidea, a decision also lol-

lowed hy Keen (1971). The snperlaniily Cyrenoidoidea

was also recognized hy Nevesskaya et al. (1971) and

placed along with Lncinoidea in the order Astartida.

Alternatively, and rather hizarrely, Starohogatov (1992)

placed Cyrenoidoidea within the infraorder Eiycinoinei

along witli Cyainioidea, Galeonnnatoidea and Leptonoi-

tlea, all contained within the order Lncinifornies.

Clearly, there e.xists ninch uncertainty concerning the

pliylogenetic position of Cyrenoididae amongst the het-

erodont bivalves hut this has never been tested hy either

morphological or molecular analyses. In 2007, we

obtained samples of Ci/reiioida floridana suitable for

molecular analysis and in this paper we present 18S and

J. D. Taylor et al„ 2009

Page 11

28S rRNA sequences for the species that enable ns to

establisli the phylogenetic position ol the lainily in rela-

tion to a wade range ol heterodont bivalve taxa previous-

ly analysed (Taylor et ah, 2007) and specilically address

the question of whether the Lncinidae and Cyrenoidi-

dae form a monophyletic group.

MATERIALS AND AIETHODS

The sample ol Cifrenoida floriclaiia, presen-ed in 100%

ethanol (BAINH 20081053), wars collected (18 Septem-

ber 2007) from Bine Hole (24°42.4' N, 81°22.8' W) a

freshwater pond on Big Pine Key, Monroe County, Elor-

ida Keys, Elorida, USA, from slioreline mnd up to 0.5 m

depth among roots of marginal reeds. Other material

from the same site is lodged at the Eield Mnsenm ol

Natural History (EMNH 314434; 317667).

Eor tlie molecular analysis, methods of DNA e.xtrac-

tion, amplification anti sequencing followed by sequence

analysis and phylogenetic reconstruction are as

described in Taylor et al. (2007). Sequences for Ci/re-

noicla florichina were analysed together wath the data set

of heterodonts listed in Taylor et al. (2007, Table 1), with

the addition of new 18S and 28S sequences for Mi/a

arenaria Linnaeus, 1758 (family Myidae) from Gydnia,

Poland. The new^ setjnences for Cifrenoida floridana anti

Mija arenaria are lodged in GenBank (Accession nnm-

bers: C. floridana F’M999789, EAI999790; M. arenaria

EM999791, 779792). Voucher specimens t)l both species

are housed in the Department of Zoology, The Natural

History Museum, Lt)udon.

Phylogenies were constructetl using Bayesian meth-

ods (MrBayes v3.1.2, Huelsenbeck and Rompiist, 2001)

using a GTR+G+I mt)tlel. The analysis for each data set

was run for 3,500,000 generations, with a sample fre-

tjuency of 100. Each analysis wais run twice. The first

15,000 trees from each run were tliscartletl so that the

final cotisensus tree was based ttn the ettmbination of

accepted trees from each run (a total of 40,000 trees).

Support for nodes w'as determinetl using pttstei'ior prt)b-

abilities (PP, calculatetl by MrBayes).

RESULTS

The combined tree based on concatenated sequences

from 18S rRNA and 28S rRNA genes is showm in

Eigure 8. The individual trees based on single genes ai'e

very similar in topolog)' to those published previously

(Taylor et ak, 2007). In all analyses Cifrenoida floridana

nests in a highly supported clade wath Corbiciila fliiininea

(Corbiculidae) and Claiiconoine virens (Glauconomidae).

This clade forms part of a major group of heterodonts

named Neoheterodontei by Taylor et al. (2007). Cifre-

noida is widely separated from both Thyasiridae and

Lncinidae that appear in the more basal parts of the

tree. The Ungnlinidae, althongh also a member of the

Neoheterodontei, lorm a separate clade distinct from

Cifrenoida.

DISCUSSION

It would have been desirable to have included the t\pe

species of Cifrenoida. namely, Cifrenoida diipontia [oan-

nis, 1835, from West Africa, in the molecular analysis

but no suitably presei'ved material wais a\’ailable. Al-

though a much smaller species, C. floridana is similar

to C. diipontia in shell characters, notably the unusual

hinge dentition, and we feel confident that they are

members of the same group, Cifrenoida dii))onlia has

three cardinal teeth in the riglit valve, the anterior of

these is tliin and elongate and tlie central tooth larger

and sliglitly bifid (Eigures 2-3) wdiile the left \ alve has

two cardinals, the posterior tooth smaller and bilid and

the anterior tooth elongate. Lateral teeth are absent. The

dentition of C. floridana is veiw similar (Eigures 4-6)

w'ith three cardinal teeth in the right valve, the central

being larger and two caixlinal teetli in the left wilve watli

the posterior tooth Ihfid and the anterior tooth elongate.

We have also e.xamined the gross anatomy of Cifrenoida

rosea (d'Aillv, 1896) from Nigeria (National Mnsenm of

Wales specimen NMW.Z.2()03.029.()2()4f ) and this has

ctenidia with hvo demibranchs, with the inner demi-

branch larger, paired triangular labial palps, and ftised

medinm-length posterior siphons. Cifrenoida floridana

is similar, wath small outer demibranchs, triangular labial

palps and short I used posterior siphons, the inl ialant with

papillae (Eigure 7), Despite the presence of siphons,

there is no pallial sinus in any Cifrenoida .species.

The inain conclusion ol this study, based on our results

lor Cifrenoida floridana, is that the Cyreuoididae sliould

be removed h'om the Lucinoidea and classified close to,

or possibly wathin, the Corbiculoidea. The status ol Cifre-

noida in relation to Corbiculidae and Glauconomidae

needs further analysis witli a larger dataset of corbicnlid

species. Eor the present the family can be classified wdth-

in a separate superfamily Cyrenoidoidea as proposetl by

Olsson (1961). Molecular evidence for a liighlv sup-

ported relationship betw^eeu Corbiculidae and (ilauco-

nomidae was reported by Taylor et al. (2007) although

the elongate shells with deep pallial sinus ami long

siphons of Glaneonoine are less similar morphologically

to Cyrenoididae and Corbiculidae. Species of Cyreuoidi-

dae and Corbiculidae occur in botli brackish and Iresh-

water habitats while Glauconomidae live intertidally

among mangroves in em ironments ol lluctuating salinity.

Eor living taxa, w^e consider that tlie superfamily Imci-

uoidea should uow' include only the lamily Lncinidae,

wath the families Thyasiridae, Ungnlinidae and Cyreuoi-

didae excluded. The position ol the entirely fossil

families Mactromyadae, llionidae, ami Paracyclidae is

unresolved althongh the latter two embrace species with

Incinid characters.

AC K N O WEE 1 ) C M E NTS

It is pleasure to tliank Riidiger Bieler and Petra Sienvald

for generous hospitality and help iu collection ol

Page 12

THE NAUTILUS, Vol. 123, No. 1

100 I

I

Neotngonia lamarcku TRiGONllDAE

Margantifera margantifera MARGARITIFERIDAE

Unopictorum (JNIONIOAE

OUTGROUPS

CRASSATELLIOAE

Anodonta cygnea

Card>ta leana CARDITIDAE

As/arte sulcata ASTARTIDAE

Eucrassatelia donaana

Eucrassatella cuinmgn

TOO I' ■ ■' Thyastra polygona

THVASIRIDAE

Mendicula /erruginosa

CRASSATELLOIDEA

Poromya iltevis POROMYIDAE

ANOMALODESMATA

LUCINOIDEA

Lunuhcardium hemicardium

CAROIIDAE

CARDIOIDEA

NEOHETERODONTEI

Chama senvpurpurata

CHAMiOAE

Chama asparsa

CHAMOIDEA

Figure 8. Molecular phylogeny of heterodont hivalves including Cijrenoida produced by Bayesian analysis for concatenated

se(jiiences from 18S and 28S rHNA genes. The tree was drawn using members ol the palaeoheterodonts Trigoniidae, Unionidae,

and Margaritileridae as outgroups. Support values are posterior probabilities. Nodes with <50% support have been collapsed.

Positions of Lucinoidea, Thyasiroidea, and Ungulinoidea marked by grey bars. Details of t;ixa in Taylor et al. (2007).

C. finridaua. We are grateful to Pat Dyal for assistance

with molecular analysis and to Graham Oliver (National

IVlnsemn ol Wales) for making a specimen of Ctireiwida

rosea available. We thank Professor Idiil Rainbow and

Department of Zoology, NIIM for continuing support.

Research on this exemplar species in the Bivalve Tree-of-

Life project (www.bivatol.org <http://www.bivatol.org/>)

was supported by the U.S. National Science Foundation

AToL program (DEB-()732854/()7329()3/0732860). The

specimens were collected under U.S. Fi.sh and Wildlife

J. D. T\iylor et al„ 2009

Page 13

Senice Special Use Permit 41580-2007-11 (to lliidiger

Bieler) for reseai'ch on native mollnscan species in the

National Key Deer Refuge.

LITERATURE CITED

Adams, H. and A. Adams. 1S.53-5S. The genera of recent

Mollusca, arranged according to tlieir organization, van

Hoorst. London, 3 volumes. \Alume 2. Cvrenoididae.

p. 452 (published IS57).

d'Ailly, A. 1896. Contributions a la connaissance des mollus-

fjues terrestres et d’eaii douce de Kameroim. Bihang Till

Kongliga Svenska Vetenskaps-Akademieris Handlingar 22:

1-137.

Amler, M.R.W. 1999. Svmoptical classification of fossil and Re-

cent Bi\ alvia. Geologica et Palaeontologica 33: 237-248.

Rieler, R. and P.M. Mikkelsen. 2006. Bi\alvia — a look at tlie

branches. Zoological lonrnal ol the Linnean SocieR 148:

223-235.

Bishop, T.D. ami C.T. Hackney. 1987. A comparative study ol

the mollusc communities of two oligohaline intertidal

marshes: spatial and temporal distribution of abundance

and biomass. Estuaries 10: 141-152.

Boss, K.J. 1982. Mollusca. In: Parker, S.P. (Ed,). Svnopsis and

classification of living organisms. McGraw-Hill, New

York, pp. 945-1166,

Dali, VV.H. 1895. Contribution to the Tertiaiv fauna of Elorida,

with especial reference to the Miocene silex-beds of

Tampa and the Pliocene beds of tlie Caloosahatchee Riv-

er. Part HI. A new classification of the Pelecvpoda.

VVagnei' Eree Institute of Science of Philadelphia Transac-

tions 3(3): 479-570.

Dali, W.H. 1901. S)mopsis of tlie Lucinacea and of the Ameri-

can species. Proceedings of the U.S. National Museum

23: 779-833.

Chavan, A. 1969. Superlamily Lucinacea Eleming, 1828. pp.

N491-N518. IN Moore, R. C. [ed.]. Treatise on Inverte-

brate Paleontology', Part N, Mollusca 6, Rix ah ia, (2). Geo-

logical Society of America, Rouldei', and Universit)' of

Kansas Press, Lawrence, 951 pp.

Eischer, P.H. 1880-87. Manuel de conchyliologie et de paleon-

tologie conchyliologique. E. Sav)', Paris. 1369 pp.

Gray, J.E. 1853. A revision of the genera of some of the

families of Conchifera or Bivalve shells. Annals and Mag-

azine of Natural History, series 2, 11: 33-44.

Huelsenbeck, J.P. and Ronquist, F. 2001. MRBAYES: Bayesian

inference of phylogenetic trees. Bioinformatics 17: 754-

755.

joannis, L. de. 1835. Olisen ations surplusiers Mollusijues dont

on ne connaissait que les coquilles, et descriptions des ces

mollusques, et de quelques especes inedites on non figur-

ees jusqu’a ce jour. Magasin de Zoologie 5: pis, 36-64.

Kat, P.W. 1982. Reproduction in a peripheral population of

Ci/reiioicia florichma (Bixalvia: Cvrenoididae). Malacolo-

gia 23: 47-.54.

Keen, A.M. 1971. Sea Shells of IVopical West America: Marine

Mollusks from Baja California to Peru. 2'"' edition. Stan-

ford University Press, Stanford, 1064 pp.

Leathaem, P. Kinner, and D. Maurer. 1976. Northern

range extension of the Florida marsh clam Ci/rcnnida

floriclana (Superfamily Cyrenoidacea). The Nautilus 90:

'9,3-94.

Nevesskaya, L.A., O.A. Scai'lato, Y. I. Starobogatov, and A. (7

Eberzon. 1971, New ideas on bivalve systematics. Paleon-

tological {ournal, 1971 (2), 141-155. Translation from

Russian of Paleontologiclieskii Zliurnal 1971 (2): 3-20.

Olsson, A. A. 1961. Mollusks ol the tropical eastern Pacific

particularly Irom the southern lialf of the Panamic-

Pacific faunal province (Panama to Peru). Panamic Pacific

Pelec\poda. Paleontological Research Institution, Ithaca,

574 pp,

Pilsbn', II. A. and J. Bequaert. 1927. The a(|uatic mollusks of

the Belgian Congo with a geographical and ecological

account of Congo malacologx’. Bulletin of the American

Museum of Natural I Ii.ston’ 53: 69-659.

Skelton, P.M'. and M.j. Benton, 1993. .Mollusca: Rostroconchia,

Scaphopoda and Bivahla. In: Benton, M.J. (ed.) The Fos-

sil Record 2. Chapman & Hall, London, pp. 237-263.

Southward, E.C. 1986. Gill symbionts in thyasirids and other

bivalve molluscs. Journal of the Marine Biological Associ-

ation of the United Kingdom 66: 889-914.

Starobogatov Y.I. 1992. Morphological basis for phylogeny and

classificati(m of BhaKia. Ruthenica 2: 1-25.

Taylor, J.D. and E.A. Clover, 2006. Lucinidae (BivaK’ia) — the

most di\ erse group of chemosvmbiotic molluscs. Zoologi-

cal Journal of the Linnean SocieR- 148: 421—438.

Tavlor, J.D, S.T. W'illiams, and E.A. ClOvei'. 2007. Evolutionaiv

relationships ol tlie bixalve family Thyasiridae (.Mollusca:

Bivahla), monophyly and superfamily status. Journal of

the Marine Biological Association of the United Kingdom

87: 565-574,

Taylor, J.D., S.T. M'illiams, E.A. C4o\er, and P. Dyal. 2007.

A molecular phylogeny of heterodont bivalves (Mollusca:

Bivalvia: Heterodonta): new analyses of 18S rRNA and

28S rRNA genes. Zoologica Scripta 36: 587-606.

Thiele, J. 19.34. Handbuch der systematischen Weichtierkunde

2 (3) (Scaphopoda. Bivalvia. Cephalopoda). Fischer, Jena,

pp. 779-1022.

\'aught, K.C. 1989. A classification of the* living Mollusca.

American Malacologists, Melbourne, Florida, 196 jrp.

Yokes, H.E. 1980, Catalogue of the BiMiKla: a systematic and

bibliographic catalogue (revised and updated). Paleonto-

logical Research Institution. Ithaca, 307 pp.

\'okes, H.E. and E.H. Yokes, 1983. Distribution of shallow-

water marine Mollusca, Yucatan Peninsula, Mexico.

Tulane UniversiR Mesoamerican Eccdogical Institute,

Monograph 1: l-i83.

Williams, S.T., J.D. Taylor, and E.A. Glover. 2004. Molecular

phylogeny of the Laicinoidea (Bivalvia): non-monophyly

and separate acquisition of bacterial chemosymbiosis.

Journal of Molluscan Studies 70: 187-202.

THE NAUTILUS 1T3(1):14-18, 2()09

Page 14

Establishment and persistence of the copse snail, Arianta

arbiistonim (Linnaeus, 1758) (Gastropoda: Helicidae) in Canada

Donald F. McAlpine

New Brunswdck Museum 277 Douglas

Avenue, Saiut |ohu New Brunswick,

CANADA E2K 1E5

[email protected]

Ronald G. Noseworthy

Yale Academy 441-3 Seohong-Dong,

Sogwipo jejudo 697-080, REPUBLIC

OE KOREA

Frederick W. Sehueler

Bishops Mills Natural Histoiy Centre

RR # 2, Oxiord Station Ontario,

CANADA KOG ITO

Maiy C. Sollows

New BrunsvMck Museum 277 Douglas

Avenue, Saint John New Brunswick,

CANADA E2K 1E5

John E. Maunder

P.O. Box 250, Pouch Cove

Newfoundland and Labrador,

CANADA AOK 3L0

ABSTRACT

Although recorded from Newfoundland in 1885, by the late

1930s the copse snail, Arianta arhnstonim (Linnaeus, 1758),

was believed to no longer be extant in North America, We

in\’estigated sites in Newtoundland, New Brunswick, and

Ontario, Canada and found that A. arbustonun is well estab-

lished in these provinces; extant populations have persisted in

Newfoundland for at least the past 30 years and in Ontario for

more than 50 years, \\4iere present in Canada, the species may

sometimes be abundant, altlurugh populations are quite local,

sometimes occupying less than 1 ha. Canadian Food Inspec-

tion .Agency records show A. arbustonun to have been inter-

cepted 7'egularly (0-3 interceptious/year) since record-keeping

starterl in 1963. Interceptions have occurred in 7 pro\inces

spanning the countiy trom Nova Scotia to British Columbia,

Nurseiy stock oiigiuating in the Netherlands appears to be the

main vector, but preliminaw molecular data from Newfound-

land populations suggests multiple European points ot origin.

Additional Kei/ivords: Introduced species, invasive species,

Newfoundland, New Brunswack, Ontario

INTRODUCTION

The cop.se snail, Arianta arbnstortmi (Figure 1), (X'curs

coinnionly across northwestern and central Europe

(Kerney and Cameron, 1979). Despite this wide.spread

European distrihntion, and the signilicant nnmher ot

European inollnscan taxa now found in North America

(Robinson, 1999), A. arhiistonnti has apparently never

become established in the United States ami has only

rarely liecome established in Canada. Dundee (1974),

il ia list of introduced mollnsks of eastern Nortli America,

noted interceptions ol A. arbnslonim by the US Depart-

ment ol Agriculture at ports in six eastern states, but

neither Mead (1971) nor Dundee (1974) reported any

established populations. Until recently, the only pub-

lished North American record of A. arbiistonim is an

1885 obseiwation from St. John s Newfoundland, reported

by Whiteaves (1904), Grimm (1996) mentioned the oc-

currence of one colony in a ravine in Toronto, Ontario, Imt

provided no details. Here we document the occurrence of

A. arbiistonim in Newfoundland, New Brunswick and

Ontario and confirm the persistence or reintroduction of

this species in Ontario and Newfoundland. We also re\4ew

records of non-native plant pests intercepted by federal

authorities from across Canada and show that A. arbiis-

tonim has been regularly imported into the countiy for

more than 40 years.

MATERIALS AND METHODS

Following discovery of A. arbiistonim in western New-

foundland by RGN in 1970 and its subsequent discovery

in New Brunswack in 2004, we accumulated data in each

region in order to delimit local distribution. We also

examined specimens deposited Ity the late F. W. Grimm

in the Canadian Museum of Nature and consulted his

unpuJ'ilished field notes at the Bishops Alills Natural

Histoiy Centre, for reference to Ontario occurrences.

These notes plus additional occurrence information

from entomologist D. Monty Wood, one of Grimm’s

correspondents, led DEM and FWS to make confirma-

toiy searches of ravine sites in Toronto. In addition,

Canadian Food Inspection Agency records, maintained

since 1963, were examined and all interceptions of

A. arbiistonim noted, along with countiy of origin, num-

ber of snails intercepted, and plant host. Voucher mate-

rial of A. arbiistonim collectetl during this study is

deposited in the collections of the Bishops Mills Natural

Histoiy Centre (EOBM), the New' Brunswick Museum

(NBM), and the Provincial Museum of Newfoundland

and Labrador (NFM).

D. F. McAlpine et al„ 2009

Page 15

Figure 1. Live Aiicmta arbustonnn, central Saint John, New

Bninswlck, May 2007. Scale bar = 1 cm. Photo M. Sollows, 2007.

RESULTS

Ontario: Collections data (Canadian Museum of

Nature 059910 and Field Museum of Natural History

267S29) provide more details on the Ontario popula-

tion reported by Grimm (1996); material was collect-

ed from the Lawi'euce Park School comple.x on the

north slope of Chatsworth Ravine, Toronto, 14-15

October 1970 by FW. Grimm and |. Cavanaugh

(43.720° N, 79.406° W; Figure 2a). Grimm’s field

notes also indicate material was collected by D.M.

Wood from Rosedale Ravine, alront 6 km away,

around 1950. Unfortunately, vouchers from this site

no longer exist. However, Wood recently stated that

his material was collected opposite Parliament Street

on the north side ol’ the ravine at 7 Dale Avenue

(43.673° N, 79.372° W, pers. comm, to DFM).

Searches of the Chatsworth Ravine by FWS (May

2006) and DFM (September 2007) revealed that the

population is still extant, at least on the basis of

numerous fresh-dead shells; however, A. arhusfonim

(EORM 1667; NBM 367; Eigure 3) were uucommon

relative to the co-occurring Ccpaea nemoralis (Lin-

naeus, 1758). Dead shells were restricted to an area

of ~L5 h wuthin the Chatsworth Ravine. Searches of

the Rosedale Ravune by DFM and FWS in August

and September 2007 revealed no A. arbustonnn, al-

though urban development and gated and fenced

properties precluded our access to some areas.

New Brunswick: lu 2004, DFM and MCS found a

well established population of A. arbustonnn in central

Saint John (Site 2, Figure 2b; 45.270" N, 66.078" W;

NBM 143, 8602). Collections were snlrsequently made

elsewdiere in the ciU (Greenhead, 45.267" N, 66.133° M7

NBM 136; Figure 4; west Saiut John, 45.249° N,

66.062° W; NB"m 315). Locally, the species is abundant.

Collection sites are 0.3-5.25 km apart and collectiv^ely

encompass an area of ~20 h straddling the St. John

River (Figure 2b).

Newfoundland: In 1970, RGN collected A. arlms-

tonim from an urban garden close to the trans-island

railway corridor on Chapel Hill (road). Deer Lake, in

western Nevvdonndland (Figure 2a; 49.16° N, 57.43° W;

NFM MO-1971, 1972; [all lat/lougs reported here are

consistent \\4th NAD83]) and from close to the Deer

Lake Airport (49.1917° N, 57.4083° W; NFM MO-

1970), but JEM was unable to relocate either of these

populations in 2006. In 1976, RGN collected A. arbus-

torum at Pettv’ Harbonr-AIaddox C(we (northern site)

near St. John’s (Figure 2c; 47.4853° N 52.7049° W;

NFM M6-1973). In 1984, RGN and JEM again collect-

ed A. arbustonim from this locality (NFM MO-389).

Subsequent obseiwations and collections of the species

in the general St. John’s area by JEM and RGN beRv'een

1986 and 2007 include: Three Island Pond between

Torbay and Bauline (47.6767° N, 52.7778° W'), just east

of Lundrigau’s Marsh (47.6031" N, 52.6813" W), Kent’s

Pond (47.5864° N, 52.7242° W), a hydro pole-line near

Oxen Pond Road (47.5825° N, 52.7535° W), Masonic

Terrace (47.5656° N, 52.7072° W), Syme’s Bridge

(47.5433° N, 52.7244° W; Figure 5), Bowring Park

(northern site) (47.5279" N, 52.7447" W), Bowning Park

(southern site) (47.5222° N, 52.7547° W; NFM MO-684,

1370, 1391) and Pettv Harbonr-Maddox Cove (southern

site)(47.4682° N, 52.7067° W).

The total Nevvdonndland population of A.arbustrorum

presently occupies ten small localities (collectiv'ely cov-

ering ~1 ha; Figure 2c) vvutliin a narrow area of alrout

23.5 X 6 km radiating north-south from the original

1885 discover site at the entrance to St. |ohn’s Harbour.

Canada Food Inspection Agency Interceptions:

Table 1 summarizes records of A. arbustonnn inter-

cepted fry the Canadian Food Inspection Agency at Ca-

nadian inspection stations since 1963. The species has

been reported on 26 occasions (range of 0-3 intercep-

tions/year) at stations in Nova Scotia, New Bi'imsvvdck,

Quebec, Ontario, Saskatchewan, Alberta, and British

Columbia, usually in association vvdth a vnudety' of garden

plants imported from the Netherlands (73%), or, less

commonly, other European countries (23%).

DISCUSSION

Whiteaves (1904) reported A. arbustonnn from “grassy

sk)pes facing the sea near tlie nai'rovv/s of St. Johns Har-

bour, Newfoundland" in mid-July 1885. Hovv'ev'er,

Brooks (1936) and Brooks and Brooks (1940) noted that

their searches of the area around St. John’s in 1934 did

not reveal the species. Pilsbry (1939) suggested that the

Newfoundlaud population might no longer persist and

deemed the species a "rather doubtful member of the

American fauna’’. The Pettv Harbonr-Maddox Cove

population was v^eiy small vvdien discovered in 1976, and

does not appear to have spi'ead much since. The Pettv

Harbour-Maddox Cov^e localities are physically sepa-

rated from the eight remaining sites by tlie steeply rising

200+ m north-sotith trending Sonthside Hills. Likewise,

the Bowring Park (southern site) population vwis also

veiy small w4ien it was discovered in 1986, restricted to

the grounds of an (4d estate. Ilow^ever, it appears to have

Page 16

THE NAUTILUS, Vol. 123, No. 1

Figure 2. a. Ea.stern North America showaiig sites (•) in Ontario, New Brunswick and Newfoundland where Arkinto arhustonnn

has lieen reported, b. Saint John, New Bnmswack wdth sites of occurrence for A. arbustonim: 1. Greenhead; 2. Saint John central;

.3. Saint John west. c. St. John’s, Newfoundland and environs whtli sites for A. arbiistonnn: 1. Three Island Pond; 2. Lnndrigan’s

.Marsh; .3. Kent’s Pond; 4. Oxen Pond; 5. Masonic Terrace; 6. Svmes’s Bridge; 7. Bowring Park north; (S. Bowring Park south; 9. Pettv'

Ilarbonr-Maddox Cove north; 10. Pett)' Ilarbonr-Maddox Cove south.

spread significantly during recent years and is probably

tJie sonrce of the eight occurrences now knowai in the

main St. |ohn’s area. Preliininaiy molecular data suggest

that the present-day Newfoundland populations are

derived from at least two separate introductions from

Europe; the Petty Ilarbonr-Maddox Cove populations

being genetically distinct from the greater St. John’s area

populations (A. Grindon, Nottingham University, pers.

comm, to DEVI). The Deei' Lake records may represent

ephemeral populations derived from snails transported

Irom St. John’s in rail cargo, since lioth localities are

located along the formei' trans-island railway corridor.

Although the area occupied by A. arl)ustorum in New

Brnn.swick suggests a long-standing population, it is not

possilde to estimate a likely date of introduction for

A. arhustonim to Saint John. Matthew and Stead (1903)

made no mention ol the species in their list of land and

Iresliwater mollusks collected in and near Saint John

about 1890-1 900. Unfortunately, the mollnsk sniwey of

Coleman (1966) conducted iu Saint John is incomplete,

even for the marine and freshwater species sampled.

An undated collection record in the Eield Mnsenin of

Natural Ilistoiy (FMNII 38439) reports a single diy

shell from “Selkirk, New Bruuswdck”. A search of recent

and histoi'ical gazetteers reveals no such location in that

proxdnce. The specimen and original label appear to be

missing. The specimen was originally in the collection of

G.K. Glide, a malacologist resident in the United King-

dom, who was active in the early 20^'’ Centuiy. Gude

produced ver)/ small labels and it seems iphte likely that

he would have abbrevdated liis label data (J. Gerber,

pers. comm, to DEM). We suggest that this record as

reported is the result of an error in transcription. While

it may refer to Selkirk, Manitoba (MB), rather than New

Brunswick (NB), the record may not even be North

American. Gude undoubtedly exchanged widely; howev-

er his research interests focused on regions outside the

North American continent, and there is no material

from the Gnde Gollectiou now in the Field Museum

labeled as being from Alauitoba.

Gonsidering that A. arhustonnn is widely distributed

ami common iu Europe, and appears to be imported not

D. F. McAlpine et al„ 2009

Pa^e 17

Figures 3—5. Representative specimens oi Arianta (ii'hiisfonim troni .3. Ontario (NRM StSfi), 4- New Brunswick (NHM 355), and

5. Newtonndland (NBM 8355). Scale Bar = 2 cm.

inlrequently into Canada, it is .surprising that the species

has not lieen recorded as more widely established in

temperate regions of Nortli America. Robinson (1999)

listed the .species as an nncommonly imported invasive,

accounting lor <0.1 % ol more than 4,900 US intercep-

tions over about a b-year period. Although this percent-

age is small, it still accounts for a sizable nnniber ol

animals. In Canada, <5% ol tropical plants from the

United States are examined but 100 % of off-continent

nurseiv stock is inspected (D. Parker, pers. comm, to

DFM). One interception at Edmonton, Alberta, in

1999-2000 originated at Coulds, South Florida. As there

are no vouchers for this interception the identification

cannot be confirmed. However, given the species re-

striction to north-temperate latitudes in Europe, estab-

lishment of A. ar])ustonun in Florida seems unlikely.

In snmmar'y, A. arhiisloriim may have persisted for

more than a centniw on Newdbnndland or may have

been repeatedly introduced; preliminan' exidcmce indi-

cates multiple introductions. It has also been present in

New Brunswick and Ontario for some time. F’nrther

investigation may reveal that this European alien is more

widely distributed at temperate latitudes in Nortli Amer-

ica than was pre\ ionsly thought.

ACKNOWLEDCiMENTS

W'e thank D. Parker, Head, Identification and Kc'gulatoiA'

Entomology, Canada Food Inspection Agencx; Ottawa,

lor making records of intercepted plant pests available

to us. D.M. Wood for sharing his knowk'dge ol the

Hosedale Ravine jiopnlation with us. N. Djan-Chekar,

Proxincial Mnsc'um of Newionndland and Labrador,

|.-M. Cagnon, Canadian Mnsemn ol Nature, Ottawa,

and |. Gerber, Field Mnsemn of Natural Historv,

Cliicago, for access to collections data and rc-sponded

to our enquiries. A. Grindon, Nottingham University

for preliininaiy molecnlar data. R. ForsN-tli drew our

attention to the Selkirk record in tlie Field Museum of

Natunrl Histoiy.

LITERATURE CITED

Brooks, S. T. 1936. The land and freshwater niollusca of

Newfoundland. Annals of the Carnegie Museum 25: 83-

108.

Brooks, S. T. anti B. W. Brooks. 1940. Geographical distribu-

tion of the recent niollusca of Newfoundland. Annals of

the Carnegie Museum 28: 53-65.

Coleman, R. VV. 1966, Certain mollusks of the environs of

St. lohn. New Brunsxrick, Canada. Iowa Academy of Sci-

ence 73: 405-407.

Dundee, D. S. 1974. Catalog of introduced molluscs of

eastern Nortli America (north of Mexico). Sterkiana 55:

1-37.

Grimm, F. W. 1996. Terrestrial Molluscs. In: I. M. Smith (ed.)

Biodiversity Assessment of the MLxedwood Plain Ecozone,

Agriculture Canada, Ottawa http://www. naturewatch.ca/

niLxedwoocFlandsnaiy. Cited 27 Fehruaiy 2006

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

Land Snails of Britain and Northwest Europe. William

Collins & Sons, London, 288 pp.

Matthew, W. D. and G. Stead (1903) Land and freshwater

shells collected near St. John, N.B. Proceedings of the

Miramichi Natural Histoiy Association 3: 48-49.

Mead, A. R. 1971. Helicid land mollusks introduced into

North America. Biologist 53: 104-111.

Pilshiy, H. A. 1939, Land Mollusca of North America (north of

Me.xico), 1(1). Monographs of the Academy of Natural

Sciences of Philadelphia 3: 1-573.

Robinson, D. G. 1999. Alien invasions: the effects of the global

economy on non-marine gastropod introductions into the

United States. Malacologia 41: 413-438.

Whiteaves, J. F. 1904. Helicigona arbustonun in Newfound-

land. Ottawa Naturalist 17: 192.

Rediscoveiy of a Caribbean living fossil: Fholadomija Candida

G.B. Sowerby I, 1823 (Bivalvia: Anomalodesmata:

Pholadoinyoidea)

Juan M. Diaz

Fernando Cast

Institute) (le Investigacion de Recursos

Biologicos "Alexander von Ilninboldt"

Apartado 8693, Bogota D.C., COLOMBIA

[email protected]. CO

Diana C. Tor res

Uuiversidad de Bogota “Jorge Tadeo Lozano”

Programa de Biologia Marina

Pidilicio .Mundo Marino, El Rodadero

Santa Marta, COLOMBIA

PhoJadomifa Candida G.B. Sowerby I, 1823, is an anom-

alodesmatan bivalve belonging to the ancient family Pho-

ladoinyidae, a group of bnrrovMng bivalves with a wide

palaeobiogeographic distribution from the Carboniferous

to the Recent (Cox, 1969). This group is characterized by

haCng posteriorly elongated shells \Cth strong radial

ribbing, a siphonal gape, a hinge with no functional teeth,

and an external opisthodetic ligament (Runnegar, 1974).

Although several recent species from different regions

of the world have been descrilied undei' Pholadonu/a,

the h'loe species, P. Candida, is the only Recent species

resembling a great number of fossil forms in size, shape,

and life habits (Cox, 1969; Waterhouse, 1969; Lazo,

2007). Accordingly, all other extant members of the Pho-

ladomyoidea are more correctly placed in the genera

Parilimi/a Melvill and Standeri, 1899 or Panacea Dali,

1905 (Cox, 1969; Runnegar, 1972; Zinsmeister, 1978;

Morton, 1980; Lazo, 2007). It seems then that P. Candida

is the only remaining species of a genus that flourished

for more than 200 million years f rom the Late Triassic to

the Recent, and this is why Runnegar (1972, 1979) and

Alorton (1980) have referred to it as a “living fossil.”

Records of Pholadonn/a Candida are extremely scarce,

mostly from the West Indies. Since living specimens

had not been found since the latter part of the 19^''

Centiny, the species was considered as possibly extinct

(Runnegar, 1972; Alorton, 1980). Ikwever, discoveries

of fresh-looking shells from Venezuela (Gibson-Smith

and Gibson-Smith, 1980) and Colombia (Diaz and

Borrero, 1995) provided evidence that it still may be

living, at least in the southern Caribbean.

Pholadanu/a Candida had been collected alive

only twice, at least in the sense of being available for

scientific studies (Alorton, 1980); both specimens

were found before 1842 in tlie same area, the Virgin

Islands. One of them was dissected by R. Owen in

1839, but some of liis illustrations were lost and

his mannscript never published (Runnegar, 1972).

The second specimen was dissected and the functional

anatomy described by Morton (1980).

On November 2004, while diving at Bahia Concha, a

sheltered bay near Santa Marta, on the Cailbbean coast

of Colombia (1 1°17'56" N, 74°08'52" W), a pair of open-

ings on the sandy bottom at a depth of about of 4 m

caught the atteiition of one of ns (DCT). Excavating

deeply around the holes, she e.xposed a large clam

(about 20 cm long) bearing white, pearly vah'es that she

hadn't seen before. The animal was photographed

(Figure 1) and, not being the subject of her study, re-

leased on the bottom. Three years later, the photograph

was shown to the first author, who immediately recog-

nized the clam as P. Candida since several years before

he had discovered empty valves of this species in the

same general area (Diaz and Borrero, 1995).

On Januaiy 26, 2008, two of us (JMD and FG) visited

Bahia Concha in order to search for other living speci-

mens of P Candida. After almost one hour diving along

the shore, we detected a pair of openings slightly protmd-

ing from the sandy bottom at 3 m depth. These structures

matched the size and shape of the apertures of the large,

bifid siphonal tube of P. Candida (Figure 2). Both aper-

tures closed sphincter-like and retracteil slightly into the

sediment when we started to dig around them. Indeed, we

dug out a specimen of the “living fossil,” though not as

large as the specimen found three years before. Unfortu-

nately, the anterior part of both v’alves broke during col-

lecting, l)ut the entire soft parts of the animal were still

present; the length of the valves was approximately 70 mm

and the siphonal tube was 55 mm long. The specimen vv^as

presented in 100% alcohol and tleposited in the marine

invertebrate collection at the Uuiversidad de Los Andes in

Bogota (IM-Andes 559).

In regard to the mode of life of P Candida, die specimen

was posidoned nearly veitically in the bottom at a small

angle, on its anteroventral margin. This obsemidon agrees

completely with the inferred life posibon of P ^igantea

Page 20

THE NAUTILUS, Vol. 123, No. 1

Figure 1. Specimen ol Fliolddomi/a Candida touiid on

Novemhei' 2004 in Bahia Concha, Colombia.

(J. cle C. Sowerhy in Pdttoii, 1S36) from the Early Creta-

cetnis ol west-central Argentina by Lazo (2007: tig. 8). This

author also stated (p. 385) that "in modem P. Candida the

slioitness ol the (ventnil) inhalant siphon relative to

the exlialant may not indicate that the animal lay on its hack

as suggested by Moiton (1980: fig. 57). The longer siphon

may hmction as a sort of tube or chimney to discharge

waste water well-above the entrance of clean water at the

inhalant siphon apeitnre.” Therefore, it seems likely that P.

CYDidida has a suspension-feeding habit rather than a pedal-

feeding system as postulated by Morton (1980) based on

the presence of a pc“dal gapt‘ and accessom mnscies.

A suspension-feeding habit has been commonly suggested

in Jurassic and Ci'etaceons Pholadoint/a species as well

(references in Lazo, 2007). The habitat of P. Candida, at

least in Balria Concha, also suggests a suspension-feeding

Figure 2. (ilosc’-np \iew (Irom ahox'c) ol the openings ol

the siplional (nhe ol Plioladoim/a Candida proirnding Irom the

bottom siirlacc.

mode of life. The habitat here is characterized by coarse

grain sediments in a shallow setting adjacent to the beach,

where w'ater motion caused fry incoming waves and drift

curremt is clearly perceptible. This is not the appropriate

en\ironment for accumulation of enough detritus on the

bottom to guarantee the alimentaiy I'equirements of a

relatively large, almost sessile, deposit feeder.

Tlie preseirt record is definitive evidence that Pholado-

mtja Candida is not e.xtinct. Moreover, tlie specimen coUect-

ed makes it now pcrssible to undertake genetic secpiencing

of the only modem representative of an ancient lineage.

Comparative molecular sequencing of P. Candida with

other anomtilodesmatan species and representatives of ad-

ditional, presumably related groups may proMde not only

an insight into the evolution of the other wddely differing

superfamilies of the Anomalodesmata, but might also

reveal clues as to the origin of the Myoida.

ACKNOWLEDGEMENTS

This research is part of the activities of the Census of Marine

Life - Caribbean Regional Committee in the Soutliem

Caribbean. We are indebted to an anonymous reviewer for

helpful suggestions to improve the manuscript.

LITERATURE CITED

Cox, L. R. 1969. Family Pholadomyidae. In: Moore R.C.

and C. Teiclieit (eds.). Treatise on Invertebrate Pdeontolog)'.

Part N, Mollnsca 6, Bivalria 2. Geolo^gical Society of

America and University- of Kansas Press, Lawi-ence, NS27-

N838.

Diaz, J.M. and F. J. Borrero, 1995. On the occurrence of

Plioladomt/a Candida Sowerby, 1823 (Bivalvia; Anomalo-

desniata) on the Caribbean coast of Colombia. Journal of

Molluscan Studies 61: 407-408.

Gibson-Sniith, J. and W. Gibson-Smith. 1980. The status of

Pholadoini/a Candida G.B. Sowerby, I. 1863. The Veliger

23: 355-.356.

Lazo, D.G. 2007. The bivalve Piudadoini/a gigantea in the

Early Cretaceous of Argentina: Taxonomy, taphonomy,

and paleogeographic implications. Acta Palaeontologica

Polonica 520: 375-,390.

Morton, B. 1980. Anatomy of the “living fossil" Pholadonnja

Candida Sowerby, 1823 (Bivalvia: Anomalodesmata:

Pholadomyacea). Videnskabelige Meddelelser fra Dansk

uaturhistorik Forening I Kobenhavai 142: 7—101.

Runnegar, B. 1972. Anatomy oPPhoJadoniija Candida (Bivalvia)

and the origin of the Phokulomvidae. Proceedings of the

Malacological Societv' of London 40: 45—58.

Runnegar, B. 1974. Evolutional-)' histon- of the bivalve

Subclass Anomalodesmata. Journal of Paleontolog)' 48:

904-9,39.

Runnegar, B. 1979. Pholadoniifa Candida Sowerby: The last

cadaver unearthed. The V'eliger 22: 171 — 172.

Waterhouse, J.B. 1969. The relationship between tlie living

genus Pholadonn/a Sowerby and upper Paleozoic pelecy-

pods. Lethaia 2: 99—1 19.

Ziusmeister, \\'. J. 1978. Review of the bivalv'e genus Pholado-

nn/a from the Tertiai-y of California and the description ol

tw'o new species. The \Aliger 21: 231—235.

Thiara scabra (O. F. Miiller, 1774): The introduction of another

Asian freshwater snail into the United States

Fred G. Thompson

Florida Museum of Natural Ilistoi'v,

Uuiversih' ol Florida,

Gaiuesville, FL 32611 -7800 USA

[email protected]

Michael W. Heyn

Biolo^ Seetiou,

Florida Department of Eu\'iroumeutal

Proteetiou 2600 Blair Stoue Rd. MS 6.515

Tallahassee. FL ;32.3y9-2400 USA

Michael. Me\m@dep. state. 11. us

Drew N. Camphell

Miliau, Swaiu & Associates, liie.

2257 \hsta ParWay, Suite 19

\\’est Palm Beach ' !• L 33411 USA

(lean iphell@iuiliau. swaiu. com

Tlie Thiaridae consists of numerous species of non-marine

snails in Africa, Asia, Australia, tropical America, and

many island archipelagoes (VIorrison, 19.54, Glanbrecht,

1999). The family is nnnsnal in that parthenogenetic

reproduction is the normal reproductive mode, which

causes each indi\idnal to be reprodnetively isolated from

all other individuals. Thus, a single specimen is all that

is re(}uired to establish a new colony in an appropriate

habitat. A recent introduction into Florida fresliwater

systems is one such species.

T] tiara scabra (O. F. Aliiller, 1774)

The Pagoda Tiara

(Figi.ire 1)

OBSERVATIONS

Identification: The species is recognized by the fol-

lowing combination of characters (Brandt, 1974). ft has

a neomelanian operculum: the nucleus is offset strongly

toward the lower columellar margin (Thompson, 2006:

fig. 38). The shell is medium-sized, generally up to

20 mm in length, and cousi.sts of fi— 8 whorls remaining

in adults. Usually early juvenile whorls are worn away.

The shell is pagodiform with strongly shouldered whorls

that bear regularly spaced stout spines or knobs. Strong-

ly impressed spiral sculpture is present, which usually is

most distinct below the peripheiy. The ground color

is tawny watli vertical, rust-colored (lames and blotches

alternating with the spines.

Distribution: Widely deployed and locally abundant

throughout its range in South and Southeast A.sia, Soutli

China, the Indo-Australian Archipelago, and westei'n

Pacific Islands (Brandt, 1974; 163—164). The senior

author has encountered this species on many occasions in

Southeast Asia. It is tolerant to many environmental set-

tings, as is reflected by its wide distribution. One such

setting is canals in semitropical regions, such as ai'e abun-

dant in South Florida. It is not surprising to us that the

snail finally was found there. It is a medium-sized, ornate

organism tliat lias potential in the a(|uarium trade. We

suspect that the species was deliberately introduced be-

cause ol the i.solated localih’ where it is lound.

Figure 1. Thiara scabra (O. F. Miiller, 1774), UF 391616.

Scale bar = 1 cm.

Page 22

THE NAUTILUS, Vol. 123, No. 1

Discussion: On September, 2006, specimens were first

collected by Todd Ennis and Cheri Hughes of Tetra Tech,

Inc. at the Port Mayaca Aquifer Injection well site

(UP 391616). The Port Mayaca site is located in the

noithwest corner of Section 14, Towiship 40 South,

Range 37 East, near the connueiice of the L-65 Canal

and St. Lucie River (C-44 Canal) in the Town of Port

Mayaca, Martin Co., Plorida (26°59'17" N, S0°36'22" W).

The site is located on a South Ploi'ida Watei- Management

District (FWMD)-owned parcel of land adjacent to the

S-153 spillway and lock, which conveys water to and from

the L-65 Canal and the St. Lucie River. The location is

approximately 2,000 feet east of the Herbert Hoover Dike,

south of the sendee access road, and appro.ximately 100

feet west of the intersection with the L-65 canal. Property

to the nortli is under cultivation of sugar cane.

Doug Strom ol Water and Air Research Water & Air

Research, Inc., Gaines\111e, Plorida. reports to us that a

co-worker, Laura Line, collected this snail in the West

Palm Beach Canal, 5 kilometers from its junction wdtli

Lake Okeechobee on April 10, 2007.

Vernacular Etymology: The vernacular name Pago-

da Tiara is taken from Reeve (1860, pi. 26, fig. 182),

from a name he proposed for a common form of this

species. Tiara comes from the generic name Thiara,

which is derived from Persian through the Classical

Greek, and means a tiara. Thiara scabra possess a coro-

na of spines on the shoulder of the whorls reminiscent of

a tiara.

ACKNOWLEDGMENTS

We thank Rob Lasley, Plorida Museum of Natural

Histoiy, for photographing Eigure 1.

LITERATURE CITED

Brandt, R. A. M, 1974. The non-marine aquatic Mollusca of

Thailand. Archi\’ fiir Molluskenkunde 105: 1—423.

Glaubrecht, M. 1999. Systematics and the evolution of

\'i\'iparity in tropical freshwater gastropods (Cerithioidea:

Thiaridae sensu lato) — an oveiwiew. Courier Forschungs-

Institut Senckenherg 215: 91—96.

Morrison, J. P. E. 1954. The relationships of Old and New

^Vorld melanians. Proceedings of the United States

National Museum 103 (3325): 357—394.

Reeve, L. [A.] 1859—1861. Monograph of the genus

Melania. Conchologica Iconica; or illustrations of

the shells of molluscous animals, vol. 12, Melania-, pis.

1-59.

Thompson, F. G. 2006. An identification manual for the fresh-

water snails of Florida, http://www.flmnh.utl.edu/malacol-

og}7fl-snaiPsnailsl.htm. Accessed July 26, 2007.

THE NAUTILUS 123(1):23, 2009

Page 23

Erratum

A review of Tijphisopsis Jousseaume, 1880, and Typhinala Jousseauine, 1881 (Gastropoda: Miirieidae) of the eastern

Faeific (2006)

Ti/phi.sopsis carolskoghindae was described by Ilonart and Hertz (2006: 56, ligs. 17-25, 47-49, 59, 63). The liolo6'jre irorn Costa

Rica was deposited in SDNHM under registration ninnber 90773. However, tliere were paratvpes oi wliicb five were stated as from

Boca de la Honda, Panama at 7°27' N, 80°5f' VV, and deposited in the followdng institutions: MZUCR (one specimen, registration

n° 6153): VINHN (one specimen, registration n° 6991); B\f(NH) (1 specimen, registration n° 2()()50371) and 2 specimens in

R. Houart coll, (no registration number).

In a recent article, Villalobos-Rojas et ah (2008) stated: carolskoghindae , Ti/pl}i.soj>sis, Ilonart and Hertz, 2006: 56-58, figs. 17-

25, 47—49, 59. 63. T\pe localitv: Playas del Coco, Cnanacaste, Costa Rica (10°55'53" N, 85°69'51" W), 24-37 m depth, on mnd

bottom. PARATYPE VIZPlCR-6153 (sliell. Figure 5). Boca de la Honda, Veraguas, Panama (7°27' N, 80°51' W), in white sand.

Remarks: The coordinates and tlie collecting locality of this paratvpe appear to be incorrect since these coordinates plot inland.”

After having again contacted the person who found these specimens several years ago, we learned that the published localitv was

in error, in part dne to the label whicli was written “B. Honda” and which was misinterpreted as “Boca de la Honda”. Tlie exact

locality where these specimens (now parahpes) were found is not Boca de la Honda, Panama, but Bahia Honda, Panama, a place

situated in the Golio de Chiricjiii, West oi Isla Cebaco. The senior author already contacted the institutions wliere the paratxpes

were deposited ami gave the exact locality data.

Abbre\4ations: BM(NH): The Natural Histon-' Museum, Ijomlon; MNHN: Mnsenm national d'Histoire naturelle, Ihiris; MZUCR;

Mnseo de Zoologia. Universidad de Costa Rica, San Jose: SDNHM: San Diego Natural Histon' Mnsenm, California, USA.

ACKNOWLEDGMENTS

Tliank.s to Andre Vassart (Costa Rica) lor his veiw uselul cooperation.

LITERATURE CITED

Houart, R. and C. Hertz. 2006. A re\4ew oi' Ti/phisoj)si.s Jonsseanme, 1880, and Ti/phisala Jonsseanme, 1881 (Gastropoda, Mnricoi-

dea) of the eastern Pacific. The Nautilus 120: 52-65.

V'illalobos-Rojas, F., G. Guzman-Mora, A. and Y. Camacho-Garcia. 2008. Catalogue ol the hpe material oi mollusks tleposited at the

Zoolog)' Mnsenm, Universitv of Costa Rica. The Nautilus 122: 155-165.

Roland Houart

Institut royal des Sciences natnrelles de Belgi(pie

I'lie Vantier, 29

1000 Bruxelles, Belgium

Carole M. Hertz

Santa Barbara Vluseum oi Natural Ilistoiy

2559 Puesta tlel Sol Road

Santa Barbara, California 93105, USA

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THEt^NAUTILUS

CONTENTS

JUL 0 7 2009

J,/brar'.h%

Volume 123, Number 2

June 30, 2009

ISSN 0028-1344

James D. Williams A new species of freshwater mussel, Anodouta hoiifieldoruin (Bivalvia:

Arthur E. Bogan Unionidae), from the Gulf Coastal Plain drainages of Alabama, Florida,

Jeffrey T. Garner Louisiana, and Alississippi, USA 25

Fernanda Pires Ohlweiler A new species ol Belocanlus (Gastropoda: Veronieellidae) from southern and

Dan Jesse Gonyalves Mota southeastern Brazil 34

Suzete Rodrigues Gomes

A. J. Reft Sensor)' structures on the siphons of woodd)oring bivalves (Pholadidae:

J. R. Voiglit Xylophagainae: Xijlopha^a) 43

Yu. I. Kantor

Guido Pastorino

An unusual new genus and a new species of Bnccinulidae (Neogastropoda)

from the Magellanic Province 49

John Slapeinsky

A new species of Pan/phantopsis (Gastropoda: Pnlmonata: Charopidae) from

Crater Mountain, Simbn (Chimbu) Province, Papua New Guinea 53

Charles F. Sturm

Juan Jose Parodiz (1911-2007): ohituaiy and bibliography

59

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THE NAUTILUS 123(2);25-33, 2009

Page 25

A new species of freshwater mussel, Anodonta hartfieldorum

(Bivalvia: Unionidae), from the Gulf Coastal Plain drainages of

Alabama, Florida, Louisiana, and Mississippi, USA

James D. Williams

Florida Museum of Natural History,

University of Florida

Museum Road and Newell Drive

Gainesville, FL 32611 USA

[email protected]

Arthur E. Bogan

North Carolina State

Museum ol Natural Sciences

MSG 1626

Raleigh, NC 27699-1626 USA

[email protected]

Jeffrey T. Garner

Alabama Division of Wildlife

and Freshwater Fisheries

350 County Road 275

Florence, AL 35633 USA

[email protected]

ABSTRACT

A new species of unionid mussel, Anodonta hartfieklonnn, is

described from Coastal Plain streams of the eastern Cult of

Mexico drainages. It occurs in the Pearl River in Louisiana and

Mississippi, Pascagoula River in Mississippi, Tombigbee River

in Mississippi and possibly Alabama, Tensaw River in Alabama

and the Escambia River drainage in Alabama and Florida.

Based on shell morphology and presence of very thin green

rays, it belongs to a species group within the genus Anodonta

which includes A. coupeiiana, A. heardi, A. implicata, and

A. suborbiculata. Anodonta harifieldontm appears to be most

closely related to A. suborbindata, but differs in several aspects

of shell mori^hology. Anodonta stdjorbiadata is widespread in

the Mississippi Basin and has been widely introduced outside

its native range. Anodonta hartficldomm occurs in floodplain

sloughs and oxbow lakes in silty sand to mud sediments. Its

conservation status is unknown as its typical habitat is under-

represented in most mussel sampling programs.

Additional keywords: Cypress Floater, new species, taxonomy,

conservation, Alabama, Florida, Louisiana, Mississippi

INTRODUCTION

Southeastern United States Unionidae have received

considerable attention during the past half century.

Thus, their tttxonomy is well understood relative to some

other groups of aquatic invertebrates (e.g. crayfishes)

(Taylor et ah, 2007). However, there are remaining

undescribed unionid species (Williams et ah, 2008).

One undescribed species of Anodonta Lamarck, 1799,

was first recognized in the Pascagoula River during the

1980s by Paul Hartfield, U.S. Fish and Wildlife Service,

Jackson, Mississippi. Subsequent surveys revealed the

presence of this species in Gulf Coast drainages from

the Pearl River east to the Escambia River (Vidrine,

1993; Williams et ak, 2008).

The genus Anodonta, as presently conceived, occurs

in most of the Nearctic and Palearctic regions. The type

species, Anodonta cijgnea Linnaeus, 1758, is from west-

ern Europe. In North America, Anodonta occurs from

Alaska and Canada to Mexico. Turgeon et al. (1998)

recognized ten species in the genus. The number of

Anodonta species worldwide is not clear due to poor

delineation of taxa and varying inteqrretations of the

species concept (e.g. Mock et ah, 2004). Ortmann

(1912) observed that “in Europe the species-making in

this group has gone beyond all the bounds of reason.”

Anodonta Lamarck, 1799, is a nonien consewatnni

(ICZN, 1926: Opinion 94; 1959: Opinion 561). Hoeh

(1990) used morphological and allozyme data to produce

a phylogeny that divided North American Anodonta,

along with the European type species A. cijgnea, into

three clades: Anodonta; Pijganodon Crosse and Fischer

in Fischer and Crosse, 1894; and Utterbackia Baker,

1927. The latter two were elevated from subgeneric to

generic status by Hoeh (1990).

MATERIALS AND METHODS

Comparative material of Anodonta coupeiiana Lea,

1840, Anodonta hartfieldorum, Anodonta heardi Gordon

and Hoeh, 1995, and Anodonta suborbiculata Say, 1831,

from several museums was utilized in this description.

These institutions include Florida Museum of Natural

History, University of Florida (UF), Gainesville, Florida;

Mississippi Museum of Natural Science (MMNS),

Jackson, Mississippi; Museum of Comparative Zoology

(MCZ), Har\/ard University, Cambridge, Massachusetts;

North Carolina State Museum of Natural Sciences

(NCSM), Raleigh, North Carolina; Ohio State Universi-

ty Museum (OSUM), Columbus, Ohio; and National

Museum of Natural History, Smithsonian Institution

(USNM), Washington, DC. Adtlitional material from

the personal collection of Robert G. Howells, Kermlle,

Texas, was also examined.

Shell measurements were made to the nearest milli-

meter using digital calipers and included total length.

Page 26

THE NAUTILUS, Vol. 123, No. 2

lieiglit, wadth, hinge line length, and distance from

innbo to posterior shell tenninns. Total length is defined

as distance between anterior and posterior margins,

measured parallel to the hinge line. Shell height is dis-

tance between dorsal and ventral margins, measured

near the midpoint of the hinge line, peipendicnlar to

shell length. Shell width is maximnm distance between

the outer surfaces of the paired valves. Hinge line length

is distance from its anterior tenninns to the anterior end

of the ligamental notch. Distance from the center of the

nmbo to the posterior shell terminus was measured in a

straight line. The angle between dorsal margin and pos-

terior margin was measured to the nearest five degrees

using a protractor (Figure 1). Shell measurement data

were analyzed using Excel spreadsheet scatter plots wath

linear regression. An ANOVA on the angle measure-

ments was performed on the three species using SPSS

vl6.0. A post-hoc comparison using Tukey’s test was per-

formed to determine the significantly different groups.

Gross anatomy of soft tissues was described from fresh

and relaxed specimens fixed in 10% formalin and trans-

ferred to 70% ethanol. The anatomical descriptions fol-

low methods described in Williams et al. (2008).

gin and posterior margin was measured. Image of Anodouta

sul)orhicuhita modified from Burch (197.5).

Anodouta lunifieldoritm new species

Cy^3ress Floater

(Figures 2, 3)

Diagnosi.s: Anodonta haiifieldontm is di.stinguished

Irom other nnionid species by a combination of the

following characteristics: thin, compressed to inflated

shell, elliptical to oval outline, ventral margin rounded;

angle between dorsal margin and posterior margin usu-

ally 140° to 150°; hinge teeth absent; periostracnm

smooth, tawny to olive or brown, ppically with veiy thin

green rays; umbo only slightly elevated above hinge

line; umbo sculpture in the form of parallel bars in

adults; inner lamellae ol inner gills connected to visceral

mass only anteriorly; supra-anal aperture small, sepa-

rated from excurrent aperture by wide mantle bridge

(may be longer than either of the two apertures); outer

gills marsupial; marsnpium occupying entire gill, w^ell

padded wdien gravid; secondary water tubes present in

gravid marsnpia; glochidinm v\4th st)4iform hooks.

Description: Length to 120 mm; shell thin; moderately

inflated; outline oval; posterior margiii narrowly rounded

to bluntly pointed; angle between dorsal margin and pos-

terior margin 140° to 155° (mean = 146°) (Table 1); ante-

rior margin broadly rounded; dorsal margin stixiight;

ventixrl margin convex; posterior ridge low, rounded; pos-

terior slope moderately steep, slightly concave, occasion-

ally extending into a veiy low dorsal wing; umbo broad,

moderately inflated, barely elevated above hinge line; um-

bo sculpture nodulous ridges in young and parallel bars in

adults; periostracnm tawny to olive or brown, typically

wtth veiy thin, olive to greenish browm rays that often are

obscure in adults. Pseudocardinal and lateral teeth absent;

nmbo cavity wide, shallow^; nacre white, sometimes with

salmon tint in nmbo cavity’ (Figures 2, 3).

In life the mantle is creamy-white to tan or golden-

tan, may be diill-orange external to pallial line, mantle

outside of apertures dull-orange to grayish-browm; ■vis-

ceral mass creamy-white to tan, may be dull-orange ad-

jacent to foot; foot dull-orange to creamy-white or tan.

Gills gold to tan or brown; dorsal margin sinuous to

concave, ventral margin convex; gill length 57-69% of

shell length; gill height 31-53% of gill length; outer gill

height 79-100% of inner gill height, outer gill height

may be greater than inner gill height in gra\4d indivi-

duals; inner lamellae of inner gills only connected to

\4sceral mass anteriorly. Outer gills marsupial; glochidia

held across gill length; well padded w4ren gravid; light-

brown to brownish-orange. Labial palps tan, may have

golden cast; straight to concave dorsally, convex ventral-

ly, bluntly pointed distally; palp length 21-40% of gill

length; palp height 46-67% of palp length; distal 27-

5(8% of palps bifurcate. Incnrrent aperture usually lon-

ger than excnrrent and snpra-anal apertures; supra-anal

and incurrent apertures occasionally of similar length.

Incurrent aperture length 8-12% of shell length;

creamy-white to dull-orange within, sometimes grayish

or rusty-browm basal to papillae; papillae in 2-3 rows,

inner row usually larger, simple, short, thick; papillae tan

to dnll-orange, larger papillae often with black edges

basally. Excurrent aperture length 5-8% of shell length;

creamy-white to dull-orange within, marginal color band

rusty-tan to dull-orange with black lines perpendicular

to margin, generally with some lines converging proxi-

mally, some individuals with lines interconirected to

form an irregular reticulated pattern; excurrent aperture

margin smooth, may undulate. Supra-anal aperture

length nsnally 5-8% of shell length, occasionally to 16%

of shell length; creamy-white to tan within, usually with-

out marginal coloration, occasionally with a thin, irregular

tan baml; supra-anal aperture margin smooth; mantle

bridge separating snpra-anal and excurrent apertures usu-

|. 1). Williams et al., 2009

Page 27

Figures 2, 3. A)iodonto liaiifiehlonim. 2. Holotspe UP' 375595, lengtli 112 mni. Fish Lake, o.xhow oH Pascagoula River, 1 air mile

[1.6 air kilometers] southeast ot Highway 614 bridge, southwest ol Wade, 30.6016°N; 88.6233°W, Jackson Countw Mississippi, 27

Oct. 2000. © Richard T. Biyant. 3. Non-tvpe specimen UF 358657, length 114 mm. Slough and gravel pits adjacent to Escambia

River, Mystic Springs boat ramp, 1 mile [1.6 kilometer] southeast of McDaxid, Escambia Counp', Florida, 30.92656°N; 87.28597°W

20 Sep. 1999. © Richard T. Bryant.

ally imperforate, of variable length, 67-480% of siipra-

anal length, occasional inclividnals with a short seconclaiy

mantle bridge anterior to primaiy bridge.

Minor soft anatomy differences were noted between

Anodonta haiifieldoniin specimens collected from the

Pascagoula and Escambia River systems. Incurrent aper-

tures of Pascagoula indmduals (S-10% of shell length)

were shorter than those of Escambia indiwdnals (10-

12% of shell length). Length ol the mantle britlge separ-

ating excurrent and supra-anal apertures varied more

widely in Pascagoula individuals (67-480% ofsnpra-anal

length) than in Escambia individuals (90-420% of snpra-

anal length). Conversely, wader x'ariation in labial palp

size was obsen'ed in Escambia individuals (palp lengtli/

gill length 21—40%; palp height/palp length 47-67%)

than Pascagoula indixidnals (palp length/gill length 25-

29%; palp height/palp length 54-57%). Gill height

in relation to gill length wars greater in Escamliia indivi-

duals (40-53%) than in Pascagoula indixiduals (31-

38%). These differences are considered sliglit and could

be an artifact ol modest sample sizes (n = 6 from each ol

the two drainages), so further comparisons are needed.

No material from the Pearl River drainage or Mobile

Basin w^as available for comparison.

Table 1. Frequency distribution of the angle measurement between dorsal margin and posteri(.)r margin of Anodonta

harifieldornnn A. heardi and A. snborbicniata . Anodonta snborbiadata wars signilicantly dilferent (p < O.OOl ) Irom the other Rvo

species. There was no significant difference among the other Rvo species (p = 0,534).

Species Angle bcRveen dorsal margin and posterior margin

Page 28

THE NAUTILUS, Vol. 123, No. 2

T>pe Material: Holot>pe: UF 375595, length 112 inm.

Fish Lake, oxbow off Pascagoula River, 1 air mi. [1.6 air

km] SE of Hw)’ 614 bridge, SW of Wade (3().6016°N;

S8.6233°\\'), jackson Count); Mississippi, 27 Oct. 2000.

Parat)pes: Pascagoula River Drainage: Mississippi:

Jackson County: MCZ 361689, length 89-107 mm

(3 dn' shells). Fish Lake, o.xbow oft Pascagoula River, 1

air mi. [1.6 air km] SE of H\\'y 614 bridge, SW of Wade

(30.6016°N; 88.6233°W), 27 Oct. 200(). MMNS 6973,

length 60-101 mm (6 dn-" shells), Pascagoula River at

mouth of Dead Ri\er Lake (30.59236°N; 88.5976rW),

27 Aug. 1986. NCSM 29799, leu^h 58-78 mm (2 alcohol

presened), Fish Lake, o.xbow off Pascagoula River, 1 air

mi. [1.6 air km] SE of Hw^' 614 bridge, SW of Wade

(.30.6016°N; 88.6233°W), 27 Oct. 200(1 NCSM 28212.

length 57-101 mm (13 alcohol preserved). Dead River

Lake (mouth) ofl Pascagoula River, [6 air km SSW center

of Wade] (30.5944°N; 8fS.5979°W), 27 Aug. 1986. OSUM

80078, length 88-106 mm (3 dn' shells). Fish Lake, oxbow

oil Pascagoula River, 1 air mi. [1.6 air km] SE of H\w 614

bridge, SW of Wade (30.6016°N; S8.6233°M'), 27'Oct.

2000. UF 428535 (6 in 95% alcohol preseiwed), Pasca-

goula Ri\er at Paper Mill Camp (30.63228° N;

88.65240°W), 21 Aug. 2008. UF 428536 (4 iu 70% alcohol

presened), Pascagoula River at Paper Mill Camp

(30.63228°N; 88.65240°W), 21 Aug. 2008. UF 428544,

length 53-120 mm (9.5 drv shells). Fish Lake, oxbow off

Pascagoula River, 1 air mi. [1.6 air km] SE of Hwy 614

bridge, SW of Wade (30.6016°N; 88.6233°W), 27 Oct.

2000. UMMZ 302000, length 90-105 mm (3 diw shells).

Fish Lake, oxbow off Pascagoula River, 1 air mi. [1.6 air

km] SE of IRw 614 bridge, SW of WTide (30.6016°N;

88.6233°W). 27 0ct. 2000, llSNM 1124163, length 79-99

mm (4 dn' shells). Fish Lake, oxbow off Pascagoula River,

1 air mi. [1,6 air km] SE of Il\w 614 bridge, SW of Wade

(30.601 6°N; 88.6233°W), 27 Oct. 2000.

Other Maternal Exainined: Escambia River Drain-

age: Alabama: Coxington County: NCSM 45095

(16 alcohol presen'ed), Cautt Resenoir. Conecuh River,

CR 86 [Dunn’s Rridge Road], [3.2 air km NE center of

Cautt] (31.42573°N;"86.4576°W), 5 Nov. 2006. NCSM

45145 (1 dn' shell). Point A Resen'oir [Conecuh River],

SW corner, 1.77 km NW from intersection of CR 70

and US 84, (31.35953°N; 86.51628°W), 8 Nov. 2005;

Alabama: Escambia County: UF 375317 (1 alcohol

presen'ed). Old Faulkner Lake, oxbow lake of Conecuh

River, 2 air mi. [3.2 air km] SE of Pollard, 0.5 air mi. [0.8

air km] N of Florida state line, 29 June 1995; Pdoritla:

Escambia County: UF 358657 (6 dn' shells), slough

and gravel jrits adjacent to Escambia River, at Mystic

Springs boat ramp, 1 mi. [1.6 km] SE of McDavid

(30,92656°N; 87.28597°W), 20 Sep, 1999. UF 376605 (5

(In' shells), slough and graved pits adjacent to E.scambia

River, at Mystic Springs boat ramp, 1 mi. [1.6 km] SE of

McDavid, 5 }nly 1992. UF 428537 (3 iu 70% alcohol

presen'ed), Escambia River at Rind Springs boat ramp

(30.92675°N; 87.28647°W), 19 Sc-p. 2007. UF 428538 (6

in 95% alcoliol presei'ved), Escambia River at Bluff

Springs boat ramp (30.92675°N; 87.28647°W), 19 Sep.

2007. NCSM 28251 (16 alcohol preserved), Escambia

River, abandoned gravel pits adjacent to Mystic Springs

boat ramp, [point estimated 1.6 air km SSE center of]

McDavid (30.S5559°N; 87.31266°W), 9 Aug. 1992,

Mobile Tensaw River Drainage: Alabama Baltl-

■win County: UF nncataloged (2 alcohol preserved),

slough off Tensaw Lake about 1 air mi. [1.6 air km] SS’W

of Hubbard Fish Camp and Landing (31.049097°N;

87.871753°W), 18 Sep. 1999. These specimens were

misplaced during a transfer from U.S. Geological Sniwey

to the Florida Museum of Natural Histon;

Pascagoula River Drainage: Mississippi: George

County: MMNS 5503 (3 diy shells), McCrea Dead River

E of Dale (30.83302°N; 88.74750°W), 29 May 2000.

Pearl River Drainage: Louisiana: St. Tammany

Parish: iVIMNS 6444 (1 diy shell). Pearl River,

Mississippi and Louisiana state line, at Walkiah Bluff,

mouth of slough about 0.5 mi. [0.8 km] upstream of boat

ramp, 18 Sep. 1986. Specimen was not examined; record

based on a personal communication with Bob [ones,

AIMNS; Missis-sippi: Marion County: MAINS 2168

(1 diw shell). Pearl River, iu viciuit)' of Columbia, 24

Apr, 1986.

Comparison with Similar Species: Anodonta hart-

fieldoruni shells resemble those of A. .suborbiculata

(Figure 4) but are less round and usually more inllated,

with a more inflated umbo that is elevated slightly above

the hinge line. It also may resemble Pi/gamHloii grmidis,

but that species has a much more inflated umbo that is

considerably elevated above the hinge line. Anodonta

Junifieldonnn may vaguely resemble Ufterbackia inibe-

cillis and Utterbackia peggi/ae folmson, 1965, but those

species are more elongate and their umbos are not ele-

vated above the hinge line. Anodonta hartfieldonon is

similar in shell moipholog)' to A. Iteardi (Figure 5), but

the two species are allopatric, with A. lieardi occurring

only in the Apalachicola Basin and easRvard in the

Ochlockonee River (Gordon and Hoeh, 1995; Brim Box

and Williams, 2()()0).

Shell proportkms of Anodonta haiijieldorum differ

from those of A. siiborbicidata and A. heardi. The most

notable differences are in the relative proportions of

shell height and length, as well as the angle beRveeu

dorsal margin and posterior margin, Anodoi}ta haiifiel-

dortini shell height, relath'e to length, is greater than

that of A. heardi bnt less than that of A. suborbiculata

(Figure 6). The angle between dorsal margin and poste-

rior margin is about ecjual iu A. hartfieldonini (mean =

146°, N = 68) and A. heardi (mean = 147°, N = 19) but

is greater than that of A. suborbiculata (mean = 129°, N

= 127). Frerjuency distributions of these angles are pre-

sented in Table 1.

Di.slribuliou: Anodonta hartfieldorum occurs (rom

the Escambia River drainage iu Florida and Alabama

west to the Pearl Rivei' diviinage iu Louisiana and Alissis-

sippi (Figure 7). It is known Irom the Escambia River,

]. D. Williams et al„ 2009

Page 29

Figures 4, 5. Anodontu species. 4. .A. siihorhinihifa. UF 370151, length 124 vmn. Coosa Hiver, \\'eiss Resenoir, inoiitli oi Big

Cedar Creek, about 2 air miles [3.2 air kilometers] east ol Alabama and Ceorgia state line, Coosa Ri\’er Mile 25S, 34.bS473°N;

85.40549°\\', Fkn'd Coimte Ceorgia, 20 Ang. 1997. (T) Ricliai'd T. Bnant. 5. A. hcanh. UF 358656, length 1 13 mm. Harrison Creek,

north side ol lirst 180° bend, above conihience of Brothers Ri\c‘r, 29.873019°N; 85.037933°\\', Franklin Connt\', Florida, 7 Sep.

1991. © Richard T. Biyant.

Escambia and Santa Rosa counties, Florida, upstream to

Gantt Resenoir, on Conecuh River, Coxington Conntx',

Alabama. In the Mobile Basin, it is kiumni from the Tom-

bigbee River drainage in Lowmdes Conntv Mississippi,

and a single site on the Tensaw Ri\er, Baldwdn County,

Alabama. Anodontn haidfieldonim is tonnd in lower

reaches of the Pascagoula River dminage in George and

Jackson counties, Mississippi, and has been reportc'd

from Pearl River in Alississippi (Vidrine, 1993; [ones

et ah, 2005) and in Louisiana where the Pearl River forms

a common bonier between the two states.

Habitat and Biology: Aiiodoitfa haiificldonnn occurs

in water with little or no current such as oxbow lakes and

sloughs. It has colonized Gantt and Point A resenoirs.

Substrates in these habitats are tvyiically composed of

mud or muddy sand, often with detritus.

Anodoiila haiifieldoniiii is a long-term brooder, pre-

sumably graxid from late summer or autumn to the

folloxx'iug spring or summer. Graxid indixiduals brooding

mature glochidia hax'e been obsen’ed in late October

and early Nox'ember in Pascagoula Rix'er and Gantt

Resenoir, Conecuh River, respectix'elxx Glochidial hosts

of this species are unkuoxxm.

Discu.ssion: A)K>donta haidfieldoniiii appears to be-

long to a species group that includes A. siihorhiciilata.

A. coupeiiaiw, A. hcardi and Anodoiita wpdicata Sax',

IS29. A common morphological feature shared among

these species is umbo sculpture that consists ol parallel

bars in adults. |ux'euiles ol this group txpicallx/ have fine

green rays radiating from the umbo, but this feature is

often obscure in adults xxith a darker periostracum. .Mo-

lecular genetic data supports the relationship ol this

group, xx'hich is coidiued to the eastern United States

(Zanatta et ah, 2007). Aiiodcmta siiboii>iail(ila is native

to the Alississippi Basin and some central Gull Coast

drainages, A. hartficldnrnm and A. hcardi occur in east-

ern Gulf Coast drainages, and A. coiipcriana and A.

iiit))licata in Atlantic Goa.st drainages. There are popula-

tions of A. siihorhicniata in the Brazos, Neches, and

Sabine Rix-er di'aiuages, east Texas, that diller somexvhat

in shell moqxhologx' (lloxx'ells et ah, 1996). Thex’ are

slightly more inllated than txpical A. siihorhiridata.

Page 30

THE NAUTILUS, Vol. 123, No. 2

Figure 6. Scatter plot of the relationship between shell height and shell length (min) in Anodonta haiifieldorum. A. heardi. and

A. suborl)icidata.

However, additional research is needed to resolve the

relationships of these populations.

There have been several reports of Anodonta stiborbi-

ciilata in the Escambia River drainage in Alabama and

Elorida. The first known Escambia drainage specimen of

Anodoida was collected in 1917 by C. A. Burke, Irom

Chumnckla Springs, Santa Rosa County, Elorida. This

specimen (UMMZ 101375) was reported as A. snborbi-

culata by Butler (1990) on the basis of a personal com-

munication from William H. Heard, but it is most likely

based on A, liailficldontni (this specimen could not be

located to confirm identification). Clench and Turner

(1956) did not encounter A. snborJncidata in their sur-

vey of freshwater mollusks of Elorida and .southern Ala-

bama. The report ol A. snborbicidata from Gantt Lake,

Covington County, Alabama (MCZ 26751S), by johnson

( 1969) is based on A. haiffieldormn. In the identification

manual of freshwater clams of Elorida, Heard (1979)

reported ami illustrated A. snhorbiculata from the

Escambia River drainage but locality data for the illu-

strated specimen were not given. A single juvenile

(48 mm; UE 134930) A. snborbicnbda Irom the E.scainbia

River, Elorida, was rejroited and illustrated by Williams

and Butler (1994). Anodonta suborbicidata appears to be

a recent colonizer of the Escambia drainage, with the first

confirmed records from the 1980s. The two species occur

s)mtopically in Gantt and Point A reseiwotrs, Gonecuh

Rh'er, Alabama.

The natural o.xbow and slough habitats of Anodoida

haiifieldortiin are often overlooked or avoided during

mussel suiweys, and these habitats have been greatly re-

duced due to channelization and impoundment of large

rivers. These factors probably contributed to the dearth

of records and museum material. A systematic sun’ey of

Gulf Goast lloodplain lakes and reseiwoirs is recjuired to

determine the current conservation status of A. haiijiel-

dontin. Additional comparative analyses of soft anatomy

and molecular genetics are also needed to firmly resolve

taxonomic relationships within the genus Anodonta.

Conseiwation Status: The fact that Anodonta haiifiel-

donnn has not been previously recognized has precluded

its inclusion in conseiwation status reviews. However, this

species does not appear to be imminently imperiled. It

can be locally abundant, but may have declined in some

lloodplain lakes and sloughs that have been negatively

J. D. Williams et al„ 2009

Page 31

Figure 7. Known range oi' Anoclonfa limifieklonini (shaded

area) in Alabama, Florida, Louisiana and Mississippi. Solid

circles represent specific localities of A. haiffichlonim. T)']3e

locality of A. haiffichlonim is indicated by the star.

affected by cliannel incision following channelization of

adjacent rivers. Anodonta haiifieklonim consemition sta-

tus will remain unresolved until a systematic sinwey of

appropriate habitat is conducted.

Etymology: The species name haiifieldoniin is in

honor of Paul D. and Elizabeth A. Hartfield in recogni-

tion of their significant contributions to conseivation and

natural history in the southeastern United States. Paul is

a biologist in the endangered species program, U.S. Fish

and Wildlife Seiwice, Jackson, Mississippi, field office,

and has been instrumental in protection and recovery

of aquatic species. Elizabeth (Libby) is director of the

Mississippi Museum of Natural Science, Jackson, Mis-

sissippi, whei'e she presided over the enhancement of

the institution, which is now one of the pi'emier

museums in the southeast. She has also played an inte-

gral role in conseiwation and enMronmental education.

The common name, C\q3i'ess Floater, is in refei'ence to

the C)qn'ess tree which is common along the flood plain

sloughs and baclcwater oxbow lakes where Anodonta

haiffieldoniin is found.

Comparative Material Examined:

Anodonta heardi

Apalachicola River Drainage: Florida: Franklin

County: UF 358656 (1 diw shell), Hanison Creek,

[tributary of Apalachicola River] at first 180° tur'n, N side

of bend (29.873019°N; 85.037933°W), 7 Sep. 1991.

Florida: Gulf County: NCSAl 30334 (1 alcohol pr'e-

serwed), Chipola River; Florida Hw)' 22 [CR 22/Lake-

gr'ove Road], [2.8 air' km NF] of Wewahitclrka

(30.12766°N; 85.17638°W), 6 Aug. 1988. UF 375520 (1

dry shell), Apalachicola River at river mile 45.3, at the

Wewahitclrka Boat Ramp, about 5 air mi. [8 air km] NF

of Wewahitclrka, 1 Sep. 1999. UF 428532 (11 dry shells),

Apalachicola River Mile 46.8, along r'ight descerrdiirg

bank of I'iver (30.1819°N; 85.1344°W), 7 Aug. 2006.

Florida: Jackson County: UF 1915 (1 dry shell),

Tarrwrt Pond", 3 nri. [4.8 km] N of Sneads, 1 Apr'. 1955.

Florida: Feon County: MCZ 267515 (1 dr")' shell),

Ochlockorree River at US Hwy 27, 11 mi. [17.7 knr] NW

of Tallahassee.

Florida: Fiberty County: UF 381286 (1 dry shell),

Florida River, fr'orrr dowrrstreanr rrear SW edge of Acorn

Lake to poiirt dowrrstr'earn of head of feeder slough iirto

Fvei'ett Sloirgh, 4 Jurre 2002.

Georgia: Crisp County: NCSM 28259 (1 alcohol

pr'esei'ved). Lake Blackshear, 0.3 mi. [0.5 knr] S of US

280 bi'idge at edge of Georgia Vetenm’s Menror'ial State

Park, [poirrt estinrated 6.2 air km F cerrter of Cobb]

(31.9624°N; 83.9224°W), Sep. 1992. UF 376024 (2 dry

sliells). Lake Blackshear, US Hwy 280 crossirrg (F side)

nriddle of Lake Br'idge, Jurre 1995.

Ochlockonee River Drainage: Florida: Leon Coun-

ty: UF 370608 (4 alcohol preserwed), Ochkrckorree

River aborrt 3.5 air mi. [5.6 air km] S [SW] of Rt. 20

bi'idge (30.34S04°N; 84.69356°W), 19 July 1993.

Anodonta suborbicvlata

Ai'kansas River Drainage: Ai’kansas: Crawford

County: USNM 124422 (2 dry shells), [Ai'kansas River,]

Vair Burerr.

Atehafalaya River Drainage: Louisiana: St. Martin

Parish: OSUM 76142 (1 dry shell), 4 mi. [6.4 knr] SF of

Herrdei'soir [6.4 km SF of Herrdei'sorr, 4.8 kirr W of Butte

La Rose] (30.28140°N; 91.73600°W), 27 Sep. 1975.

Escambia River Drainage: Alabama: Covington

County: MCZ 267518 (1 dry shell), Cleaiwiew, on

Coirecuh River, Gantt Lake at LIS Hw)/ 29. NCSM

35282 (1 alcohol pi'eserwed). Point A Resei'voir', [poiirt

estinrated 2.3 air km NF cerrter of River Falls]

(31.36796°N; 86.52074°W), 2005. NCSM 48025 (3 dry

shells). Point A Reseiwoir [Conecuh River], SW corirei',

1 .77 km NW fi'onr irrtei'section of CR 70 and US 84, [2.3

air krrr NF center of River Falls] (31.35953°N;

86.51628°W), 8 Nov. 2005. UF 375318 (1 alcohol pi'e-

seiwed) Patsaliga Creek, slough on impounded lower

end, about 0.7 air nri. [1.1 air km] N of CR 59 bridge

(31.382518°N; 86.522834°W), 24 July 1995.

Florida: Eseambia County: UF 134930 (1 alcohol

pi'eseiwed) Escambia River', Rt. 4 ci'ossrng, 2.8 km F of

Century, 13 km NNF of McDavid, 7.8 knr W of Jay,

3 Jmre 1998.

Page 32

THE NAUTILUS, Vol. 123, No. 2

Mississippi River Drainage: Illinois: Carroll Coun-

ty: UF 225860 (4 diy shells), Thomson Lake.

Tennessee: Shelby County: MCZ 152833 (7 dry

shells), Mississippi River, Presidents Island, near Memphis.

Mobile Basin Drainage: Alabama: Cherokee Coun-

ty: UF 374082 (1 dry shell), Coosa River at island,

about 0.8 mi. [1.3 km] upstream of Hwy 20 bridge (Gar-

rett Rridge), 7 Aug. 2000. UF 374282 (3 diy" shells),

Coosa River at Large Island, about 1 mi. [1,6 km] down-

stream from Maple Grove, 7 Aug. 2000.

Alabama: Monroe County: UF 374748 (10 dry

shells), slough off Alabama River, about 1 mi. [1.6 km]

upstream of Claiborne Lock & Dam, on west bank,

17 Sep. 1999.

Alabama: Tallapoosa County: UF 376505 (2 dry

shells). Lake Martin at Wind Creek State Park, about

6 mi. [9.7 km] S of Alexander City, 28 Jan. 2004.

Alabama: Walker County: OSUM 58673 (1.5 dry

shells), BullbaiTi Creek, [6.1 km S of Jasper] (33.77000°N;

87.25888°W), 14 Ang. 1993. OSUM 59644 (1 diy shell),

Bullbam Creek, [6.1 km S of Jasper] (33.77000°N;

87.25888°W), 24 Feb. 1997.

Alabama: Wilcox County: UF 244013 (2 dry shells).

Millers Ferry, 9 mi. [14.5 km] NW of Camden, 200 m N

of Rt. 28 bridge over Alabama River, East Bank Park, 24

Sep. 1988. UF 374742 (1 dry shell), slough off Alabama

River, about 1 mi. [1.6 km] upstream of Claiborne Lock

& Dam, on west bank, 17 Sep. 1999. UF 376590 (1 dry

shell), impoundment of Alabama River, East Bank Park

at Millers Feriy, just NE of Hwy 28 bridge, about 7 mi.

[11.3 km] W of Camden, 10 Sep. 1988. UF 376593 (1 diy

shell), Coosa River at Large Island, about 1 mi. [1.6 km]

downstream from Maple Grove.

Georgia: Floyd County: UF 376151 (1 dry shell),

Coosa River (Weiss Reservoir), at mouth of Big Cedar

Creek, about 2 air mi. [3.2 air km] due E of Alabama and

Georgia state line (Coosa River Mile 258) (34.18473°N;

85.40549°W), 20 Aug. 1997.

Mississippi: Prentiss County: MM NS 9072 (2 dry

shells), Tombigbee River, borrow pits at Natchez Trace

at Brown Bottom, 12 mi. [19.3 km] ESE of Baldwin

(34.46835°N; 88.42968°W), 28 Oct. 1999.

Neches River Drainage: Texas: Nacogdoches

County: NCSM 30546 (3 diy shells), Sam Rayburn

Reseiwolr, Shirley Creek Park, [point estimated at end

of CR 496 (Sowell Bridge Road) in park, 9.7 air km

WNW of Broaddus] (31.31503°N; 94.37306°W), 12

Dec. 1995.

Texas: San Augustine County: MMNS 7477 (2 dry

shells), Sam Rayburn Reservoir on the Angelina River,

5 Nov. 1995. Robert G. Howells (6 dry shells, 2 alcohol

preserved), Sam Rayburn Reservoir, Ayish Creek arm at

CR 2923, 12 Dec. 1995.

Texas: Tyler County: Robert G. Howells (1 dry

shell), B.A. Steinhagen Reservoir, 29 Dec. 1993.

Ouachita River Drainage: Arkansas: Clark County:

UF 64057 (3 dry shells). Old River, Arkadelphia.

Pearl River Drainage: Mississippi: Madison Coun-

ty: MMNS 6263 (1 alcohol preserved). Pearl River,

left ascending bank, 0.5 mi. [0.8 km] below Lowhead

Dam, above Coal Bluff Water Park (32.61559° N;

89.75558°W), 1 Oct. 1987.

Mississippi: Pearl River County: MMNS 6444 (2

dry shells). Pearl River at Walkiah Bluff, mouth of ox-

bow 0.5 mi. [0.8 km] upstream from boat launch

(30.56513°N; 89.791 14°W), 18 Sep. 1986.

Red River Drainage: Louisiana: Bienville Parish:

USNM 119969 (1 dry shell). Mount Lebanon.

Louisiana: DeSoto Parish: USNM 133381 (4 dry

shells), Frierson Mill.

Louisiana: Rapides Parish: USNM 86699 (1 valve

of shell), Red River, Alexandria.

Louisiana: Webster Parish: OSUM 76518 (4 dry

shells). Cypress Swamp, [6.7 km E of Doyline, 3.5 km

W of Sibley] (32.53757° N; 93.33049° W).

Texas: Marion County: NCSM 33214 (2 dry shells).

Lake O’ The Pines (Big Cypress Bayou), [point esti-

mated 9 air km WSW center of Kellyville] (32.81322°N;

94.69791°W), 9 July 1996.

Texas: Camp/Titus counties: Robert G. Howells (5

dry shells). Bob Sandlin Reservoir, 10 July 1996.

Tennessee River Drainage: Alabama: Lauderdale

County: NCSM 43448 (16 dry shells), Pickwick Lake,

Tennessee River, behind small islands 8.8 air km E

[center] of Waterloo [Wright Quad] (34.89613°N;

87.96736°W), 6 Feb. 2009. NCSM 43449 (4 diy shells),

Pickwick Lake, Second Creek Embayment, 2.9 km

NE of Waterloo [Waterloo Quad] (34.93369°N;

88.03724°W), 6 Feb. 2009. UF 294002 (1 dry shell).

Elk River, Wheeler Lake, above Hwy 72 bridge, 1 mi.

[1.6 km] E of Rogersville, 1 mi. [1.6 km] above conflu-

ence with Tennessee River, above launch ramp, 23 Oct.

1998.

Alabama: Limestone County: NCSM 6187 (2 dry

shells), Tennessee River Mile 306, Decatur Boat Harbor,

[point estimated 1.8 air km E center of Decatur]

(34.60472°N; 86.9625°W), 2 Feb. 2000. NCSM 30439

(4 dry shells). Elk River backwaters, [point estimated

5.2 air km NW center of Cartwright] (34.90416°N;

87. 105 1 7° W), 7 Nov. 1976.

Alabama: Madison County: NCSM 33215 (2 dry

shell), Redstone Arsenal, unnamed tributary that flows

into Tennessee River, 22-23 July 1993.

Mississippi: Tishomingo County: UF 376595 (1 dry

shell). Yellow Creek downstream of junction with Pick-

wick Lake on Mississippi Hwy 25, about 1 mi. [1.6 km]

SE of North Crossroads, 1 Mar. 1973.

Tennessee: Humphreys County: NCSM 6710 (2 dry

shells), Tennessee River at Cuba Landing, 1-40 crossing,

[point estimated 10.7 air km NE center of Sugar Tree]

(35.87826°N; 87.93317°W), 5 Feb. 1977.

Tennessee: Meigs County: UF 365491 (2 dry

shells). Sugar Creek embayment of Hiwassee River,

Chickamauga Reseivoir on CR 306, about 2 mi. [3.2

km] E of junction 306 and 58, 21 Feb. 1998.

J. D. Williams et al„ 2009

Page 33

Trinity River Drainage: Texas: Liberty County:

MCZ 227966 (2 dry shells). Wards Prairie Lake near

Romayor (30.40965°N; 94.7S743°W).

Texas: Trinity County: Robert G. Howells (2 dry

shells). Lake Livingston, Ang. 1996. Robert G. Howells

(1 dry shell). Lake Livingston, 30 July 1996.

White River Drainage: Arkansas: Lawrence Coun-

ty: MCZ (3 dry shells). Black River at Black Rock.

ACKNOWLEDGMENTS

Richard Bryant provided the outstanding photographs of

specimens herein illustrated. We also extend our appre-

ciation to the following people and their institutions for

the loan of material in their care: Adam Baldinger (MCZ),

Robert Hershler, Paul Greenhall, and Linda Ward

(USNM), Robert Jones and Todd Slack (MMNS), Diar-

maid O’Foighil and Taehwan Lee (UMMZ), John

Slapcinsky and Gustav Paulay (UF), Jamie Smith

(NCSM). and Tom Watters (OSUM). We also thank

Robert G. Howells for the loan of material from his

personal collection and for inlormation on Anodonta

populations in Texas and Louisiana. John B. Burch

provided permission to use his illustration of Anodonta

sid}orbictdata. Nathan Johnson and Jennifer Bernatis

provided statistical support. Sherry Bostick provided

technical assistance with manuscript review and layout.

Cindy Bogan and Jamie Smith commented on a draft of

this manuscript, as did two anonymous reviewers.

LITERATURE CITED

Brim Box, J. and J.D. Williams. 2000. Uiiionid mollusks of the

Apalachicola Basin in Alabama, Florida, and Georgia. Ala-

bama Museum of Natural History Bulletin 21: l-f43.

Burch, J.B. 1975. Freshwater Sphaeriacean Clams (Mollusca:

Pelecypoda) of North America. Malacological Publica-

tions, Hamburg (Michigan), 96 pp.

Butler, R.S. 1990. Distributional records for freshwater mus-

sels (Bivalvia: Unionidae) in Florida and south Alabama,

with zoogeographic and taxonomic notes. Walkerana (for

1989) 3: 239-261.

Clench, W. J. and R.D. Turner. 1956. Freshwater mollusks ot

Alabama, Georgia, and Florida from the Escambia to the

Suwannee River. Bulletin ot the Florida State Museum,

Biological Sciences 1: 97-239.

Gordon, M.E. and W.R. Hoeh. 1995. Anodonta heardi, a new

species of freshwater mussel (Bivalvia: Unionidae) from

the Apalachicola River .system of the southeastern United

States. Walkerana 7: 265-273.

Heard, W. H. 1979. Identification manual of the freshwater

clams of Florida. Florida Department of Environmental

Regulation, Technical Series 4, 83 pp.

Hoeh, W.R. 1990. Phylogenetic relationships among eastern

North American Anodonta (Bivalvia: Unionidae). Malaco-

logical Review 23: 63-82.

Howells, R.G., R.W Neck, and H.D. Murray. 1996. Freshwa-

ter Mussels of Texas. Texas Parks and Wildlife Depart-

ment, Inland Fisheries Division, Austin, Texas, 218 pp,

Johnson, R.I. 1969. Further additions to the unionid fauna of

the Gulf drainage of Alabama, Georgia and Florida. The

Nautilus 83: 34-35.

Jones, R.L., W.T Slack, and P.D. Hartfield. 2005. The fresh-

water mussels (Mollusca: Bivalvia: Unionidae) of Missis-

sippi. Southeastern Naturalist 4: 77-92.

Mock, K.E., J.C. Brim-Box, M.P. Miller, M.E. Downing, and

W.R. Hoeh. 2004. Genetic diversity and divergence among

freshwater mussel (Anodonta) populations in the Bonne-

ville Basin of Utah. Molecular Ecology 13: 108.5-1098,

Ortmann, A.E. 1912. Notes upon the families and genera of

the najades. Annals of the Carnegie Museum 8: 222-365,

pis. 18-20.

Taylor, C.A., G.A. Schuster, J.E. Cooper, R.J. DiStefano, A.G.

Eversole, P. Hamr, H.II. Hobbs, III, H.W. Robison, C.E.

Skelton, and R.F. Thomas. 2007. A reassessment of the

conservation status of crayfishes of the United States and

Canada after lO-i- years of increased awareness. Fisheries

32: 376-389.

Turgeon, D.D., J.F. Quinn, A.E. Bogan, E.V Goan, EG.

Hochberg, W. G. Lyons, P. Mikkelsen, R.J. Neves, C.E, E.

Roper, G. Rosenberg, B. Roth, A. Scheltema, EG.

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

mon and Scientific Names of Aquatic Invertebrates

from the United States and Canada: Mollusks, Second

Edition. American Fisheries Society, Special Publication

26, 526 pp.

Vidrine, M.E 1993. The Historical Distributions of Ereshwa-

ter Mussels in Louisiana. Gail Q. Vidrine Collectibles,

Eunice, Louisiana, 225 pp.

Williams, J.D. and R.S. Butler. 1994, Freshwater bivalves. In:

Deyrup, M. and R. Franz (Eds.). Rare and Endangered

Biota of Florida. Volume IV University Press of Florida,

Gainesville, pp. 53-128.

Williams, J.D., A.E. Bogan, and J.T. Garner. 2008. Freshwater

Mussels of Alabama and the Mobile Basin in Georgia,

Mississippi and Tennessee. The University of Alabama

Press, Tuscaloosa, 908 pp.

Zanatta, D.T., A. Ngo, and J. Lindell. 2007. Reassessment of

the phylogenetic relationships among Anodonta, Pi/gano-

don, and Utted)ackia (Bivalvia: Unionidae) using mutation

coding of allozyme data. Proceedings of the Academy of

Natural Sciences of Philadelphia 156: 211-216.

THE NAUTILUS 123(2):34-42, 2009

Page 34

A new species oi Belocaiiliis (Gastropoda: Veronicellidae)

from southern and southeastern Brazil

Dan Jesse Gonsalves Mota

Suzete Rodrigues Gomes

Lahoratorio de Parasitologia, Instituto Butantan

Rua Vital Brasil, 1500

05503-900, Sao Paulo, BRAZIL

tlan_iesse@hotniail .coni

[email protected]

Fernanda Fires Olilweiler

Lahoratorio de Malacologia

Divisao de Progranias PLspeciais

Superintendencia de Controle de Endeinias

Rua Cardeal Arcoverde, 2878

05408-003, Sao Paulo, BRAZIL

[email protected]

ABSTRACT

A new species is proposed liased on material collected in the

states of Minas Gerais, Sao Paulo, Santa Catarina, and Rio

Grande do Sul, Brazil. The inorpholog)', ratlula, and jaw are

described and illustrated. The new species is compared to

Belocauhis angu.stipes, the only species ol the genus currently

considered valid. The main differences are found in the penis.

The new species has small projections, similar to tubercles, on

the anterior region of the glans, which can he scattered or

arranged in Rvo, three or more longitiuliual rows. The terminal

extremity of the glans has digitiform margin. The penis base is

short and poorly delined. The accessoiy gland is completely

immersed in the tegument. The description of the new species

extends the distribution range of Belocanhts for the states of

Minas Gerais and Siio Paulo, Brazil.

Additional kci/ioords: Veronicellidae, Belocauhis, moiphologv',

land slug

O

INTRODUCTION

Veronicellidae includes terrestrial slugs of the subclass

Gyiuuophila witli pautropical distribution, wdtb no shell,

and no developed puliuouai'y cavity such as that

(ibsei'ved in other terrestiial slugs of the subclass Puhuo-

nata. Some genera have been better studied because

they include intermediate host species for the Tiematodes

Anaiostronpijhis costa licensis Aloreira and Cespedes,

1971, and Anoiostronpijhis cantoiieusis (Chen, 1935),

parasites responsible lor abdominal augiostrougiliasis

and eosinophilic meuiugoeucephalitis, respectively.

Among the veronicellid species cited as intermediate

hosts for A. costa riceiisis are Sarasinula jilehcia (Fisher,

1S68) in Central and South America, Phi/llocaiilis mricga-

tiis (Semper, LSS5), Flii/llocattlis solciformis (d’Orhigny,

1835), Sarasinula lingiiacforinis (Semper, 1885), and Bclo-

caiiliis angustipes (Ileyiiemann, 1885) in southern Brazil,

and lor A. cairtoncnsis, Sarasinula niarginala (Semper,

1885) in the state ol E.s])h ito Santo, Brazil (Graeff-Teixeira

et ah, 1989; Graeff-Teixeira et ah, 1994; Rambo, 1997;

Laitano et ah, 2001; Caldeira et ah, 2007). Veronicellid

species have also been cited as damaging to agriculturtd

crops (Pereira and Gonyalves, 1949; Araiijo, 1952; Santos,

1959; Thome, 1993; Milanez and Chiaradia, 1999; Chiara-

dia et ah, 2004; Robinson and Plollingsworth, 2004).

When Hoffmann (1925) proposed Belocauhis, he in-

cluded in the genus South American species that pi'e-

sented the following characters: penis shaped as an

asymmetrical arrow oi‘ irregularly widened, vas deferens

opening terminally or subterminally, and presence of a

small accessoiy gland behind the duct ol the bursa copu-

latrix and rectum, partially covered by the tegument.

That author included six species in the genus without,

how^ever, designating a t)pe species: B. laugsdorfi (Fer-

ussac, 1822), B. boefzkesi (Aliller, 1879), B. pferocaulis

(Simroth, 1913), B. festae (Colosi, 1921), B. pulchcr

(Colosi, 1921) and B. sloauei (Cmier, 1817). Baker

(1925) designated Vagiuuhis augustipes as the t\pe spe-

cies of Belocauhis and regarded the latter as a ,s)uionym

oi' Augustipes Colosi, 1922.

The svmonvmv proposed by Baker (1925) was not ac-

cepted by Thome (1975), wdio revised the neotropical

species of Veronicellidae. According to Thome (1975),

Belocauhis is valid and characterized by the presence ol

a small accessoiw gland between the rectum and the

lemale genital pore (which, according to Thome, is ab-

sent in Augustipes). He mentioned additional character-

istics of Belocauhis: the rectum penetrates close to tlie

female genital pore, the penial gland has uniform and

sinuous tubules at the base (wdrere they are enveloped as

a whole l)y a membrane), and the bursa copulatrix has a

kidney or oval shape, with a short duct and other con-

necting duct that penetrates at the base of the gland

accessoiy.

The six species inchuled in Belocauhis by Hollmann

(1925) w'ere transferred to other Neotropical genera by

Thome (1975): Nooovaginiila Thiele, 1931, Siiurotliiila

Thome, 1975, Colosiiis Tliome, 1975, and Verouicella

Blaimille, 1817. According to Thome (1975), Belocaii-

F. P. Ohlvveiler et al„ 2009

Page 35

his included Pvo valid species: B. angustipes, described

originally from Taqnara (State of Rio Grande do Snl),

and B. aberrans (Ileyneniann, 1885), described origi-

nally from Santa Crnz do Snl (State of Rio Grande do

Snl). However, Pitoni and Thome (1981) and more

recently Thome (1993) regarded B. aberrans as a syno-

nym of B. angiistipes, rendering Belocaiihis monopqoic.

Belocauhis angiistipes merits special attention be-

cause of its \Hde distribution in southern South America

(east of the Andes). It is recorded from Rio Grande do

Snl and Santa Gatarina, in Brazil, as well as from Uru-

guay, Argentina, and Paraguay (Pitoni and Thome, 1981;

Thome, 1993; Thome et al., 1999; Simone, 2006; Thome

et ah, 2006) (Figure 1). Belocauhis angiistipes is also

recorded from Honduras and south of the United States

where it is considered an introduced species (Thome,

1989; Gaballero et al., 1991; Thome, 1993); the .species

distribution is discontinuous.

The study of a large number of specimens ol Belocaii-

his from twelve localities revealed the existence of a new

species, which distributed throughout southern and

southeastern Brazil. Its morpholog\' is veiy similar to

that of B. angiistipes, although both species can be dis-

tinguished from each other by some characters of the

male reproductive system. The morpholog)', radula, and

jaw of the new species are described and illustrated. It is

also compared to B. angiistipes, and the main differ-

ences and similarities are pointed out. New records of

Belocauhis are provided in the states of Minas Gerais

and Sao Paulo.

MATERIALS AND METHODS

The description of the new species proposed herein is

based on the examination of 92 specimens from 35 lots

collected from twelve different localities in southern

Brazil. The material is deposited in the collections of

the Museu de Zoologia, Universidade de Sao Paulo

(MZUSP) (lots 87747, holop^pe, 87748—87750, para-

68° 64° 60° 56° 52° 48° 44°

Figure 1. Map shoMng tlie distribution oi Belocauhis icilli-

bakloi new species and Belocauhis angiistipes, considering lit-

erature records and lots recently collected of the latter species

(from Caxias do Sul and Pinlial).

tyqres), Mnseu de Giencias e Tecnologia, Universidade

Catolica do Rio Grande do Sul (MGP) (lots 7971, 7972),

and in the Malacology collection of the Superintenden-

cia de Gontrole de Endemias, Sao Paulo, (SUGEN) (lots

8968, 8982-84, 8987-8995, 8997, 9005, 9006, 9016,

9019, 9020, 9031-9037, 9039, 9042, and 9043). Speci-

mens of B. angiistipes from Gaxias do Sul, Pinhal, and

Santa Maria, Brazil, and Santa Ee and Tucuman, Argen-

tina were also examined for comparison. These are de-

posited at SUGEN (lots 9021, 9022, 9029, 9030 and

9038). Most of the material is preserved in 70% ethanol,

although some of the material from Sao Paulo was fixed

in Raillet-Henry. Animals were killed by submersion in

filtered water, and kept in hermetically-closed recipients

in the refrigerator tor 48—72 liours. Prior to fixation,

specimens were photographed using a Ganon Digital

Power Shot SD630 and obseiwed alive. Preseiwed speci-

mens were dissected under a stereomicroscope. Anato-

mical illustrations were made using a camera lucida.

Pictures of the internal structures were obtained with a

DFG 280 digital camera attached to tlie stereornicro-

scope. Digital images were merged using Automontage

Pro (Synchroscopy) and Zeiss LSM Browser. Five radu-

lae and five jaws of the new species were e.xtracted and

examined under a scanning electron microscope LEO

440 at the Museu de Zoologia, Universidade de Sao

Paulo (MZUSP) (lots 8997, 9005, and 8983). The termi-

nology and anatomical characters described and illu-

strated are in accordance to those considered diagnostic

for Veronicellidae according to Tliome et al. (2006),

Gomes et al. (2006) and Gomes (2007).

RESULTS

Belocauhis willibalcloi new species

Diagnosis; Belocauhis iciJhbaldoi bears small projec-

tions similar to tubercles in the anterior region of the

glans. These can be uneven or arranged in two, three, or

more longitudinal rows. The glans presents a widened

basal region, narrowing towards the extremity and end-

ing in a digitiform margin. The penis base is short and

poorly defined. The accessoiy gland is completely im-

mersed in the tegument.

External Morphology: The length of the examined

specimens ranges from 2.19 to 7.1 cm, the total width

from 0.63 to 1.80 cm, the sole width from 0.16 to 0.52

cm, the left htyronotum width from 0.28 to 0.90 cm, and

the right hy[3onotum vHdth from 0.32 to 0.90 cm. Notum

coloration varies from brown to light or grayish-brown

(Figures 2-7). Mostly dark, slightly or strongly conspicn-

ous black puncta are noticealrle, usually scattered. In the

majority of the specimens, there is a lighter, median,

longitudinal stripe on the notum. In specimens \\4th a

darker notum, coloration pattern tends to be more

uniform with slightly conspicuous puncta and lighter

median, longitudinal stripe. The Inyonotnm and the

sole are beige. The external borders of the hyponotum

Page 36

THE NAUTILUS, Vol. 123, No. 2

Figures 2-7. External \iew of different forms of Bclocanhis ivillibahloi new species. 2-6. Dorsal \aew showang the variation

patterns ot eoloration. 7. Neutral \iew. Ahbrexiations: an: anus position: bp: black puncta; dp: dotted line delimitating tlie

pt'rinotnni: fp: female genital pore; Is: lighter, median, longitudinal stripe; nt: notnm; rd: region of darker pigmentation; rh: right

Inponotimi; so: sole.

Figures 8-11. Hadnia and jaw ol Beloamliis tvillilx/ldoi new species, S. Entire jaw (Lot 8997). 9. Middle part ol the radula (lot

8997). 10. Centi'al pai1 of the radula, showing lateral and centi'al teeth (lot 8983). il. Lateral teeth fit (lot 8983). Abbreviations: dc:

central teeth; dl: lateral teeth; I’e: rows ol central tec-tli; ts: transversal stripes.

F. P. Ohlweiler et aL, 2009

Page 37

can present a narrow stripe of darker pigmentation.

There is a dotted line delimitating the perinotnm in the

majority of the specimens. The sole is narrow and sur-

passes the posterior limit of tlie body when the animal is

moving. The wdth of the sole is always less tlian the

width of the right h)'}3onotinn, but never equals less than

half its width. The female genital pore is located ventral-

ly, in the posterior hall of the right hy|)onotum, while the

male genital pore is located in the anterior region, under

the inferior right tentacle.

Internal Moi-jjhology (Figures 8-2.3): Salivaiy glands

formed by large and well differentiated acini. Anterior

intestinal loop located behind the digestive gland anteri-

or lobe. One pair of pallial and one of pedal nen-'es run

both parallel and together to each other from the central

newous system toward the posterior portion of the body

cavity. They are united (all lour) throughout the entire

part of the path on the sole and are slightly separated at

the posterior region. The pedal aortic arteiy runs be-

tween the pairs of pallial and pedal neiwes. It arises trom

a bifurcation of the anterior aortic arteiy near the peri-

cardium and runs between the ner\?es until they termi-

nate at region posterior of the body. The pedal gland is

long, flattened, wdth a conspicuous, median, longitudinal,

lighter stripe. The posterior extremity of the gland is free

in the body cavity and receives veiy thin and short ducts

in its extremity. The rectum penetrates in the tegument

at the height of the female genital pore, behind the ovi-

duct (Figure 12). The anus opens in the posterior region

of the body, where the free end of the sole of the foot

protects it. An opercular membrane protects the anal

opening. The bursa copnlatrix is spherical to oval-shaped

and presents a short and thickened duct that opens into

an atrium, into female genital pore. The canalis junctor

penetrates in the bursa itself (not in the bursa copnlatrix

duct) (Figure 12). At the junction between the bursa

copnlatrix and the oviduct is a small, yellovnsh accessoiy

gland completely immersed in the tegument (Figure 12).

In some specimens, the accessoiy gland can be seen

by transparency through the tegument. The penial gland

presents a short and conical or long papilla with a termi-

nal mammila and 18—26 tubules (Figures 19—22). The

tubules located at the base of the papilla in the penial

gland are sinuous, not distinguished by size. Some pres-

ent the extremity or the median region bifurcated. Fi'om

the posterior region of the penial gland extends the re-

tractor muscle, which is connected to the penis retractor

muscle and together these are inserted in the tegument.

The penis is robust, with no spathe, with a small base

and glans with a wide basal region narrowang toward

the apical e.xtremity (asymmetrical arrow shaped penis)

(Figures 13-15). The distal extremity of the glans pre-

sents a digitiform margin (Figures 16-17). On each side

of the basal region of the glans tliere is a whitisli neiwure.

On the anterior region, the glans presents small projec-

tions shaped as minuscule tubercles (Figures 16-18).

These are arranged in twa:>, three or more longitudinal

rows or are unevenly distrihuted on tlie anterior regioTi

ihaJdoi new species (lot 8997). Abfi relations: ag: accessoiy

gland; ao: portion of spermovidnct; av: anterior region of o\i-

dnct; be: bursa copnlatrix; ca: ''carrefour"; cp: fertilization

pouch; cv: coiled region of ortdnct; db: hnrsa copnlatrix duct;

dd: posterior distal \'as deferens; dl: canalis junctor; dp: posteri-

or proximal \as deferens; ga: albninin gland; bg: hennafrodict

gland; po: portion of spermovidnct; pr: prostate; pv: posterior

region of oviduct; re: part of rectum; ve: seminal v'esicle.

of the glans. In young animals, the digitifoi'in margin of

the opening may be inconspicuous or not developed. In

young specimens the tubercles may also be absent, diffi-

cult to see, or ev'en appear as small depressions (future

tubercles). The penis and the penial gland are indepen-

dently surrounded in their ovvm muscular sheath. Both

sheaths fuse to form an anterior atrium near the male

genital pore (located at the base of the inferior right

tentacle) (Figure 23).

The hrowm jaw (Figure 8) is located at the dorso-

anteiior region of the buccal bulb. It foians an arch

composed of 21— 25 tran.sversal plates, partially covei'ed

and parallel to each other, which resemble lathes wdth a

keel-shaped dorsal region; plates are ornamented wdth

strong transversal and weaker longitudinal stripes. The

radula (Figures 9-11) is composed of lateral teetli on

each side ol a central tooth. The radnlar formula varies

from C/1-I-L52-55/2. The central teeth are small, trian-

gular, and unicuspid. The lateral teeth, larger than the

central teetli, are triangular and mricuspid. On tlie

dorsal region ol the apical extremity of the lateral teetli

the cuspid stands out from the rest ol the teeth. The

lateral teeth are triangular, but the cuspid is not as

prominent as in tlie teeth closer to the central teeth.

Page 38

THE NAUTILUS, \^1. 123, No. 2

Figures 13-15. Four cliHerent views of the penis from three ailult specimens of Belocaiihis wiUihaldoi new species. 1.3. Lot 8997.

14. Lot 8995. 1.5. f.ot 9006. Ahfireviations: ba: penis base; gl: gfans; ne; whitish nemire; rin; penis retractor muscle; vcl; anterior

vas deferens.

Measiirenienls (mm): Iloloty]ie; 7.1 cm of total

length, 1.8 cm of total width, 0.5 cm of’widtli of’tlie .sole,

0.9 cm ol width ol tlie left hvqionotiim and 0.9 of width

of the right hyponotnm. Paratyjies (four specimens): to-

tal length from 4.8 cm to 6.3 cm, total width from 1.1 cm

to 1.8 cm, width of the sole from 0.3 cm to 0.4 cm, width

of the left h)'|aonotnm from 0.5 cm to 0.7 cm, and width

of the right h)|ionotum from 0.6 to 0.7 cm.

F. P. Ohlweiler et al, 2009

Page 39

Figures 16-22. Penis and penial gland of Belocaiihis wiUibaJdoi new species. 16-17. Distal extremity of the glans (lot 8997).

18. Distal extremity of the glans (lot 8995). 19. Penial gland (lot 8995). 20. Penial gland (lot 8987). 21. Papilla of the penial gland

(lot 8997). 22. Papilla of the penial gland (lot 8987). Abbreviations: dm: digitiform margin; ina: mamilla; pp: papilla of penial gland;

tg: penial gland tubules; tr: tubercle row; tu: unevenly arranged tubercles.

Type Material: Holotype; MZUSP 87747; lour para-

types: MZUSP 87748 (one specimen), MZUSP 87749

(one specimen), MZUSP 87750 (tvv'o specimens).

Tjpe Locality: Brazil, Sao Panlo State, Sao Paulo, Btdrro

Parque Fernanda 1., 23° 40' 05.89" S, 49° 47' 26.66" W

Etymology: The specific name honors Dr. Jose VVilli-

baldo Thome for his great contriliution to the knowledge

of veronicellids and other terrestrial gastropods.

Distribution (Figure 1): Brazil; Minas Gerais State

(MG): Rio Acima; Sao Paulo State (SP): Caieiras, Guanilhos,

Osa.sco, Sao Paulo; Rio Grande do Sul State (RS): Eme.stina,

Gravahu, Igrejinlia, Porto Alegi'e, Riozinho, Sapiranga, and

Vila Maria; Santa Gatartna State (SC): Chapeco.

Habitat and Habit: The specimens of Belocatihts

willibaldoi collected in MG, SP, SC, and RS were found

in urban centers and surrounding areas, in gardens and

soil, under tree trunks, wood, plastic and other objects

Page 40

TPIE NAUTILUS, Vol. 123, No. 2

Figure 23. Penial complex of Belocaiilus icillibakloi new

species (lot 8997). AbbreUations: at: common atria; bg: mns-

cnlar sheath of the penial gland; bin: muscular sheath of the

penis; dm: middle vas deferens; fin: penis retractor muscle;

inr: penial gland retractor muscle; tg: penial gland tubules;

vd: anterior vas deferens. Size of complex: 1.8 cm.

on the ground. They are active mainly at night or, after

rainy periochs, during the day.

DISCUSSION

Tire new species described liere from southern and

southeastern Brazil is tyjrical of tlie genus Belocauhis

because it presents a penis shaped as an asymmetric

arrow and an accessoiy gland connected to the female

genital atrium, which are the two main characters of the

genus according to Hoffmann (1925) and Thome (1975).

Sarasimila also includes species with an arrow-shaped

penis (Gomes 2007), but penises in Sarasimila tend to

be more symmetrical and there is no accessoiy gland. In

addition, Sarasimila species are larger species and they

have a more oval shape, with a different pattern of exter-

nal jiigmentation. The new species has relatively small

size and is rejiresented by slender sings when compared

to other species loimd in southern and southeastern

Brazil. As Belocauhis an^iisiipcs, B. icillibalcloi is a small

species when compared to those of other Neotropical

genera. It is slender with a strongly narrow sole. Exter-

nally, both species cannot be distinguished from each

other: their coloration ranges from brown to beige or

gray, in different degrees of intensity. In B. willibalcloi

the wddth of the sole of the foot is smaller than the width

ol the right hyj^ronotum, as described by Santos and

Thome (1999) for B. angiistipes.

Internally, both species are also veiy similar. There

are no differences regarding the digestive, circnlatoiy,

and neiwous systems. The main differences are ob-

served in the male reproductive system, where the

main diagnostic features in Veronicellidae are found

(Semper, 1885). Small variations are also observed in

the accessory gland, radula, and jaw. The penis of

B. loillibaldoi is robust, with no spathe, with a small

base and a glans, with a wide base narrowing towards

the apical extremity with a digitiform margin. The an-

terior region of the penis bears minuscule tubercles,

which can be scattered or arranged in two, three, or

more longitudinal rows. The new species is distin-

guished from Belocauhis angiistipes which, according

to Pitoni and Thome (1981) and Santos and Thome

(1999), presents only a short, screw-shaped socket, dis-

tal extremity widened and truncated, and glans with

rhomboid extremity. In some specimens the glans can

be bilobed. In B. angiistipes, adjacent to the penis base,

the glans is projected backwards over itself (in one side

of the penis) (Eigures 24-26). In B. willibaldoi the

penis base is shorter and less defined than in B. angu-

stipes. In both species a labium is frequently formed on

the glans extremity, which folds back covering the

opening of the vas deferens.

The penial gland, in general, is similar in both species,

differing only in the number of tubules. In B. willibaldoi,

the penial gland presents from 18 to 26 tubules, while in a

B. angiistipes it presents 13—22 tubules (Pitoni and

Thome 1981; Santos and Thome, 1999). A terminal mam-

mila was obseiwed in the papilla extremity in B. willibal-

doi. Even though Pitoni and Thome (1981) and Santos

and Thome (1999) did not menhon the existence of a

mammila in B. angiistipes, it was observed in all speci-

mens examined in this study. This mammila is not, how-

ever, so conspicuous due to the fact that the papilla is

narrower when compared to that of B. willibaldoi.

The accessoiy gland of B. willibaldoi is completely

inserted in the tegument, differing from that of B. angii-

stipes in which, according to Thome (1975) and Silva

and Thome (1995), it can be totally or partially covered

by the tegument. According to Silva and Thome (1995),

the accessoiy gland releases a lubricant secretion, prob-

ably used during copulation, toward the female genital

pore, since it oj:)ens in this region.

The morphology ol the jaw of B. willibaldoi is similar

to that described for B. angiistipes by Thome and

Chaves (1997). The jaw and the radula of both species

are distinguished only by the number of jaw plates and

the number of teetli per row. According to those

authors, the jaw of B. angiistipes includes 19-22 plates,

while B. willibaldoi includes 21-25 plates. Based on the

characters presented liy Thome and Chaves (1997) for

F. P. Ohlweiler et al., 2()09

Page 41

Figures 24-26. Penis and penial gland oi Belocaulis angustipcs (lot 9030). 24. Four different views of penis. 25. Distal extreniit)'

of the glans. 26. Papilla of penial gland; ab: glans reflected hacWard over itself ; ba: penis base; eg: distal extremity of glans, without

tubercules; gl: glans; la: labium; pp: papilla of penial gland wthont a mainilla.

the raclula of B. cmgustipes, we notice that the radular

formula in B. wiUihaJdoi (C/1 + L52-55/2) is higher, since

B. angustipes presents C/1 + L34-38/2.

Belocaulus, which was regarded by Pitoni and

Thome (1981) as a monotvpic genus, includes another

species, B. willihalcloi, wliich occurrs in the states ol

Minas Gerais, Sao Paulo, Santa Catarina, and Rio

Grande do Snl. The known records ol B. angiistipes

reach the most southern point within the distribution

of die genus, including localities in Argentina and in

Brazil (states ol Rio Grande do Snl and Santa Catar-

ina). Although both species have been recorded from

Rio Grande do Snl and Santa Catarina, it is probable

that B. angtistipes occurs more to the south, while

B. a illihahloi more to the north. With the description

of B. wilUhaldoi, the distribution range of Belocaidiis- is

e.xtended to the states ol Minas Gerais and Sao Paulo,

Brazil.

Page 42

THE NAUTILUS, Vol. 123, No. 2

ACKNOWLEDGMENTS

Research performed w4th financial support Ifom Enn-

dayao de Amparo a Pesqnisa do Estado de Sao Paulo

(EAPESP), grant n° 06/56217-4.

LITERATURE CITED

Araujo, II. B. 19.52. Controle de lesnias. Revista Agronomica

16: 363-,366.

Baker, H.B. 1925, Nomenclature of Veronicellidae (Vaginuli-

dae). The Nautilus 39: 1,3-18.

Caballero, R., }.VV. Thome, K.L. Andrews, and A. Rueda. 1991.

Bahosas de Honduras (Soleolifera: Veronicellidae): Biologia,

Ecologia, DistribuHon, Descripcion, Importancia Econom-

ica, y Claves para su Identiiicacion. CEIBA .32: 107-126.

Caldeira, R. L., C.L.G.E. Mendonya, C.O. Goveia, II. Lenzi, H.,

C. Graetl-Telxeira, W.S. Dma, E.M. Mota, l.L. Pecora, A.

M.Z. Medeiros, and O.S. Caivalho. 2007. Eirst record

of molluscs naturally infected with An^iostwngijhifi canto-

ncnsis (Chen, 19,35) (Nematoda: Metastrongylidae) in

Brazil. Memoiias do Instituto Oswaldo Cniz 102: S87-iSS9.

Chiaradia, L.A., J.M. Milanez, C. Graeff-Tei.xeira, fW. and

J.W. Thome. 2004. Lesmas: pragas da agricultura

e ameaya a saiide humana. Agropecuaria Catarinense 17:

70-74.

Gomes, S.R., J.B. Picanyo, I.L.V Mendes, and J.W, Thome.

2006. A new species oi Simivthnhi from Northern Brazil.

Zootaxa 1329: 59-68.

Gomes, S.R. 2007. Eilogenia morfologica de Veronicellidae,

filogenia molecular de Fhi/lloaiulis Colosi e descriyao de

uma nova especie para a familia (Mollusca, Gastropoda,

Puhnonata), Ph D, dissertation, Universidade Federal do

Rio Grande do Sul, 175 pp.

Graeff-TeLxeira, C., J,W Thome, S.C.C. Pinto, L. Gamillo-

Coura, and ILL. Lenzi. 1989. PliijUocaiiUs variegatus -

an intermediate host of Aiigiostroiigi/hts cosiaricensis in

south Brazil. Memorias do Instituto Oswaldo Cruz 84:

6.5-68,

Graefl-Teixeira, C., V. M, Pinto, E.B. Jiuiior, and A, A. Agostini.

1994. Natural infection of PhijllocrmUs solciformis with

lavae morpliologycally similar to L2 oi' Agiostrongijhis cost-

aricen.sis. Memorias do Instituto Oswaldo Cruz 89: 121.

Hoffmann, H. f925. Die Vaginuliden. Ein Beitrag zur Kennt-

nis ihre Biologic, Anatomie, Systematik, geographischen

Verhreitnng und Phylogenie. Jenaiscli Zeitsch Natunvis-

senschaft 61: 1-374.

Laitano, A.C., f.P. Genro, R.A. Fontoura, S.S.L. Branco,

ILL,. Maurer, G. Graeff-Tei.xeira, J.M. Milanez, L.A.

Chiaradia, and J.W. Thome. 2001. Report of the ocur-

rence of Angiosfroiigi/liis costaricoisis in southern Brazil,

in a new intermediate host from the genns Sarasimihi

(Veronicellidae, Gastropoda). Revista da Sociedade Brasi-

leira de Medicina Tropical 34: 9.5-97.

Milanez, J.M. and L.A, Chiaradia. 1999. Lesmas: praga emer-

gente no Oeste Catailnense. Agropececuaria Catarinense

12: 1,5-16.

Pereira, ILF. and L.L Gonyalves. 1949, Caramujos, caracois e les-

mas nociv'os e meios de comhate. O Biologico 15: 65-73.

Pitoni. V. L. and J.W. Thome. 1981. Revisao do genero Belo-

cauhis Hoffmann, 1925 (Mollusca, Veronicellidae).

Rev'ista Brasileira de Biologia 4: 58.5-593.

Rambo, R.P., A. A. Agostini, and C. Graeff'-Teixeira. 1997. Ab-

dominal angiostrong)'losis in southern Brazil - Prevalence

and parasitic burden in molluscs intermediate host from

eighteen endemic foci. Memorias do Instituto Oswaldo

Cruz 92: 9-14.

Robinson, D.G. and R.G. Hollingsworth. 2004. Survey of slug

and snail pests on subsistence and garden crops in the

islands of the American Pacific: Guam and the Northern

Mariana islands. Part 1, The leatherleaf slugs (Family Ver-

onicellidae). Internal Report for the United States De-

partment of Agriculture ami the Government of the

Mariana Islands, Philadelphia, 11 pp.

Santos, E. 1959. Como combater as lesmas e caracois nocivos.

Bolctim do Cainpo, 15(120): ,3-5.

Santos, PH. and J.W. Thome. 1999. Chave Ilustrada para

tleterminayao pratica tlas cinco especies de Veronicellidae

com ocorrencia no Rio Grande do Sul (Mollusca, Gastro-

poda, Soleolifera). Cadernos EDIPUGRS 13: 1-22.

Semper, C. 1885. Landmollusken, In: CW Kreidel (ed.) Reisen

in Archipel der Philippinen, Wiesbaden, 36 pp.

SiKa, M.G.O. and J.W. Thome. 1995. Microanatomia da

vesi'cula seminal, complexo de fertilizayao e glandula aces-

soria de Belocaulus angmtipes (Heynemann, 1885) (Gas-

tropoda; Veronicellidae; Soleolifera). Biociencias 3: 95-

112.

Simone, L. R. 2006. Land and Freshwater Molluscs of Brazil

Lhiiversidade de Sao Paulo. Sao Paulo, 390 pp.

Thome, J.W. 1975. Os generos da familia Veronicellidae nas

Americas (Mollusca, Gastropoda). Iheringia 48: 3-56.

Thome, J.W. 1989. Annotated and illustrated preliminary list of

Veronicellidae of the AnOlles, and Central and Nortli Amer-

ica. Jouniiil of Medical and Applied Malacology 1: 11-28.

Thome, J.W. 1993. Estado atual da sistematica dos Veronicelli-

dae (Mollusca; Gastropoda) americanos, com comentarios

sohre sua importancia economica, amhiental e na saude.

Biociencias 1: 61-75.

Thome, J.W. and D.L. Chaves. 1997. Obsei-vaycies sobre a

constituiyao da mandibula e radula de Belocaulus angu-

stipes (Heynemann, 1885) (Mollusca; Gastropoda; Vero-

nicellidae). Biociencias .5: 219-229.

Tliome, J.W, S.R. Gomes, and R.S. Silva. 1999. Ocorrencia

e distrihuiyao da familia Veronicellidae Gray, 1840 (Mol-

lusca, Gastropoda) no Rio Grande do Sul, Brasil. Biocien-

cias, 7: 157-165.

Thome, J.W, S.R. Gomes, and J.B. Picanyo. 2006. Guia ilus-

trado: Os caracois e as lesmas dos uossos bosques e jar-

dins. Pelotas, Editora USEB, 123 pp.

THE NAUTILUS 123(2):43^8, 2009 Page 43

Sensory structures on the siphons of wood-boring bivalves

(Pholadidae: Xylophagainae: Xijlophaga)

A. J. Reft*

J. R. Voight

Department of Zoology

The Field Museum of Natural Histoiy

1400 S. Lake Shore Dr.

Chicago. IL 60605 USA

ABSTRACT

Deep-sea bivalves of Xylophagainae spend their entire post-

metamorphic lives boring into wood that has fallen to the

seafloor. Although their boreholes seemingly provide a pro-

tected, imperturbable habitat, scanning electron microscopy

reveals that the siphons of three species of Xijlophaga

examined cany elaborate structures that are interpreted as

chemoreceptors or mechanoreceptors. Sensory structures

occur on the siphonal surface of Xijlophaga oregona Voight,

2007, and X. multichela Voight, 2008. The lai'ge complex papil-

lae of X. multichela are scattered on the distal incurrent siphon

and arrayed in two longitudinal rows along its dorsal surface.

The distal incurrent siphon of X. oregona carries nrinute

structures, barely projecting above the surface, that are

crowned by tufts of cilia. Both siphonal openings of

X. microchira Voight, 2007, carry cirri. At the excurrent

opening, cirri have long cilia emerging from ternrinal pits. At

the incurrent opening, cirri form two rings. The imrer cirri

appear to be unique in that cilia emerge from between scales

that cover their inner surfaces. The structures observed irray

be useful in species taxonomy and systematics, but we suspect

that their elaboration is linked to predation pressure, which

might relate to depth distribirtion.

Additional keywords: Goblet organs, scanning electron micros-

copy, depth distribution, predation, deep-sea

INTRODUCTION

Deep-sea bivalves of the Xylophagainae spend their

post-inetamorphic lives using toothed ridges on their

shells to bore into wood that has fallen to the seafloor.

Only the siphons emerge from the resulting dead-end

boreholes. Although most bivalves suspeusion-feed by

extracting food from water moving across the gills, the

Author for correspondence, [email protected]; cirrrent address:

Department of Evolution, Ecology & Organismal Biology,

1315 Kinnear Rd., Tire Ohio State Univei'sity, Cohrmbus, OH

43212 USA.

small ctenidia and the labial palps of representatives of

Xylophagainae lack significant sorting mechanisms

(Purchon, 1941). Purchon (1941) proposed that these

animals ingest wood scrapings, which are digested with

the help of endosymbiotic Iracteria (Distel and Roberts,

1997).

This paper reports scanning electron microscope

(SEM) investigations of the siphons of three species of

Xijlophaga Turton, 1822, the most diverse genus of

wood-boring bivalves, with more than 50 named species

(Voight, 20()8). Sensory structures, known from the

siphons of a few shallow-water bivalves representing a

wide taxonomic range (e.g., Hodgson and Fielden, 1984;

Pekkarinen, 1986; Fishelson, 2000), are here documen-

ted in three congeneric species. Differences among the

structures in these species are largely consistent with

inferred ecological differences.

MATERIALS AND METHODS

Although most Xylophagainae species are known only

from their type localities, recoveiy of experimental wood

deployments from the deep Northeast Pacific (Voight,

2007) provided alrimdant specimens of the

Xylophagainae and allowed for SEM study of the

siphons of Xijlophaga oregona Voight, 2007 (Field Mu-

seum of Natural Histoiy, Chicago, FMNH 308705) and

of X. microchira Voight, 2007 (FMNH 309602), from

2211 m depth. Specimens were recovered inside a

lidded box on a subsea vehicle in 2003 and 2004, respec-

tively, fixed in 8% buffered formalin in seawater, and

transferred within 48 hours to 70% ethanol. No attempt

was made to relax the specimens prior to fixation. A

single lot of X. multichela Voight, 2008 (Scripps Institu-

tion of Oceanography Benthic Invertebrate Collections,

SIO-BIC Ml 1567) was collected by trawl in 1973 from

between 106 and 113 m depth, fixed in formalin and

later moved to 80% ethanol. All specimens were dehy-

drated in ethanol and then critical point-dried with COo.

Page 44

THE NAUTILUS, Vol. 123, No. 2

Each sample was sputter-coated with gold palladium iu a

Hummer sputter-coater and examined using a Zeiss Leo

Evo 60 Scanning Electron Microscope (SEM). Eeologi-

cal data reported here are from species descriptions

(Voiglit. 2007, 2008).

RESULTS

Images ol’ these specimens are clear, despite the absence

of specific preparation for SEM studies. The lack of

appropriate ffxation is not likely to liave resulted in the

dilferent moiphologies and distributions of structures

seen, although it may have induced some artifacts in

the fine details of tlie siphon surfaces. Therefore we

locus on the morphology ol the large structures. The

species, which all have an incomplete siphon (the excur-

rent is distinctly shorter than the incnrrent siphon), are

discussed below.

Xylophaga orecona (Figures 1—4), competitu'e

DOMINANT, DEPTH 1550-2211 M

For a \4ew of the whole siphon oi Xijlophaga oregona,

see Voight (2007, Figure 8A). The excurrent opening lies

under an apparently featureless C-shaped hood of tissue

near the posterior valve (Figure 1). The incurrent sipho-

nal opening of X. oregoua lacks cirri (Figure 2). The

incurrent siphon distal to the excurrent opening is

slightly dorsally flattened; low marginal walls border the

dorsal surface (Figure 1). The surface of the incurreut

siphon carries conceutric ridges (Figures 1, 2). Distally,

veiy small (12-18 pm diameter) structures (Figures 3, 4)

emerge apparently at random from the surface ridges.

Each structure has a terminal pit from which numerous

cilia emerge (Figure 4).

Xylophaga multichela (Figures 5-8), ecology

UNKNOWN, DEPTH 106-119 M

For a full view of the siphon of Xijlophaga mitkichela,

see Voight (2008, Figure lA). In X. miilfichela, the e.x-

curreut siphon opens near the posterior valve to form a

U-shapetl base of a longitudinal groove (Figure 5). Pa-

pillae border the groove and are scattered on the lateral

and ventral distal incnri'ent siphon (Figure 6). The open-

ing of the incurrent siplion lacks cirri; however, its tip is

morjrhologically distinct with concentric ridges, rather

than a smooth or papillate surface (F'igure 7). The papil-

lae bordering the groove (Figure 6) cany terminal cilia

(Figure 8) and form fringed lappets. The papillae on the

distal siphon also have terminal cilia and appear mor-

phologically similar to, but smaller than, those lateral to

the groove. Concentric folds (annnlations) on the papil-

lae (Figure 6) and differences in the visiliility ol the

papillae among specimens in light microscopy (unpub-

lished data) suggest that the cilia-topped papillae of the

lappets and on the distal siphon are retractable.

Xylophaga microchira (Figures 9-15), early colonist,

DEPTH 1550-2656 M

The siphon of Xt/lophaga microchira is circular in cross

section and lioth siphonal openings carry cirri (Figure 9).

The opening of the excurrent siphon is near the middle of

the siphon and is flanked by veiy long cirri (np to 420 pm;

Figure 10). Cilia emerge from pits at the tips of the cirri

(Figure 11). Although the surface of the siphon is ridged,

which may be due to contraction, structures such as those

seen in Xi/Iophaga oregona (Figure 3) appear to be absent.

The incurrent opening has tM'o concentric rings of

cirri (Figure 12). The outer cirri are smooth whereas

the inner cirri, especially their inner surfaces, appear

scaly (Figures 13, 14). Cilia emerge from between the

scales (Figures 13, 14) and are densest at the peripheiy

of each cirrus (Figures 14, 15).

DISCUSSION

All structures documented here, whether on the sipho-

nal surface or at the tip of a cirrus (= tentacle sensu

Fishelson, 2000), share a terminal opening with an

emergent tuft of equal-length cilia. The absence of a

long central flagellum leads us to interpret these struc-

tures as sensoiy organs, reportedly common in bivalves

(Fishelson, 2000). Distinguishing between mechanore-

ceptor and chemoreceptor cells is difficult (Hodgson

and Fielden 1984), even if neuronal connections

are traced (Fishelson, 2000). Earlier comparison of

transmission electron microscopy (TEM (-documented

ultrastructure of sensory cells to that of known chemor-

eceptors or mechanoreceptors was said to identify mo-

dality of the cells (e.g. Jouin et ah, 1985; Chia and Koss,

1989). However, variability in the fine structure of sen-

soiy cells led Schaefer (2000: 208) to question this meth-

od. Behavioral and physiological data are integral to

assign function to sensoiy cells (Schaefer, 2000; Zhadan

et ah, 2004). Given that these representatives of Xijlo-

phaga live inside wood on the ocean floor, at depths of

over 2 km, and no material was suitably fixed for TEM

study, the modality of the sensoiy structures documen-

ted here cannot be assigned. In general, chemoreceptors

have been considered to be the most abundant sensoiy

structure on bivalve siphons (Fishelson, 2000), however,

the goblet organs of Macoma bakhica (Linnaeus, 1758)

maybe mechanoreceptors (Pekkarinen, 1984).

These SEM images reveal that the distribution, shape

and size of the sensoiy structures (Figures 1-15) differ

distinctly among these wood-boring bivalves. Xi/Iophaga

oregona (Figures 1-4) and X. mnitichela (Figures 5-8)

share aii excurrent siphon that is truncated near the

shell (Voight, 2007, 2008); their sensoiy structures lie

on the siphonal integument, in contrast with those on

cirri at siphonal openings in X. microchira, a species with

tlie excurrent opening near the middle of the siphon

(Figures 9-15). These data are consistent with the hy-

pothesis (Voight, 2007), based on dilferences in siphonal

A. J. Reft and J. R. Voiglit, 2009

Page 45

Figures 1-4. Xijlophaga oregona. 1. Excurrent siphon. Black arrow indicates hood ot tissue over the opening ot the excurrent

siphon. White arrows indicate marginal ridges that border a longitudinal flat area of the dorsal siphon. Scale bar = 200 pm. 2. Tip of

incurrent siphon. Arrow indicates the edge of the siphon; note the lack of cirri. Scale bar = 40 pm. 3. Surlace ol distal incurrent

siphon. Note tlie ridged appearance and the round projections indicated by arrows. Scale bar = 20 pm. 4. Finer detail ot a round

structure indicated in Figure 3. Scale bar = 6 pni.

allometiy and overall appearance, that the truncated

excurrent siphons are not nni(|uely derived.

The round structures of Xijlophaga oregona (Fig-

ures 3, 4) strongly resemble the goblet organs detailed

by Pekkariuen (1984 Figures 8 and 11, 1986 Figure 5)

in the veneroid bivalve Macoma halthica, which is only

distantly related to the niyoid Xijlophaga species con-

sidered here. In both species, the small (10-20 pm)

structures are associated with ridges on the distal in-

current siphonal surface, and have long cilia that emerge

from a central opening (Figure 4) (Pekkariuen, 1984,

1986). The subtle sliape differences coukl relate to

differences in fixation. The goblet organs of M. balthica

form six longitudinal rows that correspond to the

course of the main longitudinal nen-'es (Pekkariuen,

1984, 1986); sensoiy .structures iu X. oregona appear

to be randomly arranged. Apparent goblet organs,

termed ty|:)e III seusoiy organs by Hodgson and

Fielden (1984) and Ansell et al. (1999), "have also

been obsewed on incurrent siphons of the veneroid

Donax trnncnlns Linnaeus, 1758 (Fishelson, 2()0(),

Figure 5H).

Page 46

THE NAUTILUS, Vol. 123, No. 2

Figures 5-8. Xi/loj)haga iwdticlwla. 5. Opening oI tlie exeurrent siplion. Arrow indicates longjitndinal gi'oove originating at the opening.

Sc;ile har = 40 pm. (i. Two rows ol papillae (white aiTows) ionn fi'inged lappets lateral to groove that extends distally from the opening ol tlie

excurrent siphon. Black arrows indicate a row ol papillae inferior to the lappets. Scale har = 40 pm. 7. Tip of incmrent .siphon. Note tlie

ridged snriace at the siphonal tip and randomly scattered cirri (airows). Scale har = 80 pm. 8. Finer detail of a lappet from the distal

incnrrent siphon. Note the tnft ol equal-length cilia emerging from the center. Aitows indicate folds on a cirrus. Scale har = 8 pin.

Bivalve.s living in liigh-em'rgy habitats with lieaw sed-

imentation tend to have elaborate, branelied cirri on tlie

incnrrent siphon (Eislielson, 20()0). Turner (1971) sug-

gested that the elaborate cirri ol sliipwonns (teredinids)

lorm a sieve across tlie incnrrent opening to protect the

animal Irom debi'is. Ilowcwer, Lopes and Narelii (1998)

found that the tentacles did little themselves to block the

entrance to the inenrrent siphon in the teredinid Nausi-

lora fiislicnla (|ellreys, I860), ratlu-r contraction of the

siphon base sensed to block the opening. The incnrrent

siphon ol Xi/Iopha^a microchira carries structures highly

compatible with a .sieving Innction. In this species, the

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Page 47

Figures 9-15. Xijlophaf^a inicmciiira . 9. Siphon. Wliite arrow incliealc.s the ojiening ol (he inciirreiit .siphon. Mack arrow indicates

the opening ot the excnrrent siphon. Note ridgetl suriace ol siphon. Scale Itar = 400 pin. 10. Flap ol inega-cirri at excni'rent sijrhonal

opening. Tip ol cirrus indicated hy the arrow. Scale bar = SO pm. 11. Finer detail ol cirrus noted in Figure 10. Scale bar = 0 pin. 12.

Incnrrent opening with hvo rings ot cirri. I indicates inner cirri; () indicates outer cirri. Scale bar = SO. 1,3. Finer detail ol inner

cirrus. Note that the inner snriace ol the inner cirrus (1) dilters Iroin that ol tlie outer cirrus (()). Scab' bar = 20 pin. 14. Finer detail

ol an inner cirrus (I): outer cirrus (O). Note the scalv suriace aiul the arch ol cilia indicated by arrows. Scale bar = S pin. 1,5. Finer

detail of cilia ol inner cirrus. Arrows indicate area Ironi which cilia emerge. Scale bar = 4 pm.

Page 48

THE NAUTILUS, Vol. 123, No. 2

inner surfaces of the inner cirri appear scaly (Figures IS-

IS); cilia emerge from between the scales and at their

margins. The distribution of cilia is snch that if these

cirri were bent inwards to block the incnrrent opening,

penetration of the opening would perturb the maximum

number of cilia, generating the maximum sensoiy stimu-

lus. Xijlophaga microchira is considered to be

specialized for rapid colonization of new wood-falls and

shows frequent damage on the incnrrent siphon consis-

tent with cropping by predators (Voight, 2007). Poly-

chaetes are suggested to enter the incnrrent siphon of

wood-boring bivalves (Dean, 1992); this siphonal senso-

iy system may speed the bi\ alve’s defensive retraction of

the siphon.

Excnrrent siphons of most bivalves typically bear rela-

tively smaller, simpler extensions than do the incnrrent

siphons (Fishelson, 2000). However, large, fairly complex

sensory receptor-bearing stnictnres occur at the excnr-

rent opening Xijlophaga microchira (Figure 10, 11). In

X. multichela, papillae on the dorsal incnrrent siphon

bordei' the groove that appears to be a continuation of

the excnrrent siphon (Figure 6), although the papillae

have been illustrated veiy near the excurrent opening in

similar species (Turner, 2002). Near the incnrrent open-

ing of this species, smaller papillae are scattered over the

surface of the incnrrent siphon (Figure 7). The compara-

tively elaborate sensoiy arrays near the excnrrent sipho-

nal openings of these species remain enigmatic.

Given the broad ecological similarities of these wood-

boring species and their congeneric status, differences

documented here are counter-intuitive; they may reflect

fine-scale ecological differences or possibly depth distri-

bution. The species with most sensoiy structures, Xijlo-

phaga miilticliela, lives around 106 m depth; sensory

input may help it survive in the more predator-rich con-

tinental shelf depths (Vermeij, 1987), as hypothesized by

Voight (2008). Specimens of X. oregona have the fewest

sensoiy structures. Boreholes of this species are lined

with fecal chimneys, which, in addition to lowering oxy-

gen tension (Voight, 2007), may minimize or confound

chemical cues or muffle mechanical stimulation. In ex-

tremely high densities, however, siphons of this compet-

itively dominant species can extend well beyond the

wood, conceivably allowing chemical cues to be received

(personal observation, JRV). No simple environmental

variable appears to be clearly correlated with the com-

plex sensoiy structures documented here.

ACKNOWLEDGMENTS

The captains and crews of the IW Thomas G. Thompson

and the lUV Atlantis and the pilots of the ROV Jason

and the HOV Alvin made the collections possible.

National Science Foundation grant DEB-0103690 to

|RV supportetl this research. We thank B. Strack lor

SFM assistance. E. Rodriquez helped prepare the

images. A. Lindgren and M. Daly provided lielpfnl

comments on tlie text.

LITERATURE CITED

Ansell, A. D., R. Hawey, and C.R Giindier. 1999. Recoveiy from

siphon damage in Donax vittafus (Da Costa) (Bivalvda:

Donacidae). Journal of Molluscan Studies 6.5; 223-232.

Cilia, F. S. and R. Koss. 1989. The fine structure of the newly

discovered propodial ganglia of the veliger larva of the

nudibranch Onchidoris hilamellata. Cell and Tissue Re-

search 256: 17-26.

Dean, 11. K. 1992. A new arabellid polychaete living in the

mantle cavity of deep-sea wood boring bivalves (Family

Pholadidae). Proceedings of the Biological Society of

Washington 105: 224-232.

Distel, D.L. and S.J. Roberts. 1997. Bacterial endosymbionts

in tlie gills of the deep-sea wood-boring bivalves Xijlo-

phapa atlantica and Xi/lophapa woshiiiPtona. Biological

Bulletin 192: 253-261. ‘

Fishelson, L. 2000. Comparative moiyhology and cytology of

siphons and siphonal sensoiy organs in selected bivalve

molluscs. Marine Biology 137: 497-509.

Hodgson, A. N. and L.J. Fielden. 1984. The structure and

distribution of peripheral ciliated receptors in the bivalve

molluscs Donax serra and D. sordidns. Journal of Mollus-

can Studies .50: 104-112.

Jouin, C., C. Tchemigovtzeff, M.F. Baucher, and A. Toulmond.

1985. Fine stmcture of probable mechmio- and chemorecep-

tors in tlie caudal epidennis of tlie lugwonii Arcin'ro/r/ marina

(Annelida, Polychaeta). Zoomoiphology 105: 76-82.

Lopes, S.C.B.C. and W, Narchi. 1998. Functional anatomy of

Nausitora fusticula (Jeffreys, 1860) (Bivalvia: Teredini-

dae). The Veliger 41: 274-288.

Pekkarinen, M. 1984. Regeneration of the inhalant siphon and

siphonal sense organs of brackish-water (Baltic Sea)

Macoma balthica (Laniellibranchiata, Tellinacea). Annales

Zoologici Fennici 21: 29-40.

Pekkaiinen, M. 1986. Histology of the siphons of Macoma balthica

(Bivalvia: TelHiiidae). Annales Zoologici Fennici 23: 77-95.

Purchon, R.D. 1941. On tlie biology and relationsliips of tlie laniel-

libranch Xijlophaga domain (Turton). Journal of tlie Maiine

Biological Association of tlie United Kingdom 25: D39.

Schaefer, K. 2000. The adoral sense organ in protobranch

bivalves (Mollusca): comparative fine structure with spe-

cial reference to Nuctila nucleus. Invertebrate Biology

119: 188-214.

Turner, R.D. 1971. Biology of marine wood-boring molluscs.

In: Jones, E.B.C. and S.K. Eltringham (eds.) Marine

borers, fungi, and fouling organisms of wood. Organiza-

tion for Economic Co-operation and Development, Paris,

pp. 2.59-301.

Turner, R.D. 2002. On the subfamily Xylophagainae (Eaniily

Pholadidae, Bivalvia, Mollusca). Bulletin of the Museum

of Comparative Zoology 157: 223-307.

Veniieij, C.J. 1987. Evolution and escalation: An ecological histo-

ry of Life. Princeton University Press, Princeton, 527 pp.

Voight, J.R. 2007. Experimental deep-sea deployments reveal

diverse Northeast Pacific wood-boring bivalves of Xylo-

phagainae (Myoida: Pholadidae). Journal of Molluscan

Studies 73: 377-391.

Voight, ].R. 2008. Deep-sea wood-boring bivalves of Xijlo-

phaga (Myoida: Pholadidae) on the Continental Shelf

Journal of the Marine Biological Association of the United

Kingdom 88: 1459-1464.

Zhadan, P. M., A.V. Sizov, and S.S. Dautov. 2004, Ultrastructure

of the abdominal sense organ of the scallop Mizuchopecten

ijessoensis (Jay). Cell and Tissue Research 318: 617-629.

THE NAUTILUS 123(2):49-52, 2009

l^age 49

An unusual new genus and a new species of Buccinulidae

(Neogastropoda) from the Magellanic Province

Yu. 1. Kantor

A.N. Severtzov Institute of Ecology and Evolution,

Russian Academy of Sciences,

Leninski prospect 33, Moscow 119071, RUSSIA

[email protected]

Guido Paslorino

Museo Argentino de Ciencias Naturales

Av. Angel Gallardo 470 3° piso lab. SO

C1405DJR Bnenos Aires, ARGENTINA

gjrastorino@macn,gov.ar

ABSTRACT

A new genus and species of the family Buccinulidae is de-

scribed from the Southwestern Atlantic in Argentine waters.

Jern/biiccinitm maJvinense new genus and species combines

the conchological characters of Fasciolariidae wath the radula

of Buccinulidae.

AcIcJifionaJ keywords: Gastropoda, southwestern Atlantic,

Jerrybiiccimnn maloinense, new genus, new species

INTRODUCTION

The intensive collecting efforts of the United States

Antarctic Program (USAP) in the Antarctic and Magel-

lanic regions yielded rich collections, which stored at

tlie National Museum of Natural Histoiy, Smithsonian

Institution.

These collections have been the source of a vast

nnrnlier of new species of mollnsks described in several

papers and monographs (e.g. Dell, 1990; Harasewych

and Kantor, 1999; Pastorino, 1999; Harasewych et ah,

2000; Pastorino and Harasewych, 2000; Pastorino,

2002; Harasewych and Kantor, 2004; Harasew)X‘h and

Pastorino, in press). There remain to be studied, how-

ever, a number of species with novel combinations of

anatomical features and shell moqiliology. One of the

new species collected off the Falkland Islands (Islas

Malvinas) demonstrated the nnnsnal combination of a

fasciolariid-looking shell with a bnccinnlid radula.

In this paper we describe as new a species that pos-

sesses radnlar and conchological characters that pre-

clude its inclusion into any presently recognized genus

of Bnccinoidea.

MATERIALS AND METHODS

The specimens here described are housed in the collec-

tion of the National Museum of Natural History, Smith-

sonian Institution, Washington, DC (USNM). They

were collected by IW Eltanin. The shells v\4th dried-

out bodies were re-hydratated to facilitate dissections.

After cleaning with diluted bleach, air-dried, mounted

on glass slides, and coated with gold-palladium radnlae

were studied with help of a scanning electron micro-

scope at USNM. Most photographs were taken using a

digital camera. All images were digitally processed.

SYSTEMATICS

Class Gastropoda Cuvier, 1797

Order Neogastropoda Wenz, 1938

Superfamily Bnccinoidea Rafinesque, 1815

Family Buccinulidae Finlay, 1928

Genus Jern/biiccintiin new genus

Type Species: Jern/hiiccimtm malvinense new .spe-

cies, by original designation. (Currently the only species

included into the new genus is the Rpe species.)

Description: Shell lusilorm, with tall spire and long

attenuated siphonal canal. Protoconch pancispiral, orna-

mented by spiral threads and closely spaced axial ribs;

protoconch-teleoconch transition veiy weak, marked by

the appearance of the axial folds. Spiral sculpture of low

and narrow spiral ribs, raised, and rounded on the top

keel that delimitates the shell base. A.xial sculpture of

growth lines and high, closely spaced axial folds. Radula

triserial, with rectangular nnicnspid rachidian teeth and

tricuspid lateral teeth with long, stout basal plates.

Etymology: The genus is named after onr colleague

and mutual friend Miroslav (Jeny) Harasewych, cairator

ol mollnsks at the National Museum of Natural Histoiy,

S mithsonian Institntioi i .

Jern/biiccinum malvinense new species

(Figures 1-12)

Description: Shell strong, fusiform with tall spii'e and

long attenuated siphonal canal, of 2.5 protoconch and

sliglitly over 5 teleoconch whorls. Protoconch pancis-

piral, evenly rounded (Figure 5), ornamented by 5 un-

evenly spaced spiral threads and closely spaced thin, but

Page 50

THE NAUTILUS, Vol. 123, No. 2

Figures 1-10. fcrnjhnccinum nuilviiieiise newspecie.s. 1-5. Ilolotvpe, USNM S9ScS47, Falkland Island.s (l,sla.s Malvinas), 52°00’ S,

56°3fi' W, RA^ Ei.tanin, cruise 7, sta, 55S, 14 Mar. 1963, 646-(S45 in. 1—3. Shell. 1. Apertiiral view. 2. Lateral \4ew. 3. Dorsal view.

4. Operciilinn. 5. Protoconcli. 6-8. Paratvpe, USNM <898774, Falkland Islands (Islas Malvinas), W ol Beauchene Island, 53°06' S,

59°24‘ M’, RA^ Ei;r,\NiN, cruise 6, sta. 340, 03.12.1962, 567-578 in. 6-8. Shell. 6. Apertiiral view. 7. Lateral view. 8. Dorsal view.

9-10. USNM 887765, off Cape Horn, 56°06’ S, 66°19’ \V, RA' Eltanin, cruise 9, sta. 740, 18 Sep. 1963, 384-494 in, sliell length =

8.1 linn. 9. Apertiiral view ol' the sheik 10. Protoconch. Figures 1-3 ami 6-8 at same scale, scale bar = 1 cm. Figures 4, 5 at same

scale, scale har = I nun.

Yu. I. Kantor and G. Pastorino 2009

Page 51

Figures 1 1-12. Radula of Jernjbucciiuim malvineiise new species. 1 1. Dorsal \dew of the central portion of tlie raclular membrane.

12. Bending plane of the membrane. The basal projection of the lateral tooth is marked by an arrow.

di.stinct, axial ribs. Protoconch-teleoconch transition not

clear, marked by appearance of axial folds. Protoconch

diameter around 2.1 mm, exposed protoconch height

1.75 mm. Teleoconch whorls strongly convex, slightly

angulated at periphery, separated by shallow, slightly

adpressed suture. Spiral sculpture of shaip, low, and

narrow spiral ribs, separated by slightly wider inter-

spaces. Twenty-two ribs on penultimate whorl, upper

one adjoining suture, slightly vender than other ribs. Last

whorl with raised keel, rounded in cross-section. Last

whorl wnth 23 ribs above keel that delimits shell base,

ribs below keel more pronounced, 28 in total on shell

base and canal. Axial sculpture of raised growth lines

that produce reticulated structure while crossing spiral

ribs and high, closely spaced axial folds, 13 on body

wdiorl and 13 on the penultimate whorl. Folds protrude

from suture to suture on spire whorls and from suture to

keel on the last whorl. Aperture wide, oval, constituting

0.34 of shell length (without siphonal canal). Outer lip

evenly rounded and slightly reflected outward. Inner lip

with narrow callus extending to parietal wall. Siphonal

canal well defined, long, constituting about 0.17 of shell

length, slightly cuiwed to left but not crossing shell axis.

Shell covered by thin, light-yellow periostracum. Shell

color under periostracum uniform off-white.

Operculum ovate (Figure 4), elliptic, with subcentral

nucleus, external surface covered by concentric growth

lines where new growth partially overlap old ones,

resulting in lamellose surface, particularly on internal

margin. (Measurements as in holoty^De.)

Raclular ribbon (Figures 11-12) long (2.98 mm, 0.36

AL), narrow (^130 pm), triserial, consisting of 90 rows,

most posterior 9 rows nascent. Rachidian teeth narrow

(~40 pm), with anteriorly very slightly arched rectangu-

lar basal plate and single shaqr cusp. Lateral teeth with

long, stont basal projection (marked by an arrow on

Figure 12), attached at acute angle (~50°) to axis of

radular ribbon, with 3 cusps, outer largest and central

shortest situated closer to the ijmer cusp.

Type Material (Figures 1-8); (Measurements in

Table 1) Holotyjre (Figures 1-5), USNM 898847, Falk-

land Islands (Islas Malvinas), 52°00’ S, 56°36’ W, RA^

Eltanin, cniise 7, sta. 558, 14 Mar. 1963, 646-845 m.

Parat)pe (Figures 6-8), USNM 898774, Falkland Islands

(Islas Malrinas), W of Beauchene Island, 53°06'S,

59°24'W, IW Eltanin, cmise 6, sta. 340, 12 Mar. 1962,

567-578 m.

Type Locality: Falkland Islands (Islas Alalvinas),

52°00’ S, 56°36’ W, WV Eltanin, cruise 7, sta. 558, 14

Mar. 1963, 646-845 m.

Other Material Examined: Two specimens (USNM

887765) (Figures 9-10) collected off Cape Horn, 56°06’ S,

66° 19' "W, IW Eltanin, cnii.se 9, sta. 740, 18 Sep. 1963,

384-194 m.

Remarks: The paratyjve (Eigures 6-8) is a sliglitly

smaller specimen, othenvise in all respects it is similar

to holot)pe. One additional juvenile, a dead-collected

specimen (shell length = 8.1 mm) (Eigures 9-10) is rath-

er similar in shell sculpture and outline to the t)'|5es, Imt

Table 1 . .Shell dimensions of the holotyjie and paratype of

Jeni/huccimtin malvinense new species, measurements in mm.

Page 52

THE NAUTILUS, Vol. 123, No. 2

Figure 13. Geographical distribution of Jernjbitccinum mal-

vincnsc new species. Dashed line indicates .500 m isobath.

Symbols: 0 = t\pe localit)', ♦ = other material e.Kamined.

differs in having a smaller protoconch (e.xposed lieight

1.12 mm vs 1.7.5 in bolohpe).

Distribution (Figure 13): The species is knowi off

Falkland Islands (Islas Mahinas) at the depth 567-845

m and oil Cape Horn in 384-494 m.

Etymology: The species is named after the type lo-

cality, Islas Malvinas (Falkland Islands.)

DISCUSSION

The l adnlar characters undoubtedly place Jcrn/hiiccinum

in Bnccinnlidae, but the subfamilial allocation is not

clear. The single cusp of the rachidian tooth and long

basal projection of the lateral tooth suggest the affi-

nities with the subfamily Cominellinae and particu-

larly with the Antarctic species of “Pareuthiia", i.e.,

P. plicatiila Thiele, 1912, P. innocent; (Smith, 1907),

and P. hoshiaii Nnmanami, 1996 (see Nnmanami,

1996). Harasew\/ch and Kantor (2004) obsen^ed that

the Antarctic representatives of this genus differ mark-

edly ill radular moqiholog)^ from those of Magellanic

distribution, incinding the t)pe species Parcuihria

j)liin)hea (Philippi, 1844), which has a tricuspid rachi-

dian tooth. In any case, in contrast to the condition

found in lern/I)uccimmu all representatives of Cominel-

linae have tlie bicuspid lateral teetli.

The shell in the new species does not have analogues

among Antarctic and Subantarctic Buccinulidae. It

shows some resemblance to representatives of Fasciolar-

iidae, mostly due to the long and nearly straight siphonal

canal and characteristic a.xial sculpture. Thus, on first

approach, Jern/buccinum malvinense appears to com-

bine characters of both families.

ACKNOWLEDGMENTS

This work was supported in part by a Research Award

from the NSF-USAP United States Antarctic Program

Grant [ANTO636408].

LITERATURE GITED

Dell, R.K. 1990. Antarctic Mollusca, with special reference to

tlie fauna of the Ross Sea. Royal Society of New Zealand

Bulletin 27, 311 pp.

Harasewycb, M.G. and Yu. I. Kantor. 1999. A revision of the

Antarctic genus Chlanidota (Gastropoda: Neogastropoda:

Buccinulidae). Proceedings of the Biological Society of

Washington 112: 25.3-302.

Marasewych, M.G., Yu. I. Kantor, and K. Linse. 2000.

Parahucciiuiin, a new genus of Magellanic buccinulid

(Gastropoda: Neogastropoda), with a description of a

new species. Proceedings of the Biological Society of

Washington 113: .542-560.

Harasewych AI.G. and Yn. 1. Kantor. 2004. The deep-sea Bnc-

cinoidea (Gastropoda: Neogastropoda) of the Scotia Sea

and adjacent abyssal plains and trenches. The Nautilus

118: 1-42.

Harasewych, M.G. and G. Pastorino. (IN PRESS) Twplionella

(Gastropoda: Muricidae), a new genus from Antarctic

waters, with the description of a new species. The Veliger.

Nnmanami, II. 1996. Taxonomic study on Antarctic gastropods

collected by Japanese Antarctic Research E.xpeditions.

Memoirs of National Institute of Polar Research Series

E (Biology and Aledical Science) 39: 1-244.

Pastorino, G. 1999. A new species of gastropod of the genus

Trophoit Alontfort, 1810 (Mollusca: Gastropoda: Murici-

dae) from subantarctic waters. The Veliger 42: 169-174.

Pastorino, G. and M.G. Harasewych. 2000. A revision of the

Patagonian genus Xi/inenopsis Powell, 1951 (Gastropoda:

Muricidae). The Nautilus 114: 38-58.

Pastorino, G. 2002. Two new Trophoninae (Gastropoda: Mur-

icidae) from Antarctic waters. Malacologia 44: .35.3-361.

THE NAUTILUS 123(2):53-5S, 2009

Page 53

A new species of Parijphantopsis (Gastropoda: Pulmonata:

Charopidae) from Crater Mountain, Simbu (Cliimbu)

Province, Papua New Guinea

John Slapcinsky

Florida Museiiin of Natural Histoiy

and Uepartnient of' Zoology

University of Florida

Gainesville, FL 32611 USA

[email protected]

ABSTRACT

Pan/phantopsi.s braclleifi new species is described from a sub-

montane forest near Crater Mountain Biological Research

Station in Simbu Province, central Papua New Guinea. It is

distinguished from its congeners by the combination of its

large size and shaqily pointed, non-overlapping pei'iostracal

processes that are retained to maturity. It shares similarities

and is probably closely related to otlier large Paiiipliantopsis

Thiele, 1928, that have angled to carinate shell margins with

long periostracal processes and central and lateral radular

teeth that liave mesocones originating from the center of their

basal plates. It appears that much of New Guinea’s highly

endemic terrestrial snail fauna remains to be discovered. It is

imperative that the biodiversity of large groups of ta.\a is docu-

mented because this information will be crucial in efforts to

preserve rapidly diminishing rainforest habitat.

Additional keywords: Terrestiial snail, puhnonate, rainforest,

taxonomy

INTRODUCTION

The piilmonate snail family Charopidae Hutton, 1884,

was once considered to be a minor component ol tlie

terrestrial inolluscan fauna of New Guinea, in contrast

to the group’s spectacular radiations in the oceanic

islands of tlie Pacific (Solem, 1983: 305). However, re-

cent snrx'eys in Papna New Guinea suggest that inade-

quate sampling, rather than low diversit)', is tlie cause of

the perceived paucity of charopid species in New Guin-

ea (Slapcinsky, 2005). Panjphantopsis Thiele, 1928, the

most diverse genus of charopids in New Guinea, consists

of 26 described species distributed from Papua (Irian

Jaya) to the Louisiade Archipelago and New Britain.

Solem (1970) rexdewed the 14 species of the genus then

known, redescribing all species except those described

or reviewed by van Benthem Jutting (1964). However,

nearly half of this radiation has been described only

recently (Slapcinsky, 2005, 2006; Slapcinskw and Lasley,

2007), and it is clear that continued sampling, especially

in New Guinea's poorly sampled inonntain ranges, will

uncover many additional species. All Parijphantopsis

species are restricted to single mountain ranges, each

often supporting several Panjphantojisis species. The

most widely distributed species, PanjpJiantopsis ijawii

Slapcinsky, 2005, ranges 50 km along the mountains of

the East Gape Peninsula in extreme eastern Papua New

Guinea. Synapomoqrhies in Panjphantopsis from the

same mountain ranges suggest they have speciatetl on a

fine geographic scale (Slapcinsky, 2005).

Most Partjjdiantopsis species occur in moist and mossy

montane and sub-montane forests or at lower elevations

in hill forest along stream valleys. Restriction to stable

moist habitats and microhabitats might promote genetic

isolation and rapid speciation in the group. Species of

Panjj)hantopsis are unusual in being dinrnally active

(Slapcinsky and Lasley, 2007) and haxing reduced shells

of approximately three whorls compared to at least four

whorls in most other charopids. Whorl reduction

is associated wth enlargement of the shell aperture,

modifications of the kidney, ami retlnced space in the

pallial cavity for retraction of tlie visceral hump (Solem,

1970). All of these traits may make Parijjihantopsis espe-

cially susceptible to desiccation if their moist forest habi-

tats are altered. Besides shell whorl reduction and the

associated changes in pallial organs, Parypiuintopsis is

also diagnosed by the following shell .synapomorphies:

protoconch sculpture of axial and spiral riblets that usu-

ally coalesce forming spiral I'ows of pits, growth lines that

are accentuated with rib-like periostracal extensions

that often bear processes at the shell margin, and sheiks

that are not openly nmbilicate.

Between 1990 and 1993, the Fknida Museum of Nat-

ural Histoiy received a collection of terrestrial snails col-

lected by Andy Mack and lOebra Wb iglit during studies at

Grater Mountain Biological Research Station. This col-

lection included a new species ol Partjjihanfopsis which

Page 54

THE NAUTILUS, Vol. 123, No. 2

142° E 144° E 146° E 148°E

Figure 1. Map ot eastern New Guinea showdng the t\pe

locality ot Pan/phaiifopsis hiricllei/i new species

is described here. Crater Mountain Biological Research

Station is located in southeastern Sinibu Province, Papua

New Guinea, approximately 78 km SSW of Goroka and

1 1 km E of Haia Village, at 6.72° S, 145.09° E (Eignre 1).

The research station is located on the southern slope of

Crater Mountain, an arcuate chain of peaks reaching

3000 m or more in elevation and formed from an e.xten-

sively eroded strato\ olcano last active in the late Pleisto-

cene or early Holocene (Mackenzie and Johnson, 1984).

The topography of Crater Mountain is extreme, \\4th

vertical cliffs and frequent seismic actixity, which, com-

hined \\4th ample rainfall results in munerons treefalls

and landslides, lead to heterogeneous habitats and

microhabitats. Geographic and habitat heterogeneity

may contribute to the floristic richness of the area, the

richest site known in New Guinea and among the richest

in the world (Wright et ak, 1997). A 1 ha plot contained

228 tree and liana species with no strongly dominant

species. This floristically and geologically diverse site has

not previously heen sampled for terrestrial snails and is

likely to sustain additional undiscovered species.

MATERIALS AND TEXT CONVENTIONS

Specimens were hand-collected or sifted from samples

of leaf-litter. Live-collected animals were drowned and

then presented in 75% ethanol. Gross anatomical dis-

sections were made under 75% ethanol using a dissect-

ing microscope. Radulae were isolated from dissected

buccal masses using a 5% sodium hypochlorite solu-

tion. Scanning electron micrographs of radnlae were

made using a Eield Emission-SEM. Measurements

were taken using an ocular micrometer. Whorl count

was measured from the suture of the first whorl to the

body whorl and fractions of a whorl were determined

with the aid of a cardboard circle divided into ten etpial

parts of 36° (Eigure 2, line 1-2.9). Spire width was the

length of a straight line passing from the apertural edge

of the sutnre through the middle of the apex to the

opposite suture (Figure 2, hue A-B). Shell width was

the greatest width of the shell perpendicular to the shell

axis (Figure 3, line C-D). Shell height was the greatest

distance between the apex and the base of the aperture

measured parallel to the shell axis (Figure 3, line D-E).

Spire height was measured from the top of the body

whorl to the apex of the shell (Figure 3, line F-G).

Aperture width was the greatest distance from the colu-

mellar edge to the outer edge of the aperture (Figure 3,

line E-H). Aperture height was measured from the

suture to the base of the aperture, parallel to the shell

axis (Figure 3, line H-I). Shell measurements are based

on nine unbroken adults; ranges are followed by mean

and standard deviation. The lengths of radular teeth

}. Slapcinsky, 2009

Page 55

were measured from the top ol the mesocone to the

posterior edge of the basal plate. The widths of

radular teeth were measured as the greatest width of

the cusps, not the basal plate. The following abbrevia-

tions are used in figures of genital anatomy: AT = atri-

um, BC = bursa copulatrix, BT = bursa tract, EP =

epiphallus, PE = penis, PP = penial pilaster, PB =

penial retractor muscle, SO = spermoviduct, V = verge,

VA = vagina, and VD = vas deferens. All specimens are

deposited in the Florida Museum of Natural Histoiy,

Gainesville (UP).

SYSTEMATICS

Family Charopidae Hutton, 1S84

Genus Panjphantopsis Thiele, 1928

Type species: Flammulina {Panjphantopsis) lamcUi-

gera Thiele, 1928, by original designation.

Panjphantopsis bradleiji new species

(Figures 4-10)

Holotyije: UF 378116 (diy shell), Papua New Guinea,

Simbu Province, 78 km SSW ol Goroka, 11 km E ol

Haia Village, Grater Mountain Biological Research Sta-

tion, approximately 6.72° S, 145.09° E., 1100 m altitude,

D. Wright, 6 Apr. 1992.

Paratyjies: Type locality: UF 274062 (1 alcohol-pre-

served), UF 378115 (1 diy shell), 1100 m, D. Wright,

7 August 1991; UF 274059 (2 alcohol-preseiwed), UF

420747 (2 diy shells), 1350 m, D. Wright, 21 Apill

1992; UF 274061 (2 alcohol-preseived), 1100 m,

D. Wright, 6 April 1992; UF 274057 (1 alcohol pre-

served), 1130 m, D. Wright, 18 March 1992; UF

179660 (1 alcohol presented), 1160 m, D. Wright, 1 July

1990; UF 274060 (1 alcohol-preseiwed), A. Mack, 25

Januaiy 1993; UF 274058 (1 alcohol preserved),

UF378114 (1 chy shell), A. Mack; UF 274056 (1 alco-

hol-preseiwed), A. Mack; UF 179657 (1 juvenile, alco-

hol-preseiwed), 1130 m., D. Wright.

Description: Adult shell depi'essed; large for genus,

9.1-11.3 mm (10.3±0.8) in width and 4. 1-5.9 mm

(5.2±0.6) in height, with 2.8-3. 1 (2.9±0.1) rapidly

e.xpanding whorls (Figures 4-6). Suture impressed and

broadly channeled. Apical surface of whorls flattened

bePA^een suture and periphery. Shell periphery angular

to carinate above mid-point and rounded below, flatten-

ing abruptly basally. Spire flat or only slightly elevated,

0.0-0. 3 mm (0.1±0.1) and narrow 3.4-4. 1 mm (3.7T0.2)

only 0.3-0. 4 (0.36T0.02) of shell width. Teleoconch

whorls do not descend or descend only slightly, shell

heigh t/diameter ratio 0.4-0. 6 (0.50±0.04). Approximate-

ly 1.7 flattened protoconch whorls sculptured with about

12-17 rows of spiral pits that continue on teleoconch

becoming elongate and less regular. These pits are ob-

scured by thick periostracum on teleoconch but can be

oliserved in aperture througli translucent shell. Teleo-

conch whorls have growth lines accentuated with short

periostracal e.xtensions. Approximately ever)' fifth exten-

sion longer and bearing a triangular process (appro.xi-

mately f.0-1.5 mm long) at shell margin. These

processes are retained to maturation. Protoconch whit-

ish to tan. Teleoconch whorls brown; first 2.5 whorls

darkest below suture and last 0.5 whorl dark throughout.

A reflection of peristome completely covers umbilicus

at all stages of growth. Aperture is ovate to almost quad-

rate and very large \\4th an aperture-height to aperture-

width ratio of 0.4-0. 7 mm (0.63±0.10).

Epiphallus 3x diameter of vas deferens, only slightly

inflated apically, widening gradually basally, folded ap-

proximately at mid-point, and does not bear an apical

diverticnlnm (Figure 7). Penial retractor muscle short

and robust, originating from diapliragm and inserting

on epiphallus half way between epiphallar fold and junc-

tion with penis. Penis ovate and slightly inflated basally;

epiphallus joins it laterally just below rounded apex.

Penis width is 0.5 x length and 2-3 x width of epiphal-

lus. Penis wall thin and smooth and interior of retracted

penis containing large verge that, when unfolded, is

bowl-shaped (Figure 8). Interior of verge is sculptured

with tongue-shaped pilaster that extends from epiphallar

opening to near base of penis. Atrium short and broad,

with nearly same diameter as penis. Vagina relatively

long, \x4th aliout same length and only sliglitly narrower

than penis. Free oviduct short, and with nearly same

width as and poorly differentiated from vagina. Base of

bursa tract narrow, its diameter only 0.3 x diameter of

free oviduct where they meet, rapidly narrowing to 0.2x

diameter, and remaining narrow to junction with bursa

copulatrfx.

Gentral and first lateral teeth of radula are tricuspid,

9-10 pm wide, and 11-12 pm long (Figure 9). Meso-

cones of central teeth and lateral teeth tall and slender,

projecting slightly beyond their basal plates, and origi-

nating from center of their basal plates rather than from

a ridge on posterior edge of basal plate as in most other

Parijjyhantopsis species, Ectocones of central teeth

trigonal and symmetric. Ectocones and endocones of

lateral teeth are trigonal and about 0.5 x height of meso-

cones. Endocones and ectocones of lateral teeth nearly

symmetrical, endocones veiy slightly larger but other-

wise of similar shape to their ectocones. First 15 teeth

to left and right of central row are similar to first lateral

teeth, next tw'o teeth on either side grade in shape and

are difficult to classify as either lateral or marginal teetli.

Last seven teeth clearly marginal and dorsovejitrally

compressed, 11-12 pm wide and 8-9 pm long (Fig-

ure 10). Endocones of marginals nnicuspid and about

0.7x as tall as mesocones. Ectocones are nnicuspid to

irregularly multicuspid and much shorter, about 0.5 x as

tall as mesocones.

Habitat: All specimens were collected f rom sub-mon-

tane forest (Paijnians, 1976) between 1100 and 1.350 m

elevation in leaf litter and on live leaves especially of

Zingiberaceae within 1 m of the ground. Vegetation

Page 56

THE NAUTILUS, Vol. 123, No. 2

Figures 4-10. Parijj)liaut(^>sis hradlei/i new species. 4-6. Photographs of shell, holotype UF 378116, diameter 11.3 mm. Scale

har = ]() mm. 7. Photograph of genitalia, UF 274062. Scale bar =

electron micrograph of rachila, UF 274062. Scale bars = ft) m.

at the ty]K^ locality coiisi.sted ol mature micnt forest with

a tew small patclies ol late secouclaiy growth Irom abau-

rloiied gardens. Mean animal rainfall is 6400 mm and is

evenly distributed throngliont tlie year. Diurnal temper-

1 mm. 8. Drawing of penis interior, UP’ 274062. 9-10. Scanning

atnres are 15-28° C. The area’s soils range from dark

lirown loam to orange clay with variable soil nutrients

including soil calcium that ranges from 270 to 1560 ppm

(Wright et ah, 1997).

J, Slapcinsky, 2()09

Page 57

Etymology: This patronym honors botanist Ted

Bradley, Santo Domingo de Heredia, Costa Rica (retired

from George Mason University, Fairfax, Virginia) my

friend, teaclier, and field companion v\dio encouraged

my interest in taxonomy and introduced me to the rich

and underreported diversity ol the tropics.

Remarks: Pan/phantopsis bracllet/i new species is one

of the largest species in the genus and is similar in size

only to P. louisiadarum (Mollendorff, f899) and P. glohosa

(Hedley, 1890), both of whicii differ in having shells vvitli

rounded margins that lack periostracal processes. Parij-

phantopsis hracUei/i appears similar to species that have

shells with angulate to carinate mai'gins and that bear

periostracal processes that are rettxined in adults: P. corolla

Slapcinsky and Lasley, 2007, P. elegaiis (Fulton, 1902),

P. fnltoni (Coen, 1922), P. lamelligera (Thiele, 1928),

P. lebasii Slapcinsk)/, 2005, P. t/awii Slapcinsky, 2005,

and P. ijdensis Slapcinsky, 2006. The peiiostracal process-

es of P elegans, P. fnltoni, and P. i/awii overlap, forming a

continuous serrated edge at the shell margin, while

the peiiostracal processes of P. biYKllei/i, P. lamelligera,

and P. i/elensis each taper to a point, unlike the rounded

processes ofP lebasii. Of the species for which the genital

anatomy is known, P. bracllei/i is similar to P. lamelligera,

P. lebasii, and P. i/awii in lacking an apical diverticulum

on the epiphallus, unlike P. corolla and P. i/elensis. The

mesocones of both the central and lateral teeth of tlie

radula of P. bracllei/i join the basal plate near its center as

in P. lebasii. P. yawii, and P. yelensis, and unlike P. corolla.

DISCUSSION

Unlike that of many other taxa, species diversity of ter-

restrial snails has been considered to be low in tropical

rainforests (Solem, 1984). However, recent suiweys have

demonstrated that terrestrial snails are often diverse in

tropical rainforests (Winter and Gittenberger, 1998;

Schilthuizen and Rutjes, 2001) and it appears that low

abundance and sampling intensity are the reasons for

perceived low diversity of snails in rainforests. Lack of

sampling is particularly troubling because rapid defores-

tation is leading to the extinction of narrowly endemic

snail species in many tropical forests (Emberton, 1995;

Emberton et ah, 1997). Unfortunately, non-marine mol-

lusks appear to be particularly prone to e.xtinction, con-

stituting an alarming 42% of the recorded extinctions of

animal species since the year 1500 (Lydeard et ah,

2004). Much more of this loss may go unreported be-

cause terrestrial snails receive relatively little taxonomic

study in relation to their diversity. Indeed, there are

appro.ximately 24,000 described species and an esti-

mated 11,000 to 40,000 undescribed species (Lydeard

et ah, 2004). The land snail fauna of New Guinea

appears to be especially diverse but few of its mountain

ranges have ever been suiweyed for most invertebrate

groups. These mountains support diverse and highly

endemic snail faunas that are only now? being discovered

(Slapcinsky, 2005). New Guinea sustains the largest

tracts of tropical broadleaf forest remainitig in Australa-

sia and the third largest on the planet after tlie Amazon

and Gongo forests (Brooks et ah, 2006). More than 71%

of Papua New Guinea is forested and 57% of this forest

is commercially valuable and globally imperiled lowland

rainforest (Shearman et ah, 2008). These resources will

come under increasing commercial and developmental

pressures as other forests in the region are exhausted.

Already, the rate of deforestation in Papua New Guinea

is higher than previously believed, and is accelerating,

and if these rates continue it is estimated tliat 83% of the

conntiy’s forests will be cleared or degraded by 2021

(Shearman et ah, 2008). The loss of these forests will

result in the extinction of endemic species dependant

on forest habitat, many before they are ever discovered.

Future efforts to preseiwe rapidly dwindling forests will

depend on documentation of their rich biota.

ACKNOWLEDGEMENTS

I thank A. Mack and D. Wriglit for collecting the

specimens and for bringing them to the attention of

F. G. Thompson who arranged funding for fui'ther

collecting through the University of Florida Foundation,

McGinty Endowment; K. Kelley, Electi'on Microscopy

Core Laboratoiy, University of Florida for imaging the

radulae; R. Lasley for photographing the shells, and

G. Barker and R. Cowie for suggesting improvements to

the manuscript.

LITERATURE CITED

Brooks, T.M., R.A. Alittenneier, G.A.B. da Fonseca, J. Gerlacli,

M. Hoffinann, J.F Lamoreu.x, C.G. Mittenneier, J.D.

Pilgrim, and A.S.L. Rodrigues. 2006. Global biodiversity

conseivation priorities. Science 3L3: 58-61.

Emberton, K.C. 1995. On the endangered biodiversity? of

Aladagascan land snails. In: van Brnggen, A.G., S.M.

Wells, and T.C.M. Kemperman (Eds.) Biodiversity and

Conseivation of the Mollusca. Backhnys.Oegstgeest-Lei-

den, pp. 69-89.

Emberton, K.C., T.A. Pearce, P. F. Kasigwa, P. Tattersfield, and

Hahibn Z. 1997. Higli diversits? and regional endemism in

land snails of eastern Tanzania. Biodiversity and Conser-

vation 6: 1123-1136.

Lydeard C., A. Bogan, P. Bouchet, S.A. Clark, R.Il. Cowie,

K.S. Cummings, T.J. Frest, D.C. Herbert, R. llersliler,

K.E. Perez, VV. F. Ponder, B. Roth, M. Seddon, E.E.

Strong, and F.G. Thompson. 2004. The global decline of

nonmarine mollnsks. BioScience 54: 321-330.

Mackenzie D.E. and R.W. Johnson. 1984. Pleistocene volca-

noes of the western Papua New? Guinea Highlands: mor-

phology, geology, petrography, and modal and chemical

analyses. Australian Bureau of Mineral Resources Geolo-

g)? and Geophysics Report 246: 1-271.

Paijmans, K. 1976. New Guinea Vegetation. Australian Nation-

al University Press, Canberra, 212 pp.

Schilthnizen M. and H.A. Rutjes. 2001. Land snail diversity in

a square kilometre of tropical rainforest in Sabah, Malay-

sian Borneo. Journal of Molluscan Studies 67: 417-423.

Page 58

THE NAUTILUS, Vol, 123, No. 2

Sliearman, P. L., J.E. Bi-yan, J. Ash, P. Hunnam, B. Mackey,

and B. Pokes. 2008. The state of the forests of Papua New

Guinea. Mapping tlie e.xtent ami condition of forest cover

and measuring the drivers of forest change in the period

1972-2002. University of Papua New Guinea, Port Mor-

esby, 14S pp.

Slapcinsky, J. 2005. Si.x new species oi Parijphantopsis (Gastro-

poda: Pidinonata; Charopidae) from the Papuan Peninsu-

la of New Guinea. The Nautilus 119: 27-42.

Slapcinsky, |. 2006. Panfphantopsis (Gastropoda: Pulmonata:

Charopidae) from the Louisiade Archipelago of New

Guinea. The Nautilus 120: 119-130.

Slapcinsky, and R. Lasley. 2007. Three new species of Par-

ijphantopsis (Gastropoda: Puhnonata: Charopidae) trom

the Nakanai Mountains, New Britain, Papua New Guinea.

The Nautilus 121: 182-190.

Solein, A. 1970. The endodontid land snail genera Pilshn/char-

opa and Panjphautopsis (Mollusea: Puhnonata). The Veli-

ger 12: 239-264.

Solem, A. 1983. Endodontoid land snails from Pacific Islands

(Mollusea: Puhnonata: Signmrethra). Part II. Families

Punctidae and Charopidae, Zoogeography, Field Museum

of Natural History, Chicago, 336 pp.

Solem, A. 1984. A world model of land snail diversity, hi:

Solem, A. and Van Bruggen, A.C. (Eds.), World-wide

snails: biogeographical studies on non-marine Mollusea.

E.J. BrillA'V. Backhuys, Leiden, pp. 6-22.

Van Benthem Jtitting W. S.S. 1964. Non-marine Mollusea of

West New Guinea. Part 3, Pulmonata, I. Nova Guinea,

Zoology 26: 1-74.

Winter, A. J. and E. Gittenberger. 1998. The land snail fauna of

a square kilometer patch of rainforest in southwestern

Cameroon: high species richness, low abundance and sea-

sonal fluctuations. Malacologia 40: 231-250.

Wright, D.D., I.H. Jessen, P. Burke, and II. Gomez de Silva

Garza. 1997. Tree and liana enumeration and diversity on

a one hectare plot in Papua New Guinea. Biotropica 29:

250-260.

THE NAUTILUS 123(2):59-70, 2009

Page 59

fiian Jose Parodiz (1911-2007): obituaiy and bibliography

Charles F. Slurin^

Section ol Mollusks

Carnegie Mtrsenni of Natural Ilistoiy

4400 Forbes Avenue

Pittsburgh, PA 15213-4080 USA

csturm [email protected]

INTRODUCTION

Even' now and then, we meet an individual who po.s.ses.s

knowledge not only in depth in a given subject, but also

across several fields of knowledge. Often, we are not

aware of these many interests of this Renaissance jrer-

son. I knew Juan Jose Parodiz as a malacologist and

curator emeritus at the Carnegie Museum of Natural

Ilistoiy. It was after he died that I learned much more

about him and his many accomplishments.

|ose, as he preferred to he called, passed away on

4 September 2007 at the age of 95 years. He was horn

in Buenos Aires, Argentina, on 21 December 1911. He

was one of the last of the classically trained malacolo-

gists, brought up in an era before the widespread use of

computers, molecular biology, and cladistic analyses.

THE EARLY YEARS

Jose was the son of Alercedes Gonzalez Parodiz. Jose

never knew his latlier and his mother died when he was

5 years of age. He and his younger sister Avelina then

went to live with an aunt, Avelina Andrea Parodiz, whom

they regarded as their mother. She was employed as a

postmaster in Buenos Aires. This may explain Jose’s later

interests as a philatelist. Jose had three other aunts, and

one of them, Juana Pabla Parotliz, made him promise tliat

he would look after them until they tiled. This promise,

which he honored, was to play a central role in his life.

Jose went to work at tlie Aluseo Argentino de Ciencias

Naturales “B. Rivackma” (MACN) in 1927. He was

16 years old. Jose mentioned that his "mother” wanted

him to be a lawyer, however, this profession held no

interest for him. On the other haiid, what sparked his

interest in natural histoiy is unknown.

MUSEO ARGENTINO DE CIENCIAS NATURALES

(1927-1952)

At the Aluseo Argentino de Ciencias Naturales, Jose went

to work in the section of invertebrate biologvc He work'd

' Research Associate

under two men tliat he held in great esteem. Dr. Martin

Doello Jurado and Alberto Carcelles. Doello Jnrado was

the director of the museum. Carcelles, like Jose, came to

the museum witli no formal training. Jose was trained by

Doello Jurado and Carcelles. Carcelles went on to be-

come an eminent malacologist woi king with marine mol-

Insks. Jose’s first paper was co-anthored with him.

Parodiz was invoked in oceanographic expeditions in

the South Atlantic and the Southern Ocean. These expe-

ditions occurred in 193S and 1939. He spent time on the

ships ARA COMODOKO Ri\adavia and ARA B.uua Blanca

(Patagonia, Tierra del Fuego, Isla de los Estados, and

the Magallanes Strait). On these expeditions, not only

mollusks, but other invertebrates (and fish) were collect-

ed. The collections included dredged material as well as

shore-collected specimens.

After Jose started liis work in Recent invertebrates, he

was eventually appointed the head of im'ertebrate pale-

ontology (1940-1952) at AIACN. Throughout liis career,

he would continue to study both fossil and Recent mol-

hisks of SoTith and North America.

While in Argentina, Jose was a member ol the Asocia-

cion Ai'gentina de Ciencias Naturales. He seived as

secretaiy from 1945-1950 aud resigned f rom the associ-

ation in 1952. Though Jose was mainly involved with

mollusks, he had au interest in invertebrates in general.

There are unpublished manuscripts of talks that he gave

over the radio. Two of these talks were titled "Sponges”

and “Crustaceans of economic value ”. These talks were

broadcast from 1942 to 1944.

Jose also woi'ked at the Estacion Hidrobiologica de

Puerto Que(|uen (a part of AIACN). This was a research

facilit)' that was started by Doello Jurado. Jose was also

an assistant in Geolog)' and Paleontolog)' at the Univer-

sity of Buenos Aires (1930-1933). With whom he

worked at the Universitx' is unknown. Prom 1935

tlirougli 1945, Jose collaborated with Egitlio PYruglio, a

geologist, on identifying fossil mollusks.

Upon the retirement of Doello Jurado, the AIAGN

was directed by Agustin Eduardo Riggi- Jose and Riggi

did not get along. Jose felt that Riggi was politically

motivated and did not possess the same (pialifications

that Doello Jnrado brought to tlie director's position.

Page 60

THE NAUTILUS, Vol. 123, No. 2

There was a good deal of animosity in their relationship

that would come to play in decisions that |ose made iii

the Intnre.

While fose worked at the MACN, he met many emi-

nent scientists, especially from the United States.

Among the scientists were Fritz Haas, Heniy A. Pilsbiy,

and \Valdo L. Schmitt. These indiUdnals encouraged

)ose to visit and study in the United States. Jose planned

a \asit to the United States with Schmitt’s assistance;

howevei', shortly before he was to make the trip, he

cancelled it dne to a political coup that was unfolding in

Argentina.

American mnsenms hold many of the type specimens

of South American mollusks and Jose wanted to study

tliem. In 1949, Jose applied for a fellowship from the

)ohn Simon Guggenheim Memorial Foundation. He was

awarded a fellowship and in 1950 he spent six months in

the United States. Jose conducted the majority of his

research at the National Alusenm of Natural Histoiy

(Smithsonian Institution), and the remainder at the

Academy of Natural Sciences (Philadelphia) and the

Mnsenm of Comparative Zoology (Haiward University).

Prior to leaving for the United States to conduct his

studies, lose recpiested that he he kept on the payroll at

the AIACN. Riggi refused. Jose saved his vacation time

Irom 1949-f95() so that he would have some paid time

while in the Unites States. Jose was scheduled to spend a

year in the United States. The staff at the Guggenheim

Foundation allowed Jose to shorten his stay to 6 months

due to his financial situation.

W'hile in Washington, DC, )ose lived in housing

arranged for by Waldo Schmitt. The landlady had a

friend, Esther Elizabeth Sell, who worked as a secretaiy

in the Treasnr)' Department. She introduced Esther to

Jose, and Esther and Jose became romantically involved.

Prior to returning to Argentina, Jose promised Esther

that he would return and that they would wed. Needless

to say, Esther never thought that she would see him

again.

During Jose's stay in the United States, he attended

the ammal meeting of the American Malacological

Union (AMU; now American Malacological Society',

AVIS), which was held in Chicago. Years later, Jose was

to seiwe as pi'esident of this group. fOuring his visit to the

Midwest, Jose collected PJnionidae from the Meramec

River near St. Louis. His notes indicate that he also

\4sited the "Chicago Afusenm” (probably the Field Mu-

seum of Natural Histoiy) and the UniversiC of Michigan

Mnsenm of Zoology. At the AMU meeting, he met many

of the most iniluential malacologists in the United

States. From Jose’s work during his fellowship, two

papers were written (Parodiz 1950, 1962b).

Jose and Riggi bail a contentions relationship liefore

Jose went to the United States, which only worsened

njion Jose’s return to Argentina. This increased animosi-

ty was fueled iu part by laudatoiy letters sent to Riggi by

Schmitt, Clench, and Haas, people who Jose met or

studied under in the United States. These letters de-

scribed the high (juality of Jose’s scholarship. Jose’s suc-

cess in the United States put him under closer scrutiny

by Riggi.

Upon returning to Argentina, Jose was informed that

his last sinwiving aunt, Juana Pabla Parodiz, had passed

away. She passed away the day Jose left New York City

to return to Buenos Aires. He was no longer bound to

his promise to help care for her. This release, along with

a deteriorating relationship with his superior, and a

woman that he loved in the United States, set in motion

his ne.xt plan, one of permanently immigrating to the

United States.

At this point I would like to address Joses education.

Jose was often addressed as “Doctor Parodiz”; however he

appears to have had no formal academic degree. He

joined the MACN at the age of f6 and was trained by

Doello Jurado and Carcelles. He assisted someone at the

University of Buenos Aires, yet there are no documents of

what he did there, though he was fond of recounting his

stories w4th the student theater ensemble. In a 1948 issue

of Comnnicaciones del Museo Argentine de Ciencias Nat-

urales Serie Ciencias Zoologicas, the journal of MACN,

the members of the staff are listed with their titles: “Dr.”,

“Prof”, and “Lie.”. Joses name is preceded by none of

these designations. On his application for the Guggen-

heim fellowship, the section of the application which asks

for the applicant’s educational background was left nn-

completed. Lastly, queries to staff at MACN resulted in

no information regarding any formal training that Jose

may have obtained (pers. comm., M. G. Quintana, 2007).

We may draw then the conclusion that Jose had no aca-

demic degree and that his title as a doctor was a well-

deserved honor bestowed upon him by his colleagues in

recognition of his significant accomplishments.

CARNEGIE MUSEUM OF NATURAL HISTORY

(1951-2007)

Once Jose decided to move to the United States, he

needed to find a job. He wrote to contacts in the United

States and Schmitt informed him of a curatorial position

that was open at the Carnegie Mnsenm of Natural His-

toiy (CMNH) in Pittsburgh. This position had been va-

cant for a year. Jose applied for the position and was

hired. He began his duties at the CMNH in 1952. Some-

what jokingly, he once said that no one else wanted the

job due to the low salaiy.

Jose informed few colleagues in Argentina of his plans

to move the United States; he was fearful of retribution

by Riggi. Jose saved his vacation time, packed, and left

for the United States. Upon learning of Jose’s departure,

Riggi attempted to have him fired to create a black mark

on his record. Senior staff of MACN would not allow

such au action.

Prior to settling in Pittsburgh, Jose returned to

Washington. On April 26, 1952, he wed Esther Sell.

Waldo Schmitt was Jose’s best man. Jose then moved to

Pittsburgh wliere he and Esther resided together until

her death.

C. F. Sturm, 2009

Fuffe fil

Figures 1-4. Juan Jose Parodiz. 1. Ahoard the Argentinean Nav)' research vessel the AHA Haiii v Blanc:a in 1939. 2. At CMNH in

1961. .3. With wife Estlier at the AMU meeting in 1973 (Newark, Diiaware). 4. As Curator Enieritns in 1997, working in the inollusk

collection at CAINIl. Figures 1, 2, and 4 are Iroin the arcliives of CMNIl, figure 3 was Ironi the collection ol Hohcrt Hohertson,

Curator Emeritus, Academy of Natural Sciences, Philadelphia.

THE NAUTILUS, Vol. 123, No. 2

Page 62

Jose was a curator at the Carnegie Museiun ol Natural

Ilistoiw from 1952 to 1980. He retired in 1981 and was

active as a curator emeritus until his death in 2007.

Upon arriving at CMNH, Jose developed a 10-year plan.

Of this plan, he succeeded at some parts and never

completed others.

One of Jose’s plans was to reorganize the collection.

This project e.xtended well past the first ten years and

was never fully realized. While the Unionoida and Gas-

tropoda were reasonably well organized, the reorganiza-

tion of the marine bivalves was not completed at the

time of his retirement. The Sphaeiiidae, mcjst of which

were transferred to the CMNH upon Victor Sterki’s

death (1933), were virtually untouched by Jose. Jose

began a project of segregating t\pe material from the

general collection. He placed the t\pe specimens in a

separate cabinet. The majorit)’ of the t\pe specimens

that he overlooked were the t\pes of Sphaeriidae in

Sterki’s collection. Only si.\ lots of an estimated 900 lots

were transferred to the tvpe cabinet.

Another aim of Jose’s 10-year plan was to maintain

continued collaboration with malacologists in South

America. This part of his plan succeeded remarkably

well. He not only continued to undertake lieldwork in

Sontli America, but South American scientists came to

the Carnegie to study its collection and to collaborate

\Cth Jose. This will be discussed in more detail later in

this paper.

Jose sought to expand the mollnscan collection at

CMNH. He especially wanted to expand the collection

of South American mcrllnsks. The Carnegie already had

a significant collection of South Amei'ican nnionoids col-

lected by John Haseman in the 1920s. Jose embarked

upon an exchange program with other innsenms, thus

expanding the number of holdings at CMNH.

Prior curators at CMNH studied and published on

various groups of Pennsylvania mollnsks. Arnold Ort-

mann published on the nnionids, Victor Sterki on the

Sphaeriidae, and Stanley Brooks on the terrestrial Gas-

tropoda. A comprehensive study on the (reshwater gas-

tropods was lacking. Jose planned to undertake such a

study. Periodically, Jose worked on this study from when

he proposed his 10-year plan until his retirement. He

did not complete it. In 1956, he published a paper on

one aspect of freshwater gastropods in Pennsylvania

(Parodiz 1956d). In 1958, Jose published a list of the

freshwater gastropods of Pennsylvania (Parodiz 1958c).

In this paper, Jose indicated that this list was a prodi'ome

of a more e.xtensive manuscript, soon to be submitted for

publication. On this point, Jose erred.

In the late 1970s, while collaborating on a study with

C.R. Bristow, Jose mentioned that the director of

CMNH was >irging him to finisli the study of the fresh-

water gastropods of Pennsylvania. In a letter to William

J. Clencli, he mentioned that he was waiting for Clench’s

nionograjrh on the Plenroceridae and posed several

({iiestions to Clench. Clench’s monogra})h was pub-

lished; still, Jose’s mannscripl remained just a manu-

script. His 150-page mannscrijV on the freshwater

gastropods remained unpublished. For whatever rea-

sons, Jose invested significant time on this project.

He conducted field studies, reviewed collections in

museums, and corresponded with others. His work still

remains in the manuscript stage and the Pennsylvania

freshwater gastropod fauna still await a comprehensive

study.

A few years after Jose arrived at CMNH, Waldo

Schmitt offered him a position at the National Museum

of Natural Histoiy Jose declined the offer despite its

larger salaiy and a larger collection at the National Mu-

seum of Natural Histoiy. There was one significant rea-

son for Jose turning down the offer. The National

Museum of Natural Histor)' was part of the Smithsonian

Institution, an institution run by the United States gov-

ernment. Though Jose and Schmitt shared a close

friendship, Jose did not want to entertain the possibility

of finding himself, in the future, of answering to an

eventual political appointee. He did not wash to be in

the same position that he was in Argentina when Doello

Jnrado retired and wais replaced by Riggi. Jose, who

liked the director at CMNH, chose to pass up the bene-

fits of a new position at the Smithsonian and remained

in Pittsburgh, at a private institution.

In 1954, the journal Neotropica was launched. Jose

wais one of its founders. Jose remained on the editorial

board from 1954 through 1972. In addition, Jose seiwed

as the North American contact for the journal. Jose wais

also active in the journal Malacologia. Initially, Jose w'as

responsible only for translating abstracts into Spanish.

Shortly thereafter, he was asked to join the editorial

board. He seiwed in this capacity from 1962-2006. Yet a

third time, Jose wais asked to be on the editorial board of

a journal. This time, John Burch (in litteris, 1990)

requested that Jose join the editorial board of Malaco-

logical Review. During his career, Jose also seived as a

reVew^er lor several other journals including The Nauti-

lus and the Journal of Moiqohology.

Jose w'as responsible for the mollnsk display at

CMNH. This w^as developed in the early 1960s under

the sponsorship ol the Commonw'ealth of Pennsylvania.

The e.xlribit, w'hich is still extant, comprises six display

cabinets, each approximately 0.9 x 1.2 m. The exhibit is

titled “Sea Shells by the Seashore”. The display cabinets

are titled (1) How Shells are Named, (2) Shells and Such

(Classes of Mollnsks), (3) Eating and Aloving, (4) How'

Mollnsks Keprodnce, (5) Scavenging in New England,

and (6) Beach Combing in Florida. This exhibit was the

subject of an article in the Carnegie Magazine (Parodiz

1962e).

Jose received only one governmental grant during his

career. It w'as a National Science Foundation grant,

awarded for the years 1961-1962, for the study of non-

marine mollnsks of Argentina, Uruguay, and surround-

ing territories. Jose conducted Held wx)rk (w4th Alberto

Carcelles and Argentino A. Bonetto) tinring those Lvo

yeais, purchased the etjuipment that he needed, and

concluded his w'ork with 10% of his funding still intact.

Jose refused to use tlie monies for other purposes. He

C. F. Sturm, 2009

Page 63

felt that he had completed the work and any remaining

hinds should not be used tor other pinposes but

returned to the granting agency. Jose was hassled by a

few colleagues for tliis action, so much so that he chose

never again to apply for another grant. Among the

papers that resulted from this grant are Parodiz (1963c,

1965b, 1966a), Parodiz and Bonetto (1963), and Parodiz

and Hennings (1965).

Jose was parsimonious. He would combine visits to

Iriends and family in Argentina with his field work.

Since he knew about the political and economical tluc-

tuations in South America, he was able to fund his work

in South America from his museum budget. Even the

director who hired Jose, Graham Netting, realized Jose's

thriltiness. In a memo. Netting commented about this

by writing “...since you rarely succeed in spending your

entire Section budget...” (in litteris, 1969).

In 1962, Dean Putnam Jones of the University of

Pittsburgh proposed to Chancellor Litchfield that Jose

should be appointed as an adjunct member of the Grad-

uate Faculty at the University of Pittsburgh. In a letter

dated 2 January 1963, Litchfield informed Jose of his

appointment as an adjunct member of the Graduate

Faculty. Jose never acted in a professional capacity at

the University. He felt that his responsibilities as curator

were such as to preclude his serving in a professorial

role at the University. As he put it in a memo to Craig

Black (director of CMNH, 1975-1982), “I was con-

vinced that I could not fulfill both efficiently, at the same

time.”

Jose, along with Gladys McCallum, founded the Pitts-

burgh Shell Club in 1965. The first meeting was on

27 March 1965, with 13 people in attendance. Jose was

appointed councilor (advisor) of the club. The clul) pub-

lished the Pittsburgh Shell Club Bulletin (1966-1979).

Jose was a frequent contributor to the publication. In

1975, Jose was awarded the DuPont Trophy for his out-

standing exhibit at the Pittsburgh Shell Club Show. The

exhibit dealt with the evidence for evolution as seen in

fossil and Recent mollusks. In 1977, Jose w'as elected an

honoraiy member of the Pittsburgh Shell Club. The

only other person so honored was William J. Clench of

H award University.

While Jose spent most of his field time in South

America, in 1976 he had planned a trip to Guatemala.

Shortly before he was to leave, an earthquake there

forced him to cancel his trip. Instead, he w^ent to the

Yucatan Peninsula (Mexico) and stayed with a friend,

Dorothy Zapata. Her husband drove Jose around during

his stay. For three weeks, Jose collected marine mol-

lusks. In all, he visited ten stations. He published his

catalog for this trip in the Pittsburgh Shell Club Bulletin

(Parodiz 1979e). This paper includes a number of range

e.xtensions for mollusks found in the Caribbean biogeo-

graphic region.

In addition to pursuing his own research interests in

Soutli American malacology, Jose sought to help others.

One such example involved Kenneth Boss who was

studying the neotropical fossil Ampullariidae. In 1975,

he started corresponding with Jose. Several letters later.

Boss offered Jose co-authorship on a paper on which

he was working. The paper was published iu 1977

(Boss and Parodiz, 1977). Another example involved

C. R. Bristow. Bristow w^as a British geologist working

in South America. Bristow begaii his correspondence

with Jose in 1972. Six years later, he suggested that they

jointly publish a paper on the Tertiaiy non-marine mol-

lusks of Ecuador. The paper was published in 1982 with

Bristow writing the stratigraphic portion and Jose tlie

taxonomic portion (Bristow and Parodiz 1982).

Jose hosted many South American inalacologists at

CMNH. Among them were Miguel Klappenbach

(1963), A. Carcelles (several times, his longest visit being

8 months in 1965-66), Maria C. Dreher Mansur (1998),

and A. Bonetto (1959, 1963). In addition, Jose aided

researchers from all over the world witli liis knowledge

of South American malacology, paleontology, and poli-

tics. Numerous students and researchers sought Ins

opinion of their proposed studies in South America. Jose

would advise them of localities that they should visit,

museums where they would find specimens for study,

and whom to contact while in South America. If it was

appropriate, he would comment on the political situa-

tion that vv'aited for them. Jose spoke with the authoritv'

of one who had an intimate w^orking knowledge of South

American museum collections, knowledge built ov-'er the

decades from studying these collections personally. In an

era before the Internet, before one could easily search

for sucli information, Jose seived this purpose.

Jose also aided others in their work with tlie Nortli

American fauna. Some examples include identiljlng

(1) the gastropods in the stomach contents of box turtles

(1955), (2) the freshwater gastropods from the Cheat

River in West Virginia (1962), and (3) the freshwater

gastropods from the Susquehanna River in Peunsylv'ania.

Also, he provided comments to the authors of a new

unionoid from the Mesozoic of Uruguay (1993).

An intriguing aspect of Jose's w'ork were the forensic

studies in which he was inv'olved from 1953-1963. One

such case involved a worm found in a can of chicken

soup. Jose identified it as a blood vessel from a chicken.

He did not specify whether it was an arteiy or a vein.

Another case inv'olved a can of food that had been

imported and contained maggots. He determined tliat

the maggots were not from the countiy where the prod-

uct originated and tlnis absolved the company of any

wrongdoing.

Jose maintained membership in a number of profes-

sional societies. These include the Malacological Society

of London, Unitas Malacologica, the Paleontological

Research Institution, Brazilian Society of Malacology

(honoraiy member), the Malacological Society of Uru-

guay (honoraiy member), and as mentioned earlier, the

Pittsburgh Shell Club and the Argentine Association of

Natural Science. Jose was also a member of the Ameri-

can Malacological Union (later the American Malacolog-

ical Society). He seized as president of this organization

from 1964-1965.

Page 64

THE NAUTILUS, Vol, 123, No. 2

Jose was curator emeritus at CMNH from 1981-2007.

He eoutiuued to publish during this period. His last

publication dealt with a South American uuiouid (Paro-

diz and Morton, 2002).

lose was honored on three occasions by fellow scien-

tists. In 1998 he was awarded a “Diploma of Honor" for

his life’s work. This award was bestowed on him fry the

Brazilian Society of Malacology. In 1992, Balech and

lose published a hi.stoiw of the MACN. In 2001, along

with his friend Balech, he was honored as an “Illustrious

Researcher” of the MACN. Lastly and posthumously in

2008, Jose was acknowledged as “an esteemed colleague,

a distinguished malacologist, and a warm-hearted

friend ' by the American Malacologieal SocieW.

OTHER INTERESTS

Jose’s \lews of evolution paralleled, to a degree, those of

the Jesuit priest and paleontologist Teilhard de Chardin.

Jose felt there was no incompatibilitv' with a belief in

Cod and natural selection. He did feel that evolution

had some predestined direction and yet the forces of

natural selection were eUdent wherever one looked in

the natural world. His views are spelled out in The Con-

cept of the Species (Parodiz, 1977d).

Jose had a fascination with the explorations of Charles

DanUn in South America. He felt there was some con-

fusion regarding DanUn’s travels in South America. In

1981, he published Darwin in the New' Work] (Parodiz

1981). In this work, Jose described Daman’s journeys

throughout the South American continent and provided

historical commentaiy on what was transpiring in South

America at that time.

Jose was a passionate philatelist. He was a member of

the American Philatelic SocieU, the American Topical

Association, and its chapter that dealt wdth biology on

stamps. In addition to liis memberships and collecting

actix’ities, he also wrote articles about stamps. In the

Pittsburgh Shell Chib Bulletin, Parodiz contributed

three articles dealing \\4th shells on stamps (Parodiz

1972e, 1973g, 1974c). He published six articles in Bio-

philately dealing \\4th subjects such as cowries (Parodiz

1977i), biogeographic zones (Parodiz 1997b), and zoo-

logical nomenclature (Parodiz 1976d). As noted below,

Parodiz enjoyed reading novels. In Topical Times, he

wrote articles about Rvo literaiy figures that appeared

on stamps, Edith Wharton and Tfiornton Wilder

(Parodiz 1980c, 1998a).

Jose enjoyed reading novels. His friends describe him

as well read. His favorite author was Victor Hugo. In

addition, Jose was loud ol Heiny James, William Faulk-

ner, Somerset Maugham, John Steinbeck, and Edith

W'harton. At the time ol his death, Jose was working on

his own novel. I do not know the subject matter or how

close to completion it was.

Jose had an excellent command of Spanish and En-

glisli, botli written and spoken. He also had familiarity

with French, Portngnese, Italian, and Latin. Jose was a

membe]' of El Club Espahol de Pittsburgh — Pittsburgh’s

local chib for Spanish-spealdng people. One of my

patients, who was boin in Spain and was a member of

the club, said that her sons were fascinated by Jose’s

stories and the talks that he gave at the club. He spoke

about his work at CMNH, his trips to South America,

and his woi'k there. The Club honored him by awarding

him a Diploma de Honor al Merito’.

Jose never left his Argentinean culture behind. Jose

brought a love for dance to the United States. Jose

enjoyed the tango. In a letter to Anne LaBastille Bowes

of Cornell University, he wrote “I have the tango in my

chromosomes” (in litteris, 1969). In addition to main-

taining his Argentinean culture, Jose also embraced an

interest in opera and the symphony. On one occasion, he

and Esther took a train to New York City to attend a

musical performance, returning by train that same day.

This was on of the ways that the Parodiz’s could enjoy a

full life on Jose’s modest income.

THE LATTER YEARS

Esther Sell Parodiz passed away 22 Febrnaiy 2000. She

and Jose had been married for 48 years. Their relation-

ship was a close one. Jose’s pastor. Reverend Eric

Riesen, said that one did not speak of Esther or Jose

but of Esther and Jose.

Jose continued to live in their Pittsburgh home after

Esther’s death. In 2007, lie moved to Luther Crest Re-

tirement Commnnitv in Allentown, Pennsylvania. Jose

said that his advanced age was making it difficult for

him to maintain his house. He enjoyed his new home

and was veiv popular with the other residents. Jose’s

outgoing personalitv allowed him to fit in right away. In

a letter to his former pastor. Rev. Riesen, Jose men-

tioned that one of the great pleasures of Luther Crest

was its wonderful libraiy.

On September 3“', 2007, five months after moving,

Jose took ill. He was taken to the hospital where he was

diagnosed as having a heart attack (acute myocardial

infarction). It was determined that he was not a candi-

date for aggressive therapy. He passed away the next

day. Jose and Esther are interred in a cemeteiy on the

grounds of Saint Peters Union Church, in Maenngie,

Pennsylvania, a site not far from where Esther grew up

and Jose spent his final months.

Few people knew the breadth and depth of Jose

interests. May of ns knew one or two aspects of his life.

Many of ns came to realize his protean interests only

when we discussed his life at his memorial sendee. I for

one regret not haxdng gotten to know Jose in the

broader sense. As his friend S. Alan Boals put it, “De-

spite his long life, we feel cheated. We all would like a

few more hours wdth Jose to have one last discussion on

some topic of mutual interest.” I for one regret Jose’s

passing for I would like to have many such discussions.

I regret in hax ing gotten to know him so well in death

and not in life.

C. F. Sturm, 2009

Page 65

TAXA NAMED IN JUAN JOSE PARODIZ'S HONOR

ChitluircUa pawdizi Figueiras aud Rroggi, 1976

Diplodon parodizi Ronetto, 1962

Epif)l}ragmoi~f)l}ora parodizi Fernandez and Rnmi, 1984

Ojisiphanes mutates parodizi (Lepidoptera: Rhopalo-

cera) Bristow, 1991

Parodizia Medina, 1959

Potamolithiis parodizi Morton, 1986

Siphocifpraea parodizi Petnch, 1994

S))ixineIIa parodizi Hylton-Scott, 1952

StropJjocJieihis parodizi Klappenbach and Olazarri, 1965

Trophon parodizi Pastorino, 2005

TAXA DESCRIBED BY JUAN JOSE PARODIZ

New Genera and Snbgenera

1. Araucania new genus Parodiz, 1954

2. Astroborus new name Parodiz, 1949

3. Austrodiscus new name Parodiz, 1957

4. Calliostoma (Tropidotrochus) new snbgenns Parodiz,

1977

5. Cijclodo)\tina (Burri)igtonia) new snligenns Parodiz,

1944

6. Odontostomus (Ventania) new snbgenns Parodiz,

1940

7. Paleoancidosa new genns Parodiz, 1969

8. PaIeo])idiniuhis new genns Parodiz, 1949

9. Protoghjptus (Ri)uatida) new snbgenns Parodiz,

1946

10. Protoghjptus (Obstnissus) new snbgenns Parodiz,

1946

New Species, Subspecies, “Eorms,” and “Varieties”

1. Adelopoma paraguaijaua new species Parodiz, 1944

2. Araucania twomet/i new species Parodiz, 1954

3. Bidimulus corderoi new species Parodiz, 1962

4. Bidinudus mod new species Parodiz, 1962

5. Bidinudus (Lissoacme) ameghinoi madrijneusis new

subspecies Parodiz, 1944

6. Bidimulus (Scansicochlea) catamarcaniis new species

Parodiz, 1956

7. Bidimulus (Scansicochlea) In/Itoiiscottae new .species

Parodiz, 1956

8. Bidimidiis (Scansicochlea) riidiscidptiis new .species

Parodiz, 1956

9. Bidimulus (Scansicochlea) strobcli new species

Parodiz, 1956

10. Calliostoma (Tropidotrochus) jaijae new species

Parodiz, 1977

11. Cassis kctteri new species Parodiz and Tripp, 1993

12. Chilina stenostijlops new species Parodiz, 1963

13. Crepidida acideata foiiis new variety Parodiz, 1939

14. Cijclodontina (Scalarinella) nattkcnij)eri new species

Parodiz, 1944

15. Diplodon transandinus new species Parodiz, 1963

16. Diplodon (Ecuadorea) bristowi new species Parodiz,

1982

17. Dri/maeus hijltoni new name Parodiz, 1957

18. Drijinacus lynchi new species Parodiz, 1946

19. Drijinacus megastonius new species Parodiz, 1962

20. Drijinacus pereirai new species Parodiz, 1958

21. Diijinaeiis poeciliis tricinctus new subspecies Parodiz,

1962

22. Drijinacus rehderi new species Parodiz, 1962

23. Drijinacus waldoschmitti new species Parodiz, 1962

24. Epiphragmophora birabcni new species Parodiz,

1955

25. Efiiphragmophora feruglioi new species Parodiz,

1969

26. Epiphragmophora villavilensis new species Parodiz,

1955

27. Eriphijla miraflorcnsis new name Parodiz, 1969

28. Huniboldtiana cdithae new species Parodiz, 1954

29. Liojilacodcs bolivianus new name Parodiz, 1969

30. Lioplacodes feruglioi new species Parodiz, 1969

31. Littoridina vianai new species Parodiz, 1960

32. Lijinnaea klappenbachi new species Parodiz, 1969

33. Lijrodcs docllojuradoi new species Parodiz, 1960

34. Neocorbicula stelzneri new species Parodiz, 1969

35. Ncopctraciis stelzneri conispirus niinnta new form

Parodiz, 1948

36. Neopetraeiis stelzneri hijbrida new foi'in Parodiz,

1948

37. Ncopctraciis stelzneri nonogastaiius new form Paro-

diz, 1948

38. Ncopctraciis stelzneri peristomatns paraconispirus

new form Parodiz, 1948

39. Neopetraeiis stelzneri tinogastaniis new form Panxliz,

1948

40. Neopetraeiis stelzneri scaber new foi'in Parodiz,

1948

41. Ncritina loijolaensis new species Parodiz, 1982

42. Odontostomus fasciatus tenuiscidptiis new subspe-

cies Parodiz, 1962

43. Odontostomus weijenberghi minor new variety Parodiz,

1939

44. Odontostomus (Scalarinella} cordovanus striatus

new variety Parodiz, 1939

45. Odontostomus (Sjiixia) docllojuradoi new species

Parodiz, 1941

46. Odontostomus (Sjiixia) docllojuradoi minor new va-

riety Parodiz, 1941

47. Odontostomus (Spixia) columellaris new species

Parodiz, 1941

48. Odontostomus (Sjiixia) holmbcrgi new .species Parodiz,

1941

49. Odontostomus (Sjiixia) tiicuinancnsis new species

Parodiz, 1941

50. Paleoancidosa kenncrleiji new species Parodiz, 1982

51. Paleoancidosa jiatagonica new species Parodiz, 1969

52. Paleobidiinidus eocenicus new species Parodiz, 1949

53. Peronaeus izozoensis new species Parodiz, 1947

54. Peronaeus (Lissoacme) piintaniis new species Parodiz,

1947

55. Peronaeus (Lissoacme) rcedi new species Parodiz,

1947

Page 66

THE NAUTILUS, Vol. 123, No. 2

56. Peroitaeits (Lissoacme) toralhji avtis new variety

Farocliz, 1947

57. Peronaeus (IJssoacine) toralhji cornigatus new variety

Parotliz, 1947

58. Peronaeus (Lissoacme) toralhji nigruinbilicatiis new

varieU Parodiz, 1947

59. Phijsa ivicluuanni new species Parodiz, 1961

60. Plectostijhis argentinensis new species Parodiz, 1951

61. Plekocheihis (Eiinjtiis) ameghinoi new name Parodiz,

1962

62. Poiuacea (Effusa) pattersoni new species Boss and

Parodiz, 1977

63. Pomacea (Pomacea) protirceus new species Boss and

Parodiz, 1977

64. Potamides chaliana new species Parodiz, 1969

65. Potainolithus felipponei concordiamis new subspecies

Parodiz, 1966

66. Potainolithus peristomatus inisionum new snlispecies

Parodiz, 1966

67. Protoghjptus curainalalensis new name Parodiz,

1957

68. Protoghjptus delefangi new species Parodiz, 1946

69. Protoghjptus punctustriatus new species Parodiz,

1946 '

70. Protoghjptus (Riniatula) ininutissiinus new species

Parodiz, 1962

71. Pijrgulifera sehuena new species Parodiz, 1969

72. Siphocijpraea trippeana new species Parodiz, 1988

73. Strophocheilus (Megalobulinius) avus new species

Parodiz, 1949

74. Tajihius walteri new species Parodiz, 1969

75. Thauniastus patagonicus new species Parodiz, 1946

76. Triphora medinae new species Parodiz, 1955

77. Valvata windhauseni new species Parodiz, 1961

PUBLICATIONS

This bibliography for Jnan Jose Parodiz is probably in-

complete. The archives, at CMNH, of Jose’s activities

contain many manuscripts. It is uncertain whether they

were published. Occasionally, one contained a notation

indicating that it was published, but not where or when

it was published. When enough information was pres-

ent, a searcli for the publication was made. Most

attempts to find these publications were nnsnccesshil.

Parodiz was involved with tlie Treatise of Invertebrate

Paleontology. He was asked to write for the volume on

the Gastropoda, specifically the paits dealing with the

Bnlinmlidae, Orthalicidae, Odontostomidae, and Stro-

phocheilidae. It is uncertain whether he completed this

task as the volume was never published. In the decades

of the 1950s and 1960s, Jose published a number of

articles in Enciclopedia Barsa, a 8panish encyclopedia

published by the Encijclopedia Brilannica. While he is

listed as a contributor in this work, specific articles are

not attributed to a particular contributor. |ose did not

maititain a list ol the article that he wrote. Aside from

these caveats, this can be considered a complete bibliog-

raphy of Jose’s publications in both the scientific and

popular press.

ACKNOWLEDGMENTS

When recounting the life of one who has lived for

95 years, assistance is required to document the

significant event. In documenting the life of Juan Jose

Parodiz, I had such assistance. Richard Sell (nephew)

and Mercedes Scasso (niece) proUded personal insights

into Jose’s life and photographs of Jose and his wife,

Esther. Robert Winters shared personal recollections of

times when he was a volunteer working with Jose.

Reverend Eric M. Riesen, and S. Alan Boal shared

knowledge of Jose’s interests, especially those of a non-

scientific nature.

In 2003, two libraiy interns, Margaret Pett and

Michele Tourney, organized the archives of the Section

of Mollnsks at CMNH. Their labors made my work

orders of magnitudes easier. Bernadette Calleiy and

Xianghua Sun, librarians at CMNH, helped facilitate

interlibraiy loans and allowed me access to the archives

under their care. Especially helpful was a transcript of

the tapes from an oral history project that was under-

taken by Steve Wagner in 1990. Wagner inteniewed

Parodiz as part of a joint CMNH and Dmpiesne

Universit)' historical documentation program. Ellen

Peachy and Gini Horn (American Philatelic Society

Research Librar)d provided access to their collection

a)id helped in researching Jose’s philatelic publications.

Guido Pastorino and Manuel G. Quintana (MAGN)

were instrumental in providing information regarding

Jose’s years at MAGN. Maria Cristina Dreher Mansur

provided copies of some of Jose’s publications. Mary

Killer (John Simon Guggenheim Memorial Foundation)

helped facilitate my obtaining Jose’s records with the

Foundation.

Timothy A. Pearce (Assistant Curator, Section of

Mollnsks at CMNH) provided access to the Section’s

archives and proUded critical comments to drafts of this

paper. Amanda Zimmerman, a volunteer from the

Section, assisted with the design of the illustrations.

Lastly, Jose Jnves, also a volunteer in the Section, kindly

translated a number of Spanish-langnage documents. In

addition to the translations, he provided nsefnl editorial

comments on the text and put some of them into cultural

context. His assistance was greatly appreciated and

essential to the completion of this project.

I would also like to thank the reviewers of this paper,

who made useful comments that improved the quality of

the final product.

LITERATURE CITED

Carcelles, A. and f|. Parodiz. 1938. Moluscos del contenido

estoniacal de "Astropecten cingulatus" Sladen. Pliysis

12(44): 251-266.

Carcelles, A, and J.|. Parodiz. 1939. “Dorsaninae" argentinas

)' iirnguayas. Physis 17(49): 74.5-769.

C. F. Sturm, 2009

Page 67

Parodiz, J.J. 1939a. Las especies de “CrepUhila" de las costas

Argentinas. Physis 17(49): 685-709.

Parodiz, J. J. 1939h. Rexdsion de "Flagiodontes" y ''Scalarinella"

(Odontostominae). Physis 17(49): 711-7.34,

Parodiz, J.J. 1940. Ventania nuevo siibgenero de Oclontosfomus.

Notas del Miiseo de la Plata Zoologi'a 5(42): 227-234,

Parodiz, J.J. 1941. Four new species of Spixia from Argentina.

The Nautilus 54(3): 92-95.

Parodiz, J. J. 1942a. Los Odontostonhnos de la Argentina, pri-

inera parte. Physis 19(52): 191-218.

Parodiz, J.J. 1942h. Transgresiones oceanicas y fauna del mar

epicontinental argentino. Re\dsta Geografica Americana

18:203-211.

Parodiz, J.J. 1943. Los Odontostominos de la Argentina,

seguncia parte. Physis 19(53): 319-343.

Parodiz, J.J. 1944a. Contrihuciones al conocimiento de los

moluscos terrestres sudamericanos, I. Comunicaciones

Zoologicas del Museo de Ilistoria Natural de Monte\ideo

1(8): 1-9.

Parodiz, J.J. 1944b. Contrihuciones al conocimiento de los

moluscos terrestres sudamericanos, H. Comunicaciones

Zoologicas del Museo de Ilistoria Natural de Montevideo

1(11): 1-6.

Parodiz, J.J. 1944c. Contrihuciones al cf)nocimiento de los

moluscos terrestres sudamericanos. III. El genero Btili-

mnhis en el Territorio de la Pampa y en la Patagonia

Meridional. Comunicaciones Zoologicas del Museo de

Plistoria Natural de Montevideo 1(17): 1-8.

Parodiz, J.J. 1946a, Contrihuciones al conocimiento de los

moluscos terrestres sudamericanos, IV. Comunicaciones

Zoologicas del Museo de Ilistoria Natural de Montevideo

2(27):“l-14.

Parodiz, J.J. 1946b. Bulimulinae fosiles de la Argentina

apuntes paleontologicos y descripcion de una nueva espe-

cie. Notas del Museo de La Plata Paleontologia 11(92):

301-309.

Parodiz, J.J. 1946c. Los generos de los Bulimulinae Argenti-

nos. Revista del Museo de La Plata (N.S.) Zoologi'a

4: 303-371.

Parodiz, J.J. 1947a. Apuntes sohre "Ostrea" actuales y pleisto-

cenicas de Argentina y su ecologia (Resumen). Physis

20(56): 123-124.

Parodiz, J.J. 1947b. Contrihuciones al conocimiento de los

moluscos terrestres sudamericanos, V. Comunicaciones

Zoologicas del Museo de Ilistoria Natural de Montevideo

2(,38): 1-32.

Parodiz, J.J. 1948a. Contrihuciones al conocimiento de los

moluscos terrestres sudamericanos, \T. Comunicaciones

Zoologicas del Museo de Ilistoria Natural de Montevideo

2(46): 1-22.

Parodiz, J.J. 1948h, Sobre "Ostreu" actuales y Pleistocenicas de

Argentina y su ecologia. Comunicaciones del Museo

Argentino de Ciencias Naturales Serie Ciencias Zoologi-

cas 6: 1-22.

Fontes, E.M. and J.J. Parodiz. 1949. Guia de Naturali.stas

Sudamericanos. Talleres Graficos Lucania, Buenos Aires,

138 pp.

Parodiz, J.J. 1949a. Un nuevo gastropodo terrestre del Eoceuo

de Patagonia. Physis 20(57): 174-179.

Parodiz, J.J. 1949b. Notas sohre "Strophodieilus" fosiles de

Argentina. Plu/sis 20(57): 180-184.

Parodiz, J.J. 1949c. Joseph C. Betjuaert, Monograph of the

Strophocheilidae, a Neotropical Family of Terrestrial

Mollusks. Physis 20(57): 216-219.

Parodiz, J.J. 1949d. "Aiistrobonis'' n, uom. pro "Microlxviis"

Pilshiy, 1926. Physis 20(57): 189-190.

Parodiz, J. J. 1949e. Martin Doello Jurado (1884-1948). Phy.sis

20(57): 198-205.

Parodiz, J.J. 1949f Candido Firmino de VIello-Lcitao. Physis

20(57): 205-206,

Parodiz, J.J. 1949g. Mann, F. Guillermo, Biologia de la Autar-

tica Suramericaua. Physis 20(57): 227-229,

Parodiz, J.J. 1949h. Arthropoda. Organo oficial de la Asocia-

ciou Argentina de Artropodologia, vol 1 , no 1 , noviemhre

de 1947. Physis 20(57): 229.

Parodiz, J.J. 1949i, Revnsta de Biologia Marina. Physis 20(57):

229-2.30,

Parodiz, J.J. 1950. Emuneraciou taxouomica y ohsen'aciones

sohre los Buliumlidae Sudamericanos en las colecciones

de United States National Museum y Museum of Com-

parative Zoolog)' at llaiward College, Cambridge, Mass.

Manuscript publication. 116 pp. [This unjmhlished man-

uscript was the result of JPP’s work as a Cuggenheim

fellow'. Known copies are at CMNll, Section of Mollusks,

and the National Museum of Natural Ilistoiy, Smithso-

nian Institution.]

[Bonetto, A. and J.J. Parodiz]. 1951. Problemas tecnicos rela-

ciouados con la couseiwacidn y fomento de la riqueza

acuatica. 1) Estmlio sobre almejas nacan'feras. Jornadas

Icticas, (Racioualizacion de la Pesca), Organizadas por

los gobiernos de las provincias de Santa Fe y Entre Rios,

pp. 21-22. [anonymous, handwritten note imlicating that

authors were Bonetto and Parodiz].

[Parodiz, J.J. and A. Bonetto], 1951a. Problemas tecnicos rela-

cionados con la consenacidn y fomento de la. riqueza

acuatica 3) Investigaciones sohre almejas en la Estaciou

1 lidrobiologica de Rosario. Jornatlas Icticas, ( Racionaliza-

cion de la Pesca), Organizailas por los gobiernos de las

provincias de Santa Fe y Eutre Rios. p. 23. [anonymous,

handwritten note indicating that authors were Parodiz

and Bonetto].

[Parodiz, J. J. and A. Bonetto]. 1951h. Problemas tecnicos rela-

cionados con la conseiwaciou y fomento de la riqueza

aci'istica. 4) Estudios Hidroliioldgicos. Jornadas Icticas,

(Racionalizaciou de la Pesca), Organizadas por los gohier-

uos de las provincias de Santa Fe y Entre Rios. pp. 23-24.

[anonymous, handwritten note indicating that authors

were Parodiz and Bonetto].

Parodiz, J.J. 1951a. Metodos de conquiliometria. Physis

20(58): 241-248.

Parodiz, J.J. 1951b. Una nueva especie de "Plecfosti/his''

(Castr. Pulm.) de la Argentina. Physis 20(58): 334-335.

Parodiz, J.J. 1951c. Visita a la Argentina del Dr. Henry A.

Pilsbiy Physis 20(58): 345-346.

Parodiz, J.J. 1951d. Centenario de una rev'ista malacolbgica:

Journal de Conchyliologie. Physis 20(58): 346-347.

Parodiz, J.J. 1951e. Societ)' of Systematic Zoologv. Physis

20(58): 347.

Parodiz, J.J. 1953. Mayflower 1-7300. Carnegie Magazine

27: 17(1-172.

Parodiz, J. J. 1954a. Araucania twoinci/i n. gen. n. sjr del sur de

Chile (Gastrop. Pulm,). Neotropica 1(2): 17-18.

Parodiz, J. J. 1954b. A new species oi Humhohitiaua from Te.x-

as. The Nautilus 67(4): 107-108.

Parodiz, J.J. 1954c. Ensayo para determinar el grado de rela-

cion eu taxonomia. Physis 20(59): 444—452.

Parodiz, J.J. 1954d. Arneghiuo’s Centennial. Carnegie Maga-

zine 28: 240-243.

Page 68

THE NAUTILUS, Vol. 123, No. 2

Parodiz, J.J, 1955a. Una mieva especie de Triphora del Uru-

guay (Moll. Prosobr.). Neotropica 1(4): 59-60.

Parodiz, J.J. 1955b. Dos mievas Epiphragmophora de la

Argentina (Gastr. Piilm.). Neotropica 1(6): 93-95.

Parodiz, J.J. 1955c. La validez del noinbre Pahidestrina

d'Orbigny 1839. Neotropica 1(6): 9.5-96.

Parodiz, J.J. 1955d. Shell remains from the Globe Hill site -

Appendix II. 1955. In: Mayer-Oakes, Excavation at the

Globe Hill Shell Heap. West Virginia Archaeological Soci-

ety Publication 3: 29-30.

Parodiz, J.J. 1956a. Notas sobre “Plujsa" (Gastr. Pulm. Baso-

mat.). Neotropica 2(7): 19-21.

Parodiz, J.J. 1956b. Cuatro nuevas especies de Scansicochlea

de Argentina (Moll. Pulm.) sub. genero Scansicochlea

Pilsbiy 1930. 1“ parte. Neotropica 2(8): 59-64.

Parodiz, J. J. 1956c. Cuatro nuevas especies de Scansicochlea

de Argentina (Moll. Pulm.). 2“ parte. Neotropica 2(9):

77-80"

Parodiz, J.J. 1956d. Notes on the Ireshwater snail Leptoxis

[Miidalia) carinata (Bruguiere). Annals of the Carnegie

Museum 33(23): .391-405.

Parodiz, J.J. 1957a. Catalogue of the land Mollusca of Argen-

tina. The Nautilus 70(4): 127-135.

Parodiz, J.J. 19.57b. Ceiitenario de d'Orbigny. Neotropica

3(10): 1-6.

Parodiz, J.J. 1957c. New records of fresh-water gastropods

from the Bahama Islands. Annals of the Carnegie Muse-

um 35(1): 1-9.

Parodiz, J.J. 1957d. Catalogue of the land Mollusca of Argen-

tina (cont.). The Nautilus 71(1): 22-30,

Parodiz, J.J. 1957e. Alcide d'Orbigny. New York Shell Club

Notes 33: 2-5.

Parodiz, J.J. 19571. Jean Guillaume Bruguiere (1750-1798).

Minutes, Conchological Club ol Southern California

172: 2-4.

Parodiz, J.J. 1957g. Catalogue of the land Mollusca of Argen-

tina (concluded). The Nautilus 71(2): 63-66.

Parodiz, J.J. 1957h. Moluscos marinos de las ingresiones del

Pleistoceno v Holoceno. pp. 58-60. In: Bordas, A. F.

Argumentos paleontologicos y climaticos para establecer

relaciones estratigraficas del Pleistoceno-Holoceno de

Argentina. Ameghiniana 1: 51-79

Parodiz, J.J, 1958a. A new Bolndan land snail of the genus

Dnpnaeus. Breviora 95: 1-3.

Parodiz, J.J. 1958b. Land snails of Powdermill Nature Reserve.

Powdermill Nature Reserve Educational Release 13: 1-2.

Parodiz, J.J. 1958c. List of the fresh-water snails of Pennsylva-

nia, Release of the Section of Inveitebrates, Carnegie

Museum of Natural History, Pittsburgh, 3 pp.

Parodiz, J.J. 195Sd. Art and Nature Bookshelf: Cowry Shells of

the World Seas by Joyce Allan. Carnegie Magazine

.'32: 102-104.

Parodiz, J. J, 1959. Brasilia. Carnegie Magazine 33: 4.5-48.

Parodiz, J.J. 1960a. Neotype for Lijrodes giiaranitica Doering

and descripHon of a new species. The Nautilus 74(1): 2.3-26.

Parodiz, J.J. 1960b. Atyidae neotropicales (Cnistacea-Nafantia-

Caridea) en la coleccion del Museo Carnegie. Neotropica 6

(20): .38-40.

Parodiz, J.J. 1960c. Una nueva especie de Litforidina {Gasfr.

Prosobr.) en agua salada, de la Rioja. Neotropica 6(21):

89-90,

Parotliz, J.J. 1960d. A focus of interest in Marine Hall. Carne-

gie Alagazine 34: .54-55.

Parodiz, J.J. 1960e. Land shells of Westmoreland County.

Powdermill Nature Reserve Educational Release 26: 1-3.

Parodiz, J.J, 1961a. New and little known Plujsa from the

Paleocene of Patagonia. Annals of the Carnegie Museum

36(1): 1-4.

Parodiz, J.J. 1961b. Notes on Valvatidae from early Tertiaiy of

South American, wdth a new species. The Nautilus 75(1):

16-18.

Parodiz, J.J. 1961c. Moluscos terrestres, hasta ahora conoci-

dos, de la Provincia de Mendoza. Revista Cientifica de

Investigaciones del Museo de Historia Natural de San

Rafael 1(3): 59-62.

Parodiz, J.J. 1962a. Los moluscos marinos del Pleistoceno rio-

platense. Comunicaciones de la Sociedad Malacologica

del Uruguay 1(2): 29-46.

Parodiz, J.J. 1962b. New and little-known species of South and

Central American land snails (Bulimulidae). Proceedings

of the United States National Museum 113(3462): 429-

456.

Parodiz, J.J. 1962c. On South Atlantic Columbellidae. The

Nautilus 76(2): 74.

Parodiz, J.J, 1962d. Variabilidad en “Cyclodontina (Spixia)

Doello-Juradoi" Par. (Gastropoda, Pulmonata). Revista

del Museo Argentino de Ciencias Naturales “Bernardino

Rivaclavia” Ciencias Zoologicas 8(5): 69-80.

Parodiz, J.J. 1962e. Seashells by the Seashore: new exliibit at

Carnegie Museum. Carnegie Magazine 36: 333-336, 342.

Parodiz, J.J. 1963a. La extraordinaria fauna del Rio Uruguay y

sus relaciones. Comunicaciones de la Sociedad Malacolo-

gica del Uruguay 1(5): 103-110.

Parodiz, J.J. 1963b. New fresh-water Mollusca from tire Eogene

of Chile and Patagonia. The Nautilus 76(4): 145—147.

Parodiz, J.J. 1963c. Observaciones anatomicas sobre Omalonyx

patera Doer., con una nota biografica acerca de Adolfo

Doering (1848-1926). Sterkiana 12: 1-7.

Parodiz, J.J. 1963d. When the land was young. Americas 15(8):

22-25. [a Spanish version is titled Cuando la Tierra era

Joven].

Parodiz, J. J. and A. A. Bonetto. 1963. Taxonomy and zoogeo-

graphic relationships of the South American naiades

(Pelecypoda: Unionacea and Mutelacea). Malacologia 1

(2): 179-213.

Parodiz, J.J. 1964. Freshwater Mollusca from the early Tertiaiy

of Patagonia. Annual Report of the American Malacologi-

cal Union 31: 6-7.

Parodiz, J.J. 1965a. Relaciones y evidencias paleontologicas de

PotanioUthus. Comunicaciones de la Sociedad Malacolo-

gica del Uruguay 1(9): 273-278.

Parodiz, J. J. 1965b. The hydrobid snails of the genus Potamo-

lithus (Mesogastropoda-Rissoacea). Sterkiana 20: 1-38.

Parodiz, J.J. and L, Hennings. 1965. The Neocorbicula (Mol-

lusca, Pelecypoda) of the Parana- Uruguay basin. South

America. Annals of the Carnegie Museum 38(3): 1-96.

Parodiz, J.J. 1966a. Two new subspecies ol Potamolithus. The

Nautilus 80(2): 56-58.

Parodiz, J.J. 1966b. Shell collecting by king and commoner.

Shell Collecting- An Illustrated Histoiy by S. Peter

Dance. Caniegie Magazine 40: 347-350.

[Parodiz, J.J.] 1966c. How the word malacology came into

being. Pittsburgh Shell Club Bulletin 1: 10. [anonymous,

written by JJP].

Parodiz, J.J. 1967. Ty|oes of North American Unionidae in the

collection of the Carnegie Museum. Sterkiana 28: 21-30.

C. F. Sturm, 2009

Page 69

Parodiz, J.J. 1968a. Annotated catalogue of tlie genus Diplo-

don (Unionacea-Hyriidae). Sterkiana 30: 1-22.

Parodiz, J.|. 1968b. Geographical distribution of niollnsks.

Pittsburgh Shell Club Bulletin 3: 4-5.

Parodiz, J.J. 1968c. The ainatenrs’ use of genera and subge-

nera. Pittsburgh Shell Club Bulletin 3:9.

Parodiz, J.J. 1969a. The Tertiaiy non-marine Mollusca of

South American. Annals of the Carnegie Museum 40:

1-242.

Parodiz, J.J. 1969b. Subgenera in Ct/praea. Pittsburgh Shell

Club Bulletin 4: 10-11.

Parodiz, J.J. 1969c. flow many shells? Pittsburgh Shell Club

Bulletin 4: 16.

Leich, R. and J.J. Parodiz. 1970. Ludwig van Beethoven

(1770-1827). Carnegie Magazine 44: 404^07.

Parodiz, J.J. 1970a. Diphdon peraefonnis (Lea). Comuni-

caciones de la Sociedad Malacologica del Llniguay 3(19):

1-6.

Parodiz, J.J. 1970b. Atysidae, emendation for Atyidae. Pitts-

burgh Shell Club Bulletin 5: 4.

Parodiz, J.J. 1971a. A new record for Ci/praea surinamensis

Perry 1811. Pittsburgh Shell Club Bulletin 6: 6.

Parodiz, J. J. 1971b. Edgar Allan Poe Conchologist. Pittsburgh

Shell Club Bulletin 6: 17.

Parodiz, J.J. 1971c. Types in the species-group. Pittsburgh

Shell Club Bulletin 6: 19.

Parodiz, J.J. 1972a. The “superspecies” DipJodon dclodontus.

Bulletin of the American Malacological Union 37: 34.

[Parodiz, J.J.] 1972b. Editorial: shell prices. Pittsburgh Shell

Club 7: 2. [anonymous, written by jJP].

Parodiz, J.J. 1972c. Olivia saijana Ravenel, O. reticularis

Lamarck and their variations. Pittsburgh Shell Club Bul-

letin 7: 11-12. [note misspelling of Oliva]

Parodiz, J.J. 1972d. The tyj^es of genera. Pittsburgli Shell Club

Bulletin 7: 13.

Parodiz, J.J. 1972e. Shells on stamps-I. Pittsburgh Shell Club

Bulletin 7:16-19.

[Parodiz, J.J.] 1972f A weathei'wise snail. Pittsburgh Shell

Club Bulletin 7: 22. [anonymous, w'ritten by J[P].

Parodiz, J.J. 1973a. The species complex of DipJodon delodon-

tus (Lamarck) (Unionacea-Hvriidae). Malacologia 14

(1-2): 247-270. (Proceedings of the Fourth European

Malacological Congress (Geneva, 7-11 September 1971)

Eugene E. Bindur, ed.)

[Parodiz, J.J.] 1973b. Editorial. Pittsburgh Shell Club Bulletin

8: 2. [anonymous, w'ritten by JJP].

[Parodiz, J.J.] 1973c. Molluscan names, recently validated or

suppressed. Pittsburgh Shell Club Bulletin 8: 5-6. [anon-

ymous, written by JJP].

Parodiz, J.J. 1973d. How to evaluate subspecies. Pittsburgh

Shell Club Bulletin 8: 8.

Parodiz, J. J. 1973e. Gastropod conchometry. Pittsburgh Shell

Club Bulletin 8: 14-16.

Parodiz, J. J. 1973f Book Re\4ew: Australian Shells by Wilson

and Gillett. Pittsburgh Shell Club Bulletin 8: 16.

Parodiz, J.J. 1973g. Shells on stamp.s-II. Pittsburgh Shell Club

Bulletin 8: 20-23.

Parodiz, J.J. 1973h. The operculum. Pittsburgh Shell Club

Bulletin 8: 25.

Parodiz, J.J. 1974a. DipJodon cJ^armanus (d'Orbigny): a revi-

sion. Bulletin of the American Malacological Union 39:

18-19.

Parodiz, J.J. 1974b. Variation and survival. Pittsburgh Shell

Club Bulletin 9: 6-7.

Parodiz, J.J. 1974c. Shells on stamp.s-lll. Pittsburgh Shell

Club Bulletin 9: 1 1-15.

Parodiz, J.J. 1974d. A checklist and distribution of western

Atlantic Columbellidae. Pittsburgh Shell Club Bulletin 9:

17-19.

Parodiz, J.J. 1974e. An early obsemition on ConcJwJepas per-

uviana Lam. (Mnricidae). Pittsburgh Shell Club Bulletin

9: 19.

Parodiz, J.J. 19741. An introduction to the histoiy of malacolo-

gy. The Shell Case (Naples Shell Club) 4(1): 6-7.

Parodiz, J.J. 1975a. Two naiads from western Pennsylvania.

Carnegie Magazine 49: 300-301.

Parodiz, J.J. 1975b. Early American concholog)'. Carnegie

Magazine 49: 411-417.

Parodiz, J.J. 1975c. Prologo. In: J. Olazarri. Para nna Ilistoria

de la Malacologia en el Uruguay. Privately published,

Montevideo, Uruguay, p. v. [of v + 121 pp.]

Parodiz, J.J. 1975d. Recently collected freshwater shells. Pitts-

burgh Shell Club Bulletin 10: 3-4.

Parodiz, J.J. 1975e. From a Fuegian notebook. Pittsburgh

Shell Club Bulletin 10: 15-16.

Parodiz, J.J. 1975f Book Review: American Seashells

by R. Tucker Abbott. Pittsburgh Shell Club Bulletin 10:

16-17.

Parodiz, J. J. 1975g. Proptero Jaevissiina from Oklahoma (Unin-

onidae-Lampsilinae). Pittsburgh Shell Club Bulletin 10:

19. [note misspelling of Unionidae is in the printed title.]

Parodiz, J.J. 1975h. Peiple.xing OJiva and creative evolution.

Self-published, Pittsburgh, 6 pp.

Parodiz, J.J. 1976a. A new species of CaJJiostonm from the

Tertiary of Virginia. Pittsburgh Shell Club Bulletin 11:

2-3.

Parodiz, J.J. 1976b. Book Review: Cowries by John Tavlor.

Pittsburgh Shell Club Bulletin 11: 20.

Parodiz, J.J. 1976c. The tyrian puqile. Pittsburgh Shell Chib

Bulletin 11: 22-25.

Parodiz, J. J. 1976d. Principles of Zoological Nomenclature.

Biophilately 25(2): 85-89.

Parodiz, J.J. 1976e. Shells on Stamps-I. Biophilately 25(2): 96-

102.

Parodiz, J.J. 1976f Shells on Stamps-II. Biophilately 25(3):

139-142,

Boss, K. J. and J.J. Parodiz, 1977. Paleospecies of Neotropical

ampullariids and notes on other fossil non-marine South

American gastropods. Annals of the Carnegie Museum

46(9): 107-127.

Parodiz, J.J. 1977a, Alberto Carcelles (1897-1977). Comunica-

ciones de la Sociedad Malacologica del Uruguay 4(31-32):

289-293.

Parodiz, J.J. 1977b. A new subgenus and new species of Mio-

cene CaJJiostonm (Archaeogastropoda-Trochidae). Annals

of the Carnegie Museum 46(8): 101-106.

Parodiz, J.J. 1977c. New species of CaJJiostonm from the

Tertiary of Virginia. Bulletin, Conchologists of America

10: 3.

Parodiz, J.J. 1977d. The concept of the species. Section of

Invertebrates, Carnegie Museum. Pittsburgh, PA. 12 pp.

Parodiz, J.J. 1977e. Mollusca. In: Biota Aciuitica de Sudamer-

ica Austral. San Diego State University, San Diego, CA.

S.H. llulbert (Ed.). 320-329.

Parodiz, J. J. 1977f In Memoriam Alberto R. Carcelles (1897-

1977). Pittsburgh Shell Club 12: 2.

Parodiz, J.J. 1977g. Marine shells from Saudi Arabia found

"inland”. Pittsburgh Shell Club Bulletin 12: 5,

Paa;e 70

THE NAUTILUS, Vol. 123, No. 2

Parocliz, f.|. 1977h. An introduction to the histon' of nialacolo-

g)' II. The early period. The Shell Case (Naples Sliell

Club) 5(1): 6-8.'

Parodiz, |.J. 19771. Cowries. Biophilately 26(1): 7-11.

Parodiz, ].[. 1977). Shells on recent stamps from Somalia.

Biophilately 26(2): 70-73.

Parodiz, J.J. 1978a. Dr. H.M. Lemche (1904-1977). Pittsburgh

Shell Club Bulletin 13: 11-12.

Parodiz, J.J. 1978b. Re-description of Liinatia sanctivincentU

(Brooks). Pittsburgh Shell Club Bulletin 13: 13-16.

Parodiz, J.J. 1979a. Anatomy and taxonomy of Protog^li/ptus

quitensis (Pfeiffer) (Gastropoda, Pulmonata, Bulimuli-

dae). Malacologia 18(2): 11, 5-122.

Parodiz, J.J. 1979b. [Letter to the editor]. New York Shell

Club Notes 251: 6-7.

Parodiz, J.J. 1979c. Notes on Southern k^olutes. Pittsburgh

Shell Club Bulletin 14: 12-13.

Parodiz, J.J. 1979d. Monoplacophorans. Pittsburgh Shell Club

Bulletin 14: 1,3-14.

Parodiz, J.J. 1979e. Marine Mollusca collected in Yucatan.

Pittsimrgh Shell Club Bulletin 14 (Supplement): 3-20.

Parodiz, J.J. 1980a. Origin and distribution of the continental

malacofauna of South America. Ilaliotis 10(2): 112.

Parodiz, J.J. 1980b. Invertebrates. Carnegie Magazine

,54: 12-16,

Parodiz, J.J. 1980c. Edith \Miarton. Topical Times 31(4): 24—25.

Parodiz, J.J, 1981. Danvin in the New World. E. J. Brill, Lei-

den. i.\ + 143 pp.

Bristow, C.R. and J.J. Parodiz. 1982. The stratigraphical pale-

ontology of the Tertiaiy non-marine sediments of Ecua-

dor. Bulletin of the Carnegie Musetim of Natural Histoi")'

19: l-,53.

Parodiz, J.J. 1982a. Distribution and origin of the continental

South American malacofauna. Malacologia 22(1-2):

421—425. [Proceedings of the 7*'' International Malacolog-

ical Congress, Aug. 31-Sept. 7, 1980, J. M. Gaillard, ed.]

Parodiz, J.J. 1982b. Note on Invertebrata. Atiqot 15: 101.

Parodiz, J.J. 1983. Lnther and His Times, PrivateK' published.

Pittsburgh, PA. 37 pp.

Parodiz, J. J. 1988. A new species of Siphoci/praca (Gastropoda,

Cypraeidae) from the Neogene of southwest Florida. An-

nals oi the Carnegie Museum 57(3): 91-97.

Parodiz, J.J. and J.T. Tripp. 1988. Types of Mollusca in the

collection of the Carnegie Museum of Natural Histoiy,

Part 1. Bivalvia and Gastropoda (Prosobranchia and

Opistobranchia). Annals of the Carnegie Museum 57(5):

111-1,54.

Parodiz, J.J. and E. Balech 1992. El Museo Argentine de

Ciencias Naturales “B. Rivadavia” ... en pantuflas.

Mimeographed, Buenos Aires. 91 pp.

Parodiz, J.J. and J.J. Tripp. 1992. The Neogene Cassidae of

southern Elorida, with description of a new species of

Cassis (Gastropoda: Prosobranchia). Annals of the Carne-

gie Museum 61(4): 317-325,

Parodiz, J.J. 1996. The taxa of fossil Mollusca introduced by

Hermann von Ihering. Annals of the Carnegie Museum

65(3): 183-296.

Parodiz, J.J. 1997a. Paleoespecies de Rhipidodontini (Bivalvia,

Hyriidae) Taxonomia y Paleogeografia. [Abstract] In: XV

Encontro Brasileiro de Malacologia, Resumos, 21 a 25 de

Julho de 1997. Elorianopolis - Santa Catarina: Sociedade

Brasileira de Malacologia, 1999. v, 1, p. 18.

Parodiz, J.J, 1997b. Zoogeography - a bio-geographic topic.

Biophilately 46(2): 81-87.

Parodiz, J.J. 1998a. Thorton Wilder. Topical Times 49(1):

,36-37.'

Parodiz, J.J, 1998b. Understanding overprints. Topical Times

49(5): ,53-56.

Parodiz, J.J. 1999. Reminiscences on malacology in the twenti-

eth centurx' (abstract). American Malacological Society

6.5**' Annual Meeting Program and Abstracts, 4-9 July

1999, Pittsburgh, Pennsylvania, USA. p. 45.

Mansur, M.C.D., J.J. Parodiz, J. Olazarri, PA. Rodrigues,

L. M.Z. Richinitti, C. P. dos Santos, and F. de Borba Cunha.

1999. Moluscos bivalves do Rio Uruguai: Taxonomia, varia-

yoes na diversidade e morfologia. [Abstract] In: XVI Encon-

tro Brasileiro de M;dacologia Programa e Resumos, 12 a 16

de Julho de 1999. Recife - Pernambuco, Brasil: Sociedade

Brasileira de Malacologia, 1999. v. 1. pp. 5,5-56.

Sturm, Jr., C.F., J.J. Parodiz, and J. Rawlins. 1999. One hundred

years of imilacology at the Carnegie Museum of Natural

Histoiy (Abstract-poster presentation). American Malaco-

logical Society 65*'' Annual Meeting Program and

Abstracts, 4-9 July 1999, Pittsburgh, Penn.sylvania, USA.

[late entiy, abstract distributed with registration materials].

Parodiz, J.J. and L. S. Morton. 2002. Nueva posicion de Diplo-

clon htjanensis Ihering (Bivalvia, Hyriidae) del Pleistocene

tardfo. Comunicaciones Cientfficas y Tecnologicas, Uni-

versidad Nacional del Nordeste, Corrientes, 21-2,5 de

Octubre de 2002.

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THEr^NAUTILUS

CONTENTS

Volume 123, Number 3

Septe)ul)er 28, 2009

ISSN 0028-1344

Proceedings oe the Symposium

‘‘Neogastropod Origins, Phylogeny, Evolutionary Pathways

AND Mechanisms” held during the 2007 World Congress of

Malacology, Antwerp, Belgium, 15-20 July 2007

Guest Editors M.G. Harasewych and Ellen E. Strong

M.G. Harase^vych

Ellen E. Strong

Yuri I. Kantor

Alexander Fedosov

Alisa R. Kosyan

Yuri I. Kantor

Alisa R. Kosyan

Maria Vittoria Modiea

Mareo Oliverio

Mareo Ta\aani

Lorenzo Angeletti

Mark Diinech

Constantine Mifsud

Andre Freiwald

M.G. Harasewyeh

Mareo Oliverio

Mareo Oliverio

Andrea Bareo

Alexandra Riehter

Maria Vittoria Modiea

Gregoiy S. Herbert

Gregory P. Dietl

Helena Fortunato

Luiz Rieardo L. Simone

Jennifer Sliko

Luiz Rieardo L. Simone

Gregory S. Herbert

Didier Merle

Preface

71

Morphology and development of the valve of Leihlein: Possilile evidence

for paraphyly of the Neogastropoda 73

Phylogenetic analysis of the subfamily Colinae (Neogastropoda: Bnccinidae)

basetl on moiphological characters 83

The anatomy and relationships ol Troschelia (Neogastropoda: Bnccinidae):

New eridence for a closer fasciolariiddniccinid relationship? 95

Coralliophilinae (Gastropoda: Mnricidae) associated with deep-water

coral banks in the Mediterranean 106

The coralliophiline (Gastropoda: Mnricidae) radiation: Repeated

colonizations of the deep sea? 113

Extremely slow feeding in a tropical drilling ectoparasite, Viftilaria salebrosa

(King and Broderip, 1832) (Gastropoda: Mnricidae), on mollnscan hosts

from Pacific Panama 121

Unusnal anatomy of the ectoparasitic mnricid Vitiilaria salebrosa

(King and Broderip, 1832) (Neogastropoda: Mnricidae) from the Pacific

coast of Panama 137

Notice

223

THE NAUTILUS 123(3):71, 2()09

Page 7 1

Proceedings of the Symposium

“Neogastropod Origins, Phylogeny, Evolutionary Pathways

AND Mechanisms” held during the 2007 World Congress of

Malacology, Antwerp, Belgium, 15-20 July 2007

Guest Editors M.G. Harasewych and Ellen E. Strong

Preface

By any measure, neogastropods are an extraordinary ex-

ample of evolutionary sirccess. They appear'ed abrrrptly,

at least irr recognizable form, during the Albiarr (TOO

rnya), and radiated rapidly at a variety of taxonomic

levels to become the dominant predatory gastropods in

benthic rrrarine communities from the tropics to the

poles, from intertidal depths to the abyssal plain. Sever'al

gr'oups have further e.xtended their ramge into the ocean

trenches and into fresh water'.

These animals have been studied extensively, and

from nrrrnerous perspectives. Their economic impor-

tance is considerable, both as a harvestable food soirrce

and as predator's of other commercially important rnol-

lusks. Neogastropod glandnlar secretions have also been

of significant irnportarrce thr'ough the ages. Pr'odrrctiorr

of Tyrian Puqile ir'om hy|3obranchial gland secretions of

rnuricids by the Minoans and Phoenicians has been

traced to the 20^’’— 18‘'' centrrries BC. In the present

day, conotoxins, produced by the verrorrr glands of tox-

oglossans, are Ireir'rg extensively studied for pr'oveu and

potential biomedical applications.

Phylogenetic strrdies orr neogastr'opods aborrrtd at a

variety of taxononric levels. Because of the rapid prolif-

eratiorr of lineages, each with tenderrcies to modify or-

gan systems in parallel, nrost major groups are well

characterized mor-^^hologically, yet precious few charac-

ters have been identified that sirpport relationships

within and betweerr them. The absence of congruent

patterns of character distribirtion in rrrajor organ systems

has corrfounded initial attempts at phylogenetic infer-

ence based on rrrorphological characters. Mor'e r'ecent

studies trsiug DNA seqirences of rutclear and rnitochon-

dr-ial genes have also prodrrced contr adictory or eqirivo-

cal r'esrrlts, while analyses of datasets combining

molecular and morphological character's have far'ed or'rly

slightly better'.

Despite corrsiderable arrd concerted resear'clr effort

spannirrg decades, ther'e are few rprestions that can yet

be an,swered with any degree of confidence. Wherr it

comes to the most basic (prestions, we know rernar'kably

little about neogastropods. Sitch qrrestions as what?

wherr? where? how? and why? still irrtrigue trs. When

r'ephrased in the language of rnoderrr biology, they be-

come qrrestiorrs of monophyly, sister taxa, s\'rrapomor'-

phies, fossil record, evolirtionarx- rates, hiogeography, as

well irrrprir'ies that call into qrrestion oirr irnderstanding

of basic evolirtionary and gerretic rrrechanisrns. Arrswers

to many of these rprestions still evade rrs.

The first workshop focrrsing on the svsternatics, phy-

logeny arrd biology of the Neogastr'opoda was corrvened

in Merrfi, Italy "(June 14—18, 2000), and followed by

workshops on neogastropods at the Srnithsoniarr Alar'ine

Station at For't Pier'ce, Flor'ida (Augrrst 4—13, 2004) and

the Srrrithsoniarr Tropical Resear'ch Irrstitute at Naos,

Pananra (Jamrary 29— Fehr'irary 13, 2006). A four'tlr

workshop wall he hosted by Centr'o Nacional Patagonico

(CENPAT) in Prrer'to Madr-\m, Ar'gerrtina (November

9-13,2009).

The papers in this vohrrne were pr'esented as part of

the syrnposiirm on NEOGASTROPOta OiriGtNS, Phylogeny,

E\'olutionary Pathways and Mechanisms that was con-

vened dirring the 16^'' World Congress of Malacolog)'',

held in Antwer'p, Belgiirrn, on Arrgrrst 16—17, 2007.

Many of the par'ticipants in the neogastr'opod wor'ksliops

wer'e contrihirtor's to this syrrrposiurrr. The r'esear'cir pre-

sented here r'epresents a hr'oad spectnrm of appr'oaches

to varied aspects of neogastr'opod evohrtiorr. We hope

that these papers wall shed light orr some of the curr'errt

questions, br'irrg other trnr'esolved issrres into shariaer

focus and stinrrrlate further r'esear'cir orr tliis intriguing

gr'oup of gastr'opods.

We ar'e gr'atehrl to Prof. Dr'. Thierr-Y' Backeljau and to

the or'ganizers of the Congr'ess for the invitation to or'ga-

nize this symposium. Thairks ar'e dire to all of the par'ti-

ciparrts who pr'esented their resear'ch and to the marry

co-arrthor's who were preserrt, as w'ell as to oirr marry

friends and colleagires who wer'e unable to atterrd but

never'theless played a r'ole in its srrccess through their

corrtrilnrtions.

M.G. Hara.sewych

Ellen F. Strong

National Museirm ol Natrrr'al llistor'v

Smithsoiriarr Institution

Washirrgton, 14C 20013-7012 USA

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THE NAUTILUS 123(3):73-82, 2009

Page 73

Morphology and development of the valve of Leiblein: Possible

evidenee for paraphyly of the Neogastropoda

Yuri I. Kan lor

Alexander Fedosov

A.N. Severtstn Institute ol Ecolog)' and Evolution

Russian Academy of Sciences

33 Leninski Prospekt

Moscow 1 19071

RUSSIA

kantor@malaco-se\'in.msk.rn

ledosov' [email protected]

ABSTRACT

Neogastropoda are generally considered to he monophyletic,

although their monophyly is usually challenged in molecular

phylogenies. Such results suggest that serious reconsideration

of tlie synapomoqvliies and antapomorphies defining the group

is needed. One of the nncontroversial monophyletic groupings

wathin the neogastropods is the supertamily Rnccinoitlea. This

taxonomically rich clade lacks two out of three characters tfiat

are considered to be key antapomorphies of Neogastropoda,

namely accessoiy salivaiy glands and the rectal gland. The only

other autapomoiphy that unites Rnccinoidea with the rest ol

Neogastropoda is the valve of Leiblein. This stndv of the mor-

phology of the valve of Leiblein of different neogastropods

(two species of Raphitoniinae, Conidae, one of Muricidae,

one of Nassariidae, one of Buccinidae, one of Cancellariidae,

and one of Olividae) revealed its strong morphological hetero-

geneit)’. Published and original data on the einbnonic devel-

opment of the valve in Buccinidae and Muricidae demonstrate

that the valve originates from different sections of the anterior

foregut. Preliminaiw data indicate that the homology of the

valve of Leiblein within Neogastropoda is, at best, ([uestion-

able. This casts fnrtlier doubts on the monophviy of the Neo-

gastropoda, which probably include at least two stems.

Achlitional keijivonls: Gastropoda, phylogeny, monophyly,

alimentan' .system, development

INTRODUCTION

Tlie origin and phylogeny of the Neogastropoda were

the subjects of many pnhlications over the past decades.

Several hyjrotheses on the sister groups were also pro-

posed, including higher “mesogastropods” of the order

Tonnoidea (Amandrnt, 189S; Graham, 1941, and more

recently Riedel, 1994, 2()()0), and an “arcliaeogastroptxr

or primitive “mesogastropod” (Ponder, 1974). Vloipho-

logical analyses of Strong (2003) suggested otlier possi-

ble affinities for the group, wdth the nearest relatives of

Neogastropoda being Epitoniidae, Cvpraeidae and Nati-

cidae (Tonnoidea were not represented in these analy-

ses). Tl le molecular analysis of Colgan et al. (2007)

found relationships between neogastropod families (ei-

ther individually or as groups) and Tnrritellidae, Tonnoi-

dea, Stroinboidea, or Cvpraeidae.

Since the publication of Golikov and Starobogatov

(1988), mosdy overlooked by western malacologists, the

monophyly of the Neogastropotla was not contested. These

authors suggested that Bnccinilonnii (to which they attnb-

nted majorits' oi neogastropods, but also include Triphoi-

idae) and Gonilormii (in whicli they included Gonoidea,

Mitridae, Gancellailidae, and Fvrenoidea) oi'iginated in-

tlependently, the former ones fi'om Amberleyoidei while

the latter from Tnrbinoidei. The idea that Gonoidea

(= Toxoglossa) stands well apart from the rest of the neo-

gastropods was also supported by Sheritlan, Vin Mol, and

Bouillon (1973) and Shimek and Kolm (1981).

Kantor (2002) summarized the major apomorphies of

the Neogastropoda and concluded that they are mono-

phyletic. Among recent morpholog>'-based phylogenetic

analyses. Strong (2003) and Ponder et al. (2008) sup-

ported the monophyly ol the Neogastropoda. In Ponder

et al. (2008) a Bayesian analysis of a combinetl morphol-

ogical dataset and the molecular data also supported the

monophyly of the Neogastropoda.

The monophyly of Neogastropoda has usually been chal-

lenged, albeit weakly, in molecular analyses (llarasevwch

et ak, 1997; Golgan et ak, 2000, 2003, 2007; Rit'del, 2()00;

McArthur and Ilarasewycli, 2003). More details of

different molecular data analvses can be lonnd in Golgan

et ak (2007). It should be spc'cifically noted that in a

number ol analyses the Tonnoidea were nested wdthin the

Neogasti'opoda.

A sniwey of e.xisbng theones and suggestions reveals

that nearly eveiw possible evolntionan-’ scenario foi' the

Neogastropoda, and nearlv all possible relationships have

already been proposed. A con.sensns has not yet becni

Page 74

THE NAUTILUS, Vol. 123, No. 3

achieved, and tlie situation is not becoming clearer

witli atldition of more moqrhological and/or molecular

data.

A major current problem is the incongmence between

molecular and morphological analyses botli in terms of

the monopliyly of Neogastropoda and the composition of

the clade. The answer may lie in the erroneous interpre-

tations ol the s\niapomoiphies and antapomoiphies defin-

ing Neogastropoda. Taylor and Aloi'ns (1988) and, more

recently, Kantor (2()02) summarized and discussed in de-

tail the antapomoiphies of Neogastropoda. Three autapo-

moiphies of neogastropods have been found so far: the

presence of a second paii' of salivaiy glands (accessoiy

salivaiy glands, differing in morphology and histology

from the primaiy salivaiy glands), the presence of a valve

of Leiblein, and the presence of an anal, or rectal gland.

It has been nnanimonsly accepted that these three struc-

tures are homologous wdthin the Neogastropoda.

One of the nncontroversial inonophyletic groups

within Neogastropoda is the snperfamily Bnccinoidea

Ralinesque, 1815. This clade was considered as highly

ad\ anced by Kantor (1996), or the sister taxon to the rest

of the neogastropods (Ponder and Lindberg, 1997). Buc-

cinoidea lack both accessoiy salivaiy glands and a rectal

gland, leaving the valve of Leiblein as the single remain-

ing antapomorphy that is present in all major branches

ol the Neogastropoda.

The valve or phaiynx ol Leiblein is usually described

as pear-shaped organ, consisting of a posteriorly directed

cone-shaped protuberance that is enclosed in a chamber

formed by the expanded walls of the anterior portion of

the mid-esophagus (Browm, 1969). The protuberance, or

flaps {sensti Fretter and Graham, 1962) are fringed with

extremely long cilia that lieat veiy languidly.

The major Innction ol the structure is to prevent re-

gurgitation of food from the more posterior part of the

gilt during the elongation of the proboscis. It reacts

partially mechanically but also chemically — exposure to

secretions of the digestive gland or stomach contents

caused the Haps to close (Brock, 1936).

Snqrrisingly, the anatomy of the valve has not been

studied e.xtensively. In addition to the description of the

valve oi Ili/aiiassa oJ)soleta (Say, 1822) (Nassariidae) by

Browm (1969), the \alve wms described in detail only

lor Nttcclla lapilliis (Linnaeus, 1758) (Graham, 1941;

Andrews and Thorogood, 2005). Despite these veiy

limited data, the homology of the valve was never ques-

tioned. In light of the need to re-evahiate the phyloge-

netic value of antapomorphies for Neogastropoda, we

undertook a comparative study of the valve in different

branches of the Neogastropoda.

MATERIALS AND METHODS

Alaterial for this study was collected in a number of

localities; details are given in the corresponding descrip-

tions for each species. For most species, the valve to-

gether wnth parts of anterior and mid-esophagns were

dissected out Irom tlie body prior to lixation, then fixed

in 4% formalin or 75% alcohol. In the laboratory, the

valves were dehydrated and embedded in Paraplast; se-

rial sections were cut at 7 pm thickness and stained with

Masson's trichrome.

For the studies of the embiyonic development of Biic-

ciiiiiin tindafiini Linnaeus, 1758, the egg cases were col-

lected by SCUBA diving in the vicinity of the Biological

Station of Moscow State University in Kandalaksha Bay,

on the White Sea. The egg cases were maintained in the

laboratoiy in a running seawater aijuarium. Capsules

wmre dissected periodically, and embiyos preseiwed in

phosphate-buffered 2.5% glutaraldehyde (pH 7.6).

Fixed embiyos were dehydrated in graded ethanol

series and embedded in epone-araldite medium. Sec-

tions were cut at 2-2.5 pm thickness and were stained

with methylene blue and tohiidine blue in borax. Sections

were examined using a Carl Zeiss Axioplan 2 microscope

and photographed with an Axio-Cam digital camera.

RESULTS

Nassariidae

Nassaiiiis hiteostoma Broderip and Sowerby, 1829

(Figure 1)

Material Examined: Two specimens sectioned, Pana-

ma, Pacific Ocean: Venado Island, at low tide on sandy

bar, ()8°52’48.6" N, 79°35’36.9" W, coll. Yu. Kantor, 2006.

The valve of Leiblein is large, pear-shaped, about twice

as broad as the anterior esophagus, situated immediately

in front of the circumoesophageal neiwe ring. Its histolo-

gy is veiy similar to that described by Browm (1969) for

Ili/cnuissa obsoleta. The cone-shaped papilla (Figure 1,

csp) is lined by colnmnar ciliated epithelium which is

continuous with that of the anterior esophagus. The cells

on the top of the papilla bear extremely long cilia of

about 400 pm in length and that span most of the valve

length. At the base of the papilla there is a ring of tall,

ciliated, light-staining cells confluent with the papilla. In

longitudinal section, this ring of cells looks like a triangle

(Figure 1, Isc). This ring is usually called a mucous pad

(Fretter and Graham, 1962; Andrews and Thorogood,

2005), and it is thought that its main function is to pro-

duce mucus that binds the particles. This ring of cells is

seen as a whitish circle through the valve walls.

The thickened part of the valve is composed of pseu-

dostratified columnar ciliated epithelium (Figure 1,

pse). The cells are stained dark blue. No traces of the

dorsal folds of the anterior esophagus were found.

The outer surface of the valve has an extremely thin

layer of mnscle fibers (in contrast to the relatively thick-

layers of longitudinal and circular muscles that form the

wall of the anterior esophagus) and a rather thick layer

of connective tissue (Figure 1, ct).

Buccinidae

Triumjdiis distoiia (Wood, 1828)

(Figure 2)

Yu. I. Kantor and A. Fedosov, 2009

Page 75

Material Examined: Two specimens sectioned, Pana-

ma, Pacific Ocean: Playa Bicpie, in rock crevices, liigh in

intertidal zone, 08°52’42.3" N, 79°39T8.8" W, coll. Yn.

Kantor, 2006.

The valve is large, snbcylindrical, about 1.5 times as

broad as the anterior esophagus, situated at some dis-

tance in front of the neiwe ring. Histology of the valve is

rather similar to that of N. hiteosfoma, altbongb due to

the fixation conditions it seems slightly distorted. The

cilia of the cells of the cone-shaped papilla reach at

least 1100 pm in length. The ring of the light-staining

cells conihient with the papilla is less pronounced

(Figure 2, Isc). The psendostratified epithelium lining

the valve forms two different zones. The anterior zone,

rather narrow and adjoining the cone-shaped papilla is

stained veiy dark blue (Figure 2, pse) and similar in

histolog)' and staining properties to that of Nassarius

hiteostoma. This type of epithelium is sharply replaced

by light staining columnar epithelium, composed of two

t\pes of cells. The first one e.xtends from basement

membrane to the lumen (Figure 2, Ipse), bears cilia,

and has small nuclei that are located close to the apical

tip. The second t)pe of cells extends to approximately

2/3 the height of the tissue layer, and does not reach

the lumen. Their nuclei are situated in the basal 1/3 of

the cytoplasm. This ty|3e of epitlielium occupies a much

longer zone of the valve and adjoining part of the mid-

esophagus, so that in total it is three times as long as

the expanded part of the valve proper.

The outer surface of the valve has an extremely thin

layer of muscle fibers and no connective tissue.

Muricidae

Muiicanthus radix (Gmeliu 1798)

(Figures 3-5)

Material E.xaminecl: Two specimens sectioned, Pana-

ma, Pacific Ocean: Venado Island, on rocks at low tide,

08°52’48.6" N, 79°35’36.9" W, coll. Yu. Kantor, 2006.

The valve is large, pear-shaped, 3.5 times as broad as the

anterior esophagus, situated immediately in front of

tlie neiwe ring. The columnar ciliated epithelium, lining

the anterior esophagus, is shaqrly replaced by veiy tall

columna)' epithelium at the entrauce to the \ alve. These

tall epithelial cells form the large cone-shaped papilla

\\4th broad lumen. The cells on the top and external wall

of the papilla bear long cilia (Figure 5, cil), around 750

pm in length. The mucous pad at the base of papilla is

absent. The thickened part of the \ alve is composed ol

tall, columnar, folded ciliated epithelium (Figure 3, cle).

Due to the staining properties, the nuclei were not seen.

The location of torsion is seen from the exterior, lies in

the middle of the valve. The dorsal groove of the anterior

esophagus internipts the cone-shaped papilla and can be

traced along the entire x'idve length (Figure 4, clg).

The outer surface of the valve has a very thin layer of

muscle fibers (in contrast to the relatively thick layers

ol longitudinal and circular muscles that form the wall of

the anterioi' esopliagus) and no connective tissue.

Couidae, Haphitominae

Faramoniana rufozonofa (Angas, 1877)

(Figures 6-8)

Material Examined: One specimen sectioned. Western

Australia, Rottnest Island, Cape Vlamingh, intertidal

rocks, coll. ].D. Taylor, 1996.

The valve of Leiblein is veiy small, funnel-shaped,

situated immediately posterior to the buccal mass and

in front of the neiwe ring. It is about twdce as broad as

the esophagus. The wall of the valve consists of a single

layer of ciliated epithelial cells, slightly tallei' cells form

the cone-shaped papilla. These cells bear long cilia

(around 120 pm in length). No other structures can be

recognized wdtliin the valve.

Cancellariidae

Plesiotriloii vivas Ilabe and Okntani, 1981

(Figures 9-10)

Material Examined: Two specimeus sectioned, Philip-

pines, Bohol/Snlu seas, IW DA-BFAR, PANGLAO

2005 Deep-Sea Cruise, st. CP 2359, 8°49.9’ N,

123°34.9’ E, 437-476 m.

The enlargement of the esophagus (Figure 9, vl), wliich

was recognized as the \alve of Leiblein by Graham

(1966), lies immediatelv posterior to the buccal mass iu

the anterior part ol tlie extremely long, coiled proboscis,

and is partially covered by the tulmlar salix'an' glands.

The structure is coiled, formiug at least two complete

whorls, meaning that tins is not a site of torsion (where

the rotation ol the esopliagus would not exceed 180°).

Through the semi-transparent walls of the \ alve, the nar-

row strip of opaque wliite tissue ruuniug along the entire

length of the vahe is clearly seen. On external \4ew, it

looks similar to the ring of tall, ciliated light-staining cells

(= the mucous pad) in tlie valve of other neogastropods.

In histological sections, this strip is represented by

light-staining, low, non-ciliated, large epithelial cells

with large, oval nuclei. The remaining wall of the valve

is lined wth veiy tall psendostratified ciliated epitheli-

um, composed of two cells t)qres. Cells of tlie first L pe

extend from the basement membrane to the lumen, bear

cilia, and have small, narrow, elongated nuclei that are

located in the upper 1/3 of the c)4oplasm. Cells of the

second t\pe do not reach the lumen and have rounded

nuclei that are situated in the basal most part of

the cytoplasm. The cone-shaped papilla is absent. The

dorsal groove and folds were clearly seen w'ithin the

vah’e. The relatively liroad lumen of the x'ah'e was filled

with blue-staining secretion.

The outer surface of the valve has a \'en’ thin layer

of muscle fibers (in contrast to the relatively thick layers

ol longitudinal and cT'cular muscles that form the wall ol

tlie anterior esophagus) and hardly any connectix e tissue.

Olixhdae

Oliva hulhosa (Riidiug, 1798)

(Figures 11-12)

Piiy;e 76

THE NAUTILUS, Vol. 123, No. 3

Yu. I. Kantor and A. Fedosov, 2009

Page 77

Material Examined: One specimen sectioned, Aden

Bay, sandy beach 6 km west of Aden; Bed Sea, coll. D.

Ivanov.

The valve is large, at least 3.5 times as broad as

the esophagus, pear-shaped, and situated immediately

in front of the neiwe ring. The coTie-shaped papilla

(Figure 11, csp) is lined by columnar ciliated epithelium

which is continuous with that of the anterior esophagus.

The cells oti the top of the papilla hear long cilia about

200 pm in length (Figure 12, cil). Probably due to the

contraction of the papilla, its inner lumen was not oh-

seiwed. At the base of the papilla there is a ring ol tall

e.xtremely light-staining cells conllnent with the papilla

(Figurell, Isc). The thickened part of the valve is com-

posed of pseudostratified columnar ciliated epithelium

(Figure 11, pse). The cells are stained dark bine. No

traces of the dorsal folds of the anterior esophagus were

found inside the valve. The outer surface of the valve has

an extremely thin layer of muscle fibers.

DISCUSSION

Morphological Comparisons of the Valve of Leiblein

Among Different Lineages of Neogastropoda: Gra-

ham (1941) described the significant differences in tlie

foreguts oi NnceUa and Biicciwim, and suggested their

independent origins from different groups because they

e.xhibit different positions of torsion in the mid-esopha-

gus. In Nucella, torsion occurs wdthin the valve, while in

Buccinnm the position ol torsion is posterior to the

nerve ring. Ponder (1974) did not consider the position

of torsion to be of great importance, and did not dispute

the homology of the valve. At the same time he pointed

out the significant differences among taxa in the position

of the valve relative to the buccal mass. While in most of

the Neogastropoda the valve lies immediately in front of

the nerve ring, in Cancellarioidea it is situated just

behind the hnccal caUty, with the mid-esophagus posi-

tioned in front of the neive ring (Graham, 1966).

The data presented confirm the significant moipholog-

ical variability of the “valve of Leiblein” found in different

lineages of the Neoga.stropoda. The most-divergent

“valve” from the few described in literature was found in

Plesiotriton ( Cancel lariidae), in which it is coiled and

forms at least two complete whorls. The cone-shaped

papilla and the ciliaiy valve are completely absent

(Figures 9-10). The way in which it functions is unclear.

The position of the valve itself in the most-anterior part of

the proboscis is unusual for the Neogastropoda, hut it

position in relation to the hnccal mass is similar to that in

Conoidea.

In the remaining families studied, the valve of Lei-

hlein demonstrates a higher degree of similarity, being

pear-shaped and possessing the cone-shaped papilla ei-

ther formed by or lined with epithelium with veiy long

cilia, vaiying from 120 pm (Paramontana rufozonoia) to

1100 pm {Triuinphis distorta) in length. In relation to

the circumoesophageal nerve ring the valve in adults is

always positioned in front of the ring. The other charac-

ter common to all the studied species is that the walls of

the valve lack any substantial muscle layer, unlike the

walls of the adjoining part of the esophagus.

Within Conoidea presence of a valve was recorded only

in two species, Parcnnoniana nifozonata and KeiDiia

haniardi (Brazier, 1878) (Kantor and Taylor, 2002). Botli

species have a valve of veiy similar structure, which is

formed by only a single layer of cells. It should he noted

that these species are characterized by a veiy small shell

(less then 5 mm). Therefore the valve seems to he veiy

much simplified due to the minute size of the mollusks.

Some significant differences can he found among the

valves studied to date, mostly in the presence/ahsence of

the dorsal groove of the anterior esophagus within the

valve. It can he clearly ohsen'ed in nearly all families

studied — Muricidae, Cancellaiiidae, Volntidae (Ponder,

1970), Costellaiiidae (Ponder, 1972), and Voliitomitridae

(Kantor and Harasewych, 1992). It is absent in studied

Biicciuoidea, including Fasciolariidae (Marcus and

Marcus, 1962), as w'ell as in Olhidae and Conidae (our

data). Another difference among valves has to do witfi the

position of the valve in relation to the site of the torsion.

Torsion is situated posterior to the valve in all taxa except

Muricidae. We were not aide to trace the tonsion site in

Conidae due to the minute size of the animal.

Another difference ohsei'ved w^as the presence/

absence of the ring of the ciliated light-staining cells (mu-

cous pad). It WATS mentioned for eveiy stuched species

possessing the valve of Leildein, Init surprisingly it wals

absent in Muricandius (although present in Nucella). It

was similarly absent in two species of Conidae.

De\'elopment of the Valve in Ontogeny: These

differences ohsei'ved in the histology of the valve of

Leiblein prompted us to check whether its development

is identical in the emhiyogenesis of different neogastropod

lineages.

Figures 1-8. Tire valve of Leiblein. 1. Na.s.sarius luteostoina Broderip and Sowerhy, 1829, longitudinal section through the valve.

2. Triuinphis distorta (Wood, 1828), longitudinal section through the valve. 3-5. Miiricanthus radix (Gnielin, 1791). 3. Longitudinal

section through anterior esophagus and valve. 4. Enlarged fragment of the longitudinal section showing tlie dorsal groove of

the anterior esophagus interrupting the cone-shaped papilla. 5. The tip oi the cone-shaped papilla showing the long cilia.

6-8. Paramontana nifozonata (Angas, 1877). 6. Semi-diagrarnmatic longitudinal section through proboscis, buccal mass, and \'alve.

7. Enlarged semi-diagrainmatic section througli the valve. 8. Histological section through the valve. Abbreslations: aoe, anterior

esophagus; bip, buccal lips; bin, buccal mass; bsc, buccal sac; bt, buccal tube; cil, cilia; cle, columnar folded ciliated epithelium;

con, circumoesophageal newe ring; csp, cone-shaped papilla; ct, connective tissue; dg, dorsal groove; Ipse, light staining columnar

epithelium; Isc, light-staining cells; pse, pseudostratified epithelium; sd, salivaiy duct; sg, salivai'}' gland; sp, septum of the

rhynchocoel; vg, venom gland.

Page 7(S

THE NAUTILUS, Vol. 123, No. 3

Figures 9-12. \'alve of Leihlein. 9-10. Flcsiofritoii viviis llahe et Okutaiii, 1981. 9. Anterior part of the proboscis dissected to

show the position of the \ alve in rel;ition to the Iniccal mass. 10. Longitudinal section through the anterior esophagus and the coils ol

the valve. 11-12. Oliva hiilhosa (Rdding, 1798). 11. Longitudinal section through anterior esophagus and valve. 12. Longitudinal

section through x’ah'e and mid-esophagus, showing the long cilia ol the cone-shaped papilhi. Abbreviations: p, proboscis; vL, \ al\e of

Leiblein. Other abbre\iations see in captions to Figures 1-8.

There is veiy little published data on the development

ol the vaK e of Leiblein in ontogeny. Ball et al. (1997a, h)

examined the ontogeny of the foregnt in Niicella lapiUus

(Figures 13-15).

Ahro (1969) (snrmnari/.ed by Fretter, 1969) examined

the emhiyology ol Nassariiis incrassatus (Strdm, 1768)

and N. rcticiilahis (Linnaens, 1758) (Nassariidae). Page

(2005) n'-exarnined the development ol the (oregnt and

proboscis in a dillerent nassariid species, Nassarius

incitdicii.s (Ccjnld, 1850) with planktotrophic lamie and

illustrated it by a sc-ries ol outstanding photographs.

We complemented the data on tlie Nassariidae by ob-

sen'ations ol direct developing embrvos ol Biicciinim

midfil II III .

Published and original data on the embiyonic devel-

opment ol the valve in Bnccinidae and AInricidae

demonstrated that it originates Irom dillerent sections

ol the anterior foregut. From the diagrams of Ball et al.

(1997a, b), it is obvious that in Niicella, the buccal mass

with the radnla originated from the ventral ontpocketing

of the esophagus during the early stages ol proboscis

fonnation (Figure 13). The valve of Leiblein appeared

in the next stage (Figure 14), as development of the

esophagus posterior to the uen'e ring. Later, the pro-

gressive elongation ol the proboscis pulls the salivary

glands, radnlar sac, and the valve through the neiwe ring

into their final positions (Figure 15). Thus, the valve is

formeil as part ol the anterior larval esophagus.

Yu. I. Kantor and A. Fedosov, 2009

Page 79

Figures 13-19. I2iagraminatic lateral view of tlie development ol the foregut and proboscis. 13-L5. Nucclla l(ii)illiis (after Ball

et ah, 1997a, h, modified). 13. Stage fi, the buccal muss has developed. 14. Stage 7, the valve ol Leiblein lies posterior to the cerebral

commissure. 1.5. Stage S, the saiKe of Leiblein, acinous salivaiT glands, and the radular sac lie anterior to the neiwe ring. 16-19.

Selected stages of the foregut development in Buccinoidea (based on Page, 2005, on Nassariiis mcndiais (Gonld, 1S50) and

obseiwations on Biiccinmn umhihnn). Sali\an' glands are omitted for simplicity 16. Foiination of tlie ventral ontpocketing.

17. Formation of buccal caxity, anterior esophagus, valve of Leiblein, ami radular sac. Lar\al esophagus still open. 18. Lanai

esophagus resorbed, lamil mouth opening is sealed. 19. Postinetamorphic new rnontli is formed. Abbreviations: ascl, duct of

accesson' salixaiy gland; a.sg, accesson- salivan' gland; bhc, buccal commissure; be, buccal cavity; Ini, lar\al mouth; loe, lar\al

esophagus; ni, mouth; out. ontpocketing; ps, proboscis sheatli; sn, snout. Other abbresiations as in captions to Figures 1-12.

The situation with Nassariidae and Biicciiiiini differs

significantly. In these groups, the lanail esophagus is

initially a ciliatetl tube that extends from the inoutli to

the stomaeli. A patch of enlarged, non-ciliated cells is

embedded within the \'entral wall of the tlistal lanal

esophagus and forms an out[:)ocketing (Figure 16, 20-21).

Eventually, the f)ut|X)cketiug will enlarge and become ex-

tensively elaborated to form the entire post-metainoiphic

foregnt. At first, the lutnre hnccal ca\dtX', radnlar sac, and

valve of Leiblein form Iroru different chambers of the

onginal ontpocketing (bdgures 17, 22-23). With the en-

largement of these structures as w^ell as elongation ol the

proboscis, dramatic changes take place. Most notable

among these events are the complete occlusion of the

lanal month and the degenerahon and loss of the distal

lan al esophagns between the lan al mouth and the point

Page 80

THE NAUTILUS, Vol. 123, No. 3

100 gm

out

100 gni 20

4..6'

100j,ira

1 00 gm

w0ii

i0j?k»M4

Figures 20-23. Selectetl stages of development of Bucciiunn iiiidatitiii. 20. Mid-sagittal section through a lana. \'entral

ontpocketing formed. The stage corresponds to Figure 16. 21. Enlarged region ol Figure 20. 22-23. Formation ol hnccal cavit)',

anterior esophagus, \'alve of Leihlein, and radnlar sac. Lan ai esophagus still open. The stage corresponding to Figure 17. 23. En-

larged regiont ol Figure 22. Ahhrex iations: ne, nurse eggs; ocl, odontophore. Otlier abbreviations see in captions to Figures 1-19.

where the post-inetanioiphic foregut extends from the

ventral sirle of the lanal esopliagirs (Figure 18).

Later, tlie new delinitive month niptnres through the

transient epithelial seal tliat formed over the laival month

(Figure 19) (Page, 2005, figs. 2 B, C). Tims the overall

similaritv ol adult loregnts ol Bnceinoidea and Mnrieidae

in fact is achieved ihrongh veiy different prt)cesses.

W'e want to emphasize that the radnlar apparatus in

both stems originates Irom homologous strnetnre.s — the

ventral ont]X)cketings ol the esophagus. In contrast, the

“valvc’" originated Irom dilferent parts ol the loregnt —

Irom the posterior chamher ol ventral ontpocketing in

Buccinidae and part ol the anterior lanal esophagus

in Vlnricidae.

Our attempts to examine the entire development of

the foregut in emlm’os oi Buccinin)} inidatuui failed due

to asvmchronons development ol the embiwos even wdth-

in the same egg cluster and egg capsules. Therefore it

was not possible to obtain the embivos t)u c(m.sec|nent

developmental stages wdth any reliable timing. Never-

theless, we were able to obsene the early stages which

roughly corresponded to approximate hallway point ol

obligator lanml development (21 days post-hatcliing) in

Ndssaritis meudicus (Page, 2005).

The major difference between stndietl na.ssariids and

Biicciinmi is that the na.ssariids are characterized by feed-

ing planktonic larvae, while Biiccinmn has direct develop-

ment, feeding on nimse eggs inside the egg capsule and

Yu. I. Kantor aud A. Fedosov, 2009

Page 81

hatching in the crawling stage. The nurse eggs are con-

sumed in rather early stages, and are clearly seen in tlie

lamil esophagus (Figures 20-2, ne). We have not ob-

served the stage with the degenerated laiwal esopliagus,

hut have seen an example of the strongly differentiated

initial ontpocketing gi\ing rise to the radnlar sac, in which

the radnlar teeth were seen, and the buccal mass with

odontophore and future anterior esophagus with valve of

Leiblein was situated in e.xactly tlie same positicjn as in N.

mcndicus (Figures 22-23). Therefore, it is presumed tliat

the development of the valve in Buccium undutum is

analogous to that in Nassarhis.

It should be emphasized that the development of

tl ie valve seems to be unrelated to the mode of embiyo-

genesis. Similar developmental patterns were (bund

in related species with planktotrophic {Nassariiis) and

lecitotrophic lamie (Bitccinitm), while unrelated species

with lecitotrophic laiwae {Buccinum and NuceUa) dif-

fered in the development of the valve. Both Biiccimnn

and NuceUa feed on the nurse eggs during the first

stages of the development (Fretter and Graham, 1962).

Our preliminaiy data demonstrated significant dilfer-

ences in the morphology of the valve of Leiblein in

different groupings of Neogastropoda, and different ori-

gins of the valve during embiyogenesis, at least in Mnr-

icidae and Bnccinidae. This suggests that, despite the

superficial similarity, the homology of the valve of Lei-

blein ndthin Neogastropoda is at best questionable.

If this supposition is correct, then Bnccinoidea do not

share any of the pre\aonsly h\qiothesized antapomor-

phies with the rest of neogastropods. This raises the

prospect of a paraphyletic Neogastropoda that includes

two stems, one including the Bnccinoidea, the other

containing the remaining neogastropod families.

AC KNOWLE DC M E NTS

The senior author was to e.xpress his thanks to Drs. M.G.

Harasexxych and E.E. Strong of the National Museum of

Natural Histoiy, Smithsonian Institution, who organized

and invited him to the Workshop in Panama (2006),

during which the bulk of material was collected. Prof

fohn Taylor kindly provided sections of Parauioiitaiia

rufozonata and Kcrmia barnardi. We would like to thank

the staff of the Chair of Invertebrate Zoology of Moscow

State University lor providng access to their histological

equipment. An anonymous referee and Dr. Winston

Ponder added valuable comments and corrections to the

manuscript. The work was in part supported by the grant

of Russian Foundation of Basic Research 09-()4-()()911.

LITERATURE CITED

Abro, A.M. 1969. Studies in the functional anatomy of some

British monotocardian veligers, witli observations on the

musculature of tlie buccal mass of Littorina littorea (L.).

Ph.D. dissertation, Univ'ersity of Reading, Berksliire, UK.

Amaudrut, A. 1898. La partie anterieure de tube digestif et le

torsion chez les mo!lus(|ues gasteropodes. Annales des

Sciences natnrelles, 8'' series, Zoologie 7: 1-291, pis. 1-10.

Andrews, E.B. and K.E. Tliorogood. 200.5. An ultrastructurai

study of the gland of Leiblein of iiuuicid and nassariid

Neogastropods in relation to function, with a discussion

oil its homologies in other Caenogastropods. Journal ol

Molluscan Studies 71: 269-.300.

Ball, A.D., E.B. Andrews, aud |.D. Taylor. 1997a. The ontogeny

of the pleurembolic proboscis: NuceUa laijilliis (Gastro-

poda: Muricoidea). |ournal of Molluscan Stmlies 6.3: 87-95.

Ball, A.D., J.D. Taylor, aud E.B, Andrews. 1997b, Develop-

ment of the acinous and accessoiv salivaiy glands in

Nucclla lapilhts (Gastropoda: Muricoidea). Journal of

Molluscan Studies 63: 24.5-260.

Brock, F, 19.36, Suche, Aiifnahme und enzymatische Spaltuug

der Nahrung durch die Wellhoruscfmecke Bncciiimn

undatum L. Zoologica, Stuttgart 34(92): 1-1.36 (not seen).

Brown, S.G. 1969. The structure and function of the digestiv'e

system of the mud snail Nassariiis ohsolctiis (Say). Mala-

cologia 9: 447-500.

Colgan, D.J., W. E. Ponder, and P. E. Eggler, 2000. Gastropod

evoliitionar)' rates and phylogenetic relationsfiips assessed

using partial 28S rDNA and histone 113 sequences. Zool-

ogica Scripta 29: 29-63.

Colgan, D.J., W. E. Ponder, E. Beacham, and J.M, Macaranas.

2003. Molecular phylogenetic studies of Gastropoda

based on six gene segments representing coding or non-

coding and mitochondrial or nuclear DNA. Afolluscan

Research 23: 123-148.

Colgan, D.J., W.E. Pomler, E. Beacham, and |.M. Macaranas.

2007, Molecular phylogenetics of Caenogastropoda (Gas-

tropoda: Mollusca). Molecular Phylogenetics and Evolu-

tion 42: 717-7.37.

Fretter, V. 1969. Aspects of metamoqrhosis in prosobranch

gastropods. Proceedings of the Malacological Society of

Loudon, .38: 37.5-385.

Fretter, V. and A. Graham. 1962. British Prosobranch Mol-

luscs. Ray Society', Loudon, xM + 755 pp.

Golikov, A.N. and Ya.l. Starobogatov'. 1988. Problems of pliv-

logeny and system of the prosobranclhate gastropods.

Proceedings of the Zoological Institute, Leningrad 187:

4-77 [In Russian],

Graham, A. 1941. The oesophagus of the stenoglossan proso-

branchs. Proceedings of the Royal Society of Edinburgh,

series B, 61: 1-23. "

Graham, A. 1966, The fore-gut of some marginellid and can-

cellariid prosohranchs. Studies in Tropical Oceanography',

Miami, 4: 1.34-151.

Ilarasewych, M.G., S.L. Adamkewicz, J.A. Blake, D. Saiidek,

T. Spriggs, and C.J. Bull. 1997. Neogastropod phylogeny:

A molecular perspective. Journal ol Molluscan Studies 63:

327-351.

Kantor, Yu. 1. 2002. Moiphological prerequisites for under-

standing Neogastropod phy'logeny. In: M, Oliverio and

R. Chemello (eds). Systematics, Phvlogeuy and Biologv'

of the Neogastropoda. Bollettino Malacologico, Supple-

ment 5: 2-15,

Kautor, Yii.l. and M.G. Ilarasewych. 1992a. Morphology of

the digestive system of Volutoiiiifra alaskaiia flail, 1902

(Gastropoda, Pectinibranchia, N'olutomitridae), with notes

on the possible mechanism of feeding. Ruthenica, Rus-

sian Malacological Journal 2(1): 27-35.

Kantor, Yii.l. and J.D. Taylor. 2002. Foregut anatmny aud

relationships of raphitomine gastropods (Gastropoda:

Page 82

THE NAUTILUS, Vol. 123, No. 3

Conoidea: Rapliitominae). In: M. Oliverio and R. Che-

niello (eds). Sy,steinatics, Phylogeny and Biology of the

Neogastropoda. Bollettino Malacologico, Supplement 5:

161-174.

Marcus, E. and E. Marcus. 1962. On Lcucozonia nassa. Bole-

tim da Facultade de Eilosofia, Ciencias e Letras, Univer-

sidade de Sao Paulo, 261, Zoolog)' 24: 11-.30.

McArthur, A.G. and M.G. Marasewych. 2003. Molecular sys-

tematics of the major lineages of the Gastropoda. In:

C. Lydeard and D.R. lindherg (eds.). Alolecnlar System-

atics and Phylogeography of Mollusks. Smithsonian

Books, Washington, pp. 140-160.

Page, L. R. 2005. Development of foregut and proboscis in the

hnccinid neogastropod Nassariiis mcndicus: evolntionaiy

opportunit)' exploited by a developmental module. Jour-

nal of Morphology 264: 327-338.

Ponder, \TF. 1970. The morphology oi' Aid tlu>e arahica (Mol-

Insca: Mrlutidae). Malacological Review 3: 127-165.

Ponder, \V. F. 1972. The morpholog)' of some mitriform gastro-

pods with special I'eference to their alimentaiy^ and repro-

ductive systems (Neoga.stropoda). Midacologia 11: 295-342.

Ponder, \V. F. 1974. Origin and evolution of the Neogastro-

poda. Malacologia 12: 295-338.

Ponder, W'.F, D.J. Colgan, J.M. Healy, A. Niitzel, L.R.L.

Simone, and E.E. Strong. 2008. Caenogastropoda. In: W.

F. Ponder and D.R. Lindherg (eds.). Phylogeny and Evo-

lution of the Mollusca. University of California Press,

Berkeley: 331-383.

Ponder, W. F. and D.R. Lindherg. 1997. Towards a phvlogeny

of gastropod molluscs — an analysis using morphological

characters. Zoological Journal of the Linnean Society 119:

8.3- 265.

Riedel, F. 1994. Recognition of the snperfamily Ficoidea Meek

1864 and definition of the Thalassocynidae fain. nov. (Gas-

tropoda). Zoologische Jahrbnchei; Ahteilung fiir Systematik,

Geographic und Biologic derTiere 121: 457—474.

Riedel, F. 2000. Ursprnng und Evolution der “hoheren" Cae-

nogastropoda. Berliner Geowassenschaftliche Ahhandlun-

gen E 32: 1-240, 21 Tafeln.

Sheridan, R., J.J. Van Mol, and J. Bouillon. 1973. Etude

mophologique du tube digestif de quelques Turridae

( Mollusca-Gastropoda-Prosobrancl lia-Toxoglossa) de la

region de Roscoff Cahiers de Biologie Marine 14: 159-188.

Shimek, R.L. and A.J. Kohn. 1981. Functional moqrhology

and evolution of the to.xoglossan radula. Malacologia 20:

42.3- 438.

Strong, E.E. 2003. Refining molluscan characters: moiphol-

og)', character coding and a phvlogeny of the Caenogas-

tropoda. Zoological Journal of the Linnean Society 137:

447-554.

Taylor, J.D. and N.J. Morris. 1988. Relationships of Neogas-

tropoda. Malacological Review, Supplement 4: 167-179.

THE NAUTILUS 123(3):83-94, 2009

Pa^e 83

Phylogenetic analysis of the subfamily Colinae (Neogastropoda:

Bnceinidae) based on morphological characters

Alisa R. Kosyan

Yuri I. Kantor

A.N. Severtsov Institute ot Ecology and Evolution

Russian Academy ot Sciences

33 Leninski Prospekt

Moscow 119071, RUSSIA

kosalisa@rai nliler.ni

ABSTRACT

Colinae Gray, 1857, the most abundant and diverse sulilamily

ol whelks in the northwestern Pacific and Far-Eastern Seas ot

Russia, includes several conchologically similar genera or suh-

genera ot unclear status and composition. Based on morphol-

ogical and anatomical studies ol 38 species attributed to the

genera Coins Rciding, 1799, Phcijnsns Dali, 1902, Latisipho

Dali, 1916, Ankicofiisns Dali, 1918, Retifusns Dali, 1916, Rcfi-

inohnia McLean, 1995, and Pararctifnsus Kosnge, 1967, a par-

tial generic re\'ision and phylogenetic analysis based on 34

characters is produced. Tlie resulting majorits' rule consensus

tree well resolves the genera Plicifisus, Retifnsns, Panirctifn-

sii.s. and Anhicofusns. The genus Retimolinia appears to be a

junior synonym ot the genus Rctifnsus. Species ot the hetero-

geneous genus Coins included in this study do not form a

clade, indicating that this genus, as presently understood, is

paraphyletic. Our results demonstrate the importance and util-

ity of anatomical characters for resolving the systematics of the

e.xtremely div'erse and v’ariable family Bnceinidae.

Additional kci/icords: Taxonomy, phylogeny, cladistics, nortli-

westeni Pacific

INTRODUCTION

Althoiigli the miniher ol papers dedicated to the molecu-

lar phylogeny of neogastropods continues to increase,

there is no parallel increase in data on their morphology

and anatomy. Tins is especially tme tor tlie Bnceinidae, a

large and evolntionarily successtnl family ot predatoiy ma-

rine gastropods that are widespread in polar, temperate,

and tropical waters ol the World Ocean, and which have

significant commercial value. In the northwesteni Pacific,

Bnceinidae is one ot the dominant tamilies, and in waters

ot the Russian Far-East, it is the most abundant and di-

verse tamily, comprising more tlian 30% of the total num-

ber of gastropod species (Kantor and Sysoev, 2006). Six

buccinid subfamilies are present in the northwestern Pa-

citic: Bnccininae Rafinesqne, 1815; Colinae Gray, 1857;

Beringiinae Golikov and Starobogatov, 1975; Ancistrolepi-

tlinae Ilabe and Sato, 1973; Parancistrolepidinae Ilabe,

1972; and Volntopsiinae Ilabe and Sato, 1973. The snb-

lamily Colinae (previously better knowai under the name

Neptuneinae Stimpson, 1865) is the most diverse with

respect to the number ot genera and species in the north-

western Pacific (Kantor and Sysoev, 2005, 2006). It

includes 16 ot the 34 genera and 116 ol the 263 species ol

Bnceinidae recorded in the fauna ot Russia.

The best known representative of this subfamily is the

diverse genus NeptnnecK which has had two recent revi-

sions (Golikov, 1963; Fraussen and Ternn, 2007). Other

genera, with species that do not grow to commercial

size, have not attracted sidlicient attention ot malacolo-

gists. Among tliem are sev'eral conchologically similar

genera with unclear taxonomic status and species com-

position, inclnding: Coins ROding, 1799, Latisipho Dali,

1916, Plicifusus Dali, 1902, Aniacofnsns Dali, 1918, Rcti-

fnsus Dali, 1916, Pamretifusus Kosnge, 1967, and Reti-

mohnia McLean, 1995.

Species and genera within Bnceinidae luwe generally

been diagnosed based primarily on conchological char-

acters, with radnlar moqvhologv contributing only occa-

sionally to their taxonomy. Anatomical characters have,

thus tar, hardly been used tor these purposes.

The aim ot this publication is to clarify the status and

composition of the genera Coins Rikling, 1799, Plicifn-

sns Dali, 1902, Latisipho Dali, 1916, Aniacofnsns Dali,

1918, Rctifnsns P)all, 1916, Rctiinohnia McLean, 1995,

and Pararctifnsus Kosnge, 1967, based on conchological,

anatomical and radnlar characters, as w'ell as to evaluate

the utilitv' ot moiphological characters tor resoKing the

ta.xonomy ot Colinae.

MATERIALS AN14 METHODS

We dissected and analyzed the anatomy ot 38 species

ot Colinae, defining 34 cliaracters coded as 82 chaixicter

states that w'ere used to perform the phylogenetic

analyses of these taxa (Table 1, Appendix 1). Of these,

7 characters described shell structure, 5 characters the

Page 84

THE NAUTILUS, Vol. 123, No. 3

soft body and the mantle, 5 characters the reproductive

systems, 12 characters the digestive system, and 5 char-

acters tlie strnctnre of the radnla. The material for the

study was ohtmned Irom the Zoological Institute (Saint

Petersburg, Russia), the P, P, Shirshov Institute of Ocean-

olog)^ of Russian Academy of Sciences ( AIoscow), and the

Zoological Mnsemn of Moscow State University. In totd,

nearly 200 specimens were dissected. While processing

this material, standard zoological methods were used,

such as manual dissection, histology and scanning elec-

tron microscopy for the e.xamination of radnlae. Phyloge-

netic analyses were rnn using Panp*4 (Swofford, 1998).

RESULTS

Brief DESCRiimiONS of the Taxonomically Infor-

MATi\E Morphological Characters of the Studied

Genera: The gross anatomy of Colinae is Apical of the

Bnccinidae in general features (Figures 1-2). The opercu-

lum may have a terminal {LatisipJio, Cohi.s, Aiilacofusus;

Figures 1, 26), or suhspiral nucleus [Pararetifiisiis; Fig-

ure 4). The mantle ca\’itv' spans approximately one whorl

of the body (Figure 3). The ctenidinm (ct), osphradinm

(os) and, in females, the capsule gland (eg) can be ob-

sen ed by partial transparency ol the mantle. Relative sizes

of the ctenidinm and osphradinm vaiv in different species.

Penis moiphology was used successfully by Golikov

(1963, 1980) for taxonomic studies of the genera Neptii-

nea and Bticchuim- however, in our study, the structure

of the distal section of penis varied ven' little. In Latisi-

j)Jn\ Plicifusns, Rvo species of Coins, and several Rctifn-

sus species, the seminal duct opens at the tip of a large,

cone-shaped papilla (Figures 6, 8-9, sp) that is encircled

by a fold of skin (cf). In the remaining Retifusns species,

the seminal papilla is veiy small and becomes narrower

towards its tip (Figure 10, sp). In the genus Para-

retifnsns, the seminal papilla is absent, \\4th the male

orifice situated terminally at the tapering tip of the penis

(Figure 5). The strnctnre of the pallia! gonoduct in

females appeared to be even more conser\/ative in the

genera studied, In the majorirt' of species we e.xamined,

the vagina is strongly developed (Figure 11), occupving

a ventral position on the capsule gland. Only in the

genus Pararctifusns it is situated terminally.

The montli opening is situated at the tip of a more or

less elongated proboscis (Figure 12, mo). While con-

tracted, the proboscis is situated within the rhynchodenm

(Figiire 12, rd). The anterior section of the rhynchodenm

is immovable and attached to the body haemocoel walls by

multiple tensor muscles. The posterior seebon of the

rhvnchodeuni is capable of being everted. The proboscis

is retracted by retractor muscles attached to the rhxmcho-

denm walls (Figure 12-15, prr). The longest proboscises

in the contracted state were found in Anlacofnsns and in

some species ol Coins, where they are folded vithin the

rhynchodenm (Figure 14). In other genera (Plicifusns,

Rctifnsns, ami Latisipho), the jiroboscis remains straight

within the rhvnchodeuni (Figures 12-13), and elongates

mostly due to eversion of the po.sterior, movable seebon.

The proboscis wall is foi'med of an epithelium, one or

A\'0 layers of circular muscle fibers, and Rvo layers of

longitudinal muscle fibers. The sequence of layers in the

majority of studied genera (Anlacofnsns, Latisipho, Reti-

fnsns) is, (from outer to inner surfaces): epithelium, cir-

cular muscle layer, longitudinal muscle layer, circular

muscle layer, and an innermost longitudinal muscle layer

(Figures 18-20). In Rvo studied species oi Plicifusns, the

sequence of layers differed, consisting of: epithelium,

longitudinal muscle layer, circular muscle layer, and lon-

gitudinal muscle layer in P. hastarins (Figure 24), with

the addition of an innermost, second layer of circular

muscle fibers in P. rlu/ssns.

Within the proboscis is the buccal mass with radula.

Gomparative lengths of the buccal mass varied among

taxa and have taxonomic significance. Each row of the

radula (Figures 36-41) consists of Rvo lateral teeth and

one central tooth, each normally bearing 3 cusps. Al-

though the teeth are similar in shape, the finer details

are specific for genera (see below in the discussion).

The anterior esophagus opens into a large (Rctifn-

sns, Pararctifusns) or medium-sized (Latisipho, Plicifn-

sns, Coins, Anlacofnsns) valve of Leiblein (Figure 12,

vl). The gland of Leiblein is present in all studied

species (Figures 12-13, gl). Salivaiy glands differ in

shape and in size (Figures 12-14, sg), being largest in

Rctifnsns and Anlacofnsns. The salivaiy ducts leave the

inner side of each gland and run along the esophagus

to their openings into the posterior part of the buccal

ca\4ty. The diameter and the structure ol the wall of

the ducts vary among different genera. In Latisipho,

Plicifusns, and Coins, the ducts are thin and coiled

(Figure 13, scl), while in Anlacofnsns, Rctifnsns, and

Pararctifusns, they are thick, sometimes with swellings

in a form of a sac (salivaiy sacs) (Figure 15, ss). In

Anlacofnsns, the walls of salivaiy ducts have an addi-

tional layer of longitudinal mucles (Figure 21, Ini).

The posterior esophagus opens into the stomach. The

structure of the stomach is generally of the same t)qre

in the majorit)’ of the species studied, but the length

of the posterior mi.xing area can differ among genera

(Figure 16-17, pma).

Phylogenetic: Analyses: Volntopsins norvegiens (Gme-

lin, 1791) (Bnccinidae: Volutopsiinae) And Ancistrolcp is

okliotcnsis Dali, 1925 (Bnccinidae: Ancistrolepidinae),

whose anatomy is knowm (Kantor, 1982, 1988), were

used as outgroups. A heuristic search )4elded 2624 trees,

each 147 steps in length. Gonsistency index (Gl) =

0.3197, homoplasy index (HI) = 0.6803, retention index

(Rl) = 0.6942. Figure 25 shows the 50% majority-rule

consensus tree.

Several clades can be distinguished within the

ingroup (Glades 1 to 6, Figure 25).

Glade 1, which is supported in 93 percent of trees,

corresponds to the genus Plicifusns, and contains 12

species, including the Rqre species o\ Plicifusns. At the

moment, we prefer to treat it as a monophyletic genus

pending examinations of additional species.

A. R. Kosyan ami Yu. I. Kantor, 2009

Paue 85

Figures 1-11. Anatomy. 1-2. Plicifn.siis hamhiisiis. .3. Mantle ol Flicifiisiis Ijastariii.s. 4. Opercniiini ol rararrUfiisus kantori.

5. Penis ol Pararetifiisus kanfoii. 6, 8. Penis ol Lalisipho liallii, ventral \ie\v. 7. Frontal-dorsal \iew ol the soft hotly ol Coins uiinor.

with mantle removed. 9. Upper section of penis ol Coins minor. 10. Penis ol Rctifnsns jcssocnsis. 1 1. Pallial female reprodncti\'e

system ol Plicifnsns rhi/.ssns, capsule glaiul openetl dorsally. Ahhre\ iations: be, hm'.sa copniatrix; cf, circular fold of skin around tlie

seminal papilla, eg, capsule gland; ciul, coinmellar muscle; ct, ctenidimn; clg, digestix'e gland; eye, eye; f'o, Icnnale orifice; lul, heath

lig, hxpohranchial gland; kd, kidney; ni, mantle etlge; op, opercnhini; os, ttsphradinm; p, penis; prp, propotlinm; prpg, propotlial

grt)ove; re, rectum; s, siphon; so, male orifice; sp, seminal papilla; va, \agina

Plicifnsns Dali, 1902

Tritonofusns {Plicifnsns) Idall, 1902: 523.

Type Species: Fnsns krot/eri Moller, 1842, by original

designation.

Diagnosis: The genus is eharaeterizetl hv an elongat-

ed, small to medium-sized fusiform shell with well-

developed axial ribs and nnmerons spiral cortls (from

30 to 60 cords on penultimate whorl) that cttver the

entire sliell surface (Figures 31, 33). The central tot)th

of the radnla is large and brttad, and has hvo to four

Page S6

THE NAUTILUS, Vol. 123, No. 3

A. R. Kosyan and Yu. I. Kantor, 2009

l^age 87

(usually tliree) sharp cusps (Figure 36), The lateral teeth

usually have three or four cusps, with the central cusps

always smaller than the lateral ones. The salix aiy ducts

are veiy thin and convoluted. The stomach is large, as

compai'ed to the proboscis, and narrow, with a small

posterior mixing area.

Remarks: Phcifu.siis was described by Idall (1902) as

subgemis of Tritonofiisiis Mdrch, 1857, which is an ob-

jective synonym oi' Colas Roding, 1799, since it is based

on the same t\q:>e species. Plicifasas has been treated as

a distinct genus by the majorih' of subsequent authors.

Genus Composition: The majoritv' of the included

species were described wdthin this genus [or attributed

to the subgenus Tritoaofasus {Plicifasas)]. Qaasisipho

tonpiaias Petros; 1982, is the tvqre species of the mono-

tvpic genus Qaasisipho Petrov, 1982, from the upper

Pliocene-lower Pleistocene of eastei'n Kamchatka. This

species smwives in the Recent fauna, and its anatomy

confirms that the tspe species belongs within Plicifasas.

Thus Qaasisipho becomes junior subjective svnonvm of

Plicifasas. Some species were originally described or

attributed to Retifasas [e.g., Plicifasas (Rctifastis) scis-

saratas Dali, 1918]. Tritoaofasas (Plicifasas) rhijssas

Dali, 1907 was placed in the genus Helicofasas Dali,

1916 (tvpe species by original designation Trito)iofasas

(Plicifasas) aaraatias car. laticordafas Dali, 1907) by

many Russian authors (e.g., Kautor and Sysoev, 2005,

2006).

The results of our study place the following species

wathin the genus Plicifisas:

Plicifasas kweijcri (Moller, 1842) [= Fasas arcticas

Philippi, 1850]

Plicifasas plicatas (A. Adams, 1863)

Plicifasas scissarafas (Dali, 1918)

Plicifasas croceas (Dali, 1907)

Plicifasas ekieodes (Dali, 1907)

Plicifasas rlu/ssas (Dali, 1907) [= Plicifasas (Latifasas)

wakasaaas Dali, 1918; Tritoiuifasas (Plicifasas) aar-

aatias Dali, 1907; Pliciftisas (Aaiacofasas) ria/ssoides

Dali, 1918]

Plicifasas hastarias Tiba, 1980

Plicifasas haadjasas Tiba, 1980

Plicifasas ohtasatas Goliko\’ ia Golikov and Scarlato,

1985

Plicifasas olivaccas (Aurivillius, 1885) [= Plicifasas

(Retifasas) iacisas Dali, 1919]

Plicifasas oceanodroaiae (Dali, 1919)

Plicifasas tonjaatas (Petrov, 1982)

A second, well delined clade witli 100% bootstrap snp-

poi't includes 20 .species in our study, and is composed of

several well supported subclades (chides 2, 3, 4, 5) and

two unresoK'ed species.

Glade 2, although not supported in all trees, contains

three northern Atlantic species of the genus Colas Riid-

ing, 1798 (Figure 26), including G. i.slaadicas, the t\pe

species. The other two species, often attributed to Colas:

C. miiav (lOall, 1925) and C. kajiaiais Tiba, 1973, do not

emerge as members of this clade. Tliese results reflect

the high heterogeneity of Colas, which is widely

distributed iu the Atlantic and Arctic Oceans and in the

northern Pacific. Alauy more species need to be studied

in detail before the taxonomy ol C(das is clearly under-

stood.

Glade 3 includes three species belonging to the genus

Pararctifasas, iucluding its Rpe species.

Pararctifasas Kosuge, 1967

Rrt/fa.va.v (Pararctifasas) Kosuge, 1967: 62.

T>q)e Species: "Pla/aiorla/achas?” teiuiis Okutani,

1966 (by original designation).

Diagno.sis: The genus is cliaracterized by a small shell

with a relati\’ely liigh last whorl. The spiral sculpture

consists of a few ele\'ated, sharp or rounded ribs; axial

folds are absent (Figures 28, 30). The radiila is similar

to that of Retifasas roseas, R. Iatici)igalatas. R. siaiilis,

R. itaraj)iis, and Pi. attoiaatas (Figure 38) (see below for

description).

Remarks: The t\pe species was originally placed iu

Plapaorlapichas (Gouoidea), but examination of the rad-

ulai' and moqihological characters undoubtedly placed it

wathiu Buccinidae (Kosuge, 1967).

Genus Composition: \Tn' few species have been

placed in Pararctifasas. In addition to the species stud-

ied here (below) only one, P. dcdonderi Franssen and

Hadorn, 2001, from Pliilippines was tentatively attribu-

ted to Pararctifasas but later excluded by Kosyau

(2006a).

Pararctifasas teaais (Okutani, 1966)

Pararctifasas hnitori Ko.syan, 2006

Pararctifasas kosaaci Kosyan, 2006

Tlie genus was proposed as a subgenus of Retifasas

and is close to it in radiilar strnctnre and anatomy, but

differs in shell sculpture. The spiral cords ol Pararctifa-

sas shells are veiy similar to the cords of Aaiacofasas

Figures 12-17. Anatomy. 12. Right lateral \'iew of the foregut of Plicifasas hastarias. l.'l. Left lateral \iew of the foregut ol

Plicifasas rhijssas. 14. Right lateral view of the loregut of Aaiacofasas hcrcadceai. 15. Dorsally opened prolioseis of Retifasas

roseas. 16. Opened .stomach o( Aaiacofasas pcriscclidas. 17. Opened stomaeh of Plicifasas hastarias. Ahhre\iations: aclg, opening

of anterior duct of digestive gland; agl, ampulla of gland of Leibleim aoc, anterior esophagus: bin, buccal mass; gl, gland of

Leibleim int, intestine; mo, mouth opening; n, neimes; nr, nen'e ring; oclr, odoiitophore retractors; oeo, oesophageal opening; pclg,

opening of posterior duct of digestive gland; pnia, posterior mixing area; poe, posterior esophagus; pr, profioscis; prr, proboscis

retractors; r, radula; rcl, rlpmcliodeuiu; scl, salivaiy duct; sg, salix an' gland; tfl, tvphlosole; vl, valve ol Leihlein

88

THE NAUTILUS, Vol. 123, No. 3

lOOjim

50 (jm

A. K. Kosyan and Yu. I. Kantor, 2009

Page 89

• \ olufopsius

• P. Uroeycri

• P pticatus

• P. croceus

• P oceunodronute

• P. hamhusus

mP otivaceiis

• P (ortimifus

mP rhyssus

• P. scissnnifiis

• P ehieodes

• P hiistarius

• P. ohtusafiis

• C. isltuuUciis

• C. ^nicHis

• C. jeffreysUintis

• C. minor

• Par. tenuis

• Par. kantori

• Par. kosu^ei

• R. virens

• R. jessoensis

• R. yanainii

• R. frieiei

• R. faticingulatus

• R. simi/is

• R. a/fcnuutus

• R. rosetts

• R. ifurupus

> 4. hrevicauda

4. hcrendeeni

4. omhronius

f. periscelidns

• C. kujianus

• L. haUii

• L. hypoUspus

• N. antiqna

• N. ja^'udinae

• y <>uihini

4 ncistrolepis

Figure 25. Fifty-percent inajorip'-mle consensus tree ob-

tained from 2624 trees, each 147 steps in length.

perisceUdiis; however, the anatomy of Pararetifiisus dif-

fers considerably.

Clade 4, which is conchologically most heterogenous,

contains 9 species previously classified within the genera

Rctifusus, Mohnio, Refimohiiia, and Plicifusns. The old-

est \ alid name for this group is Rctifusus.

Rctifusus Dali, 1916

Flicifusus (Rctifusus) Dali, 1916: 8.

cusps of nearly ecpial length. The central teeth may

be of hvo t\'|res. 71. jessocusis, R. circus, R. i/auauiii. and

R. fiiclci have five or six sharp cusps increasing in length

from the peripheiy to the center (Figure 37). The central

teeth ol7l. roseus, R. laticiuguJatus, 71 siniilis, 71 iturupus,

and 71 attcnuatus have only tliree shaip cusps, and

the central cusp is usually longer tlian the lateral cusps

(Figure 38). The salivaiy ducts are veiy thick and straiglit.

The stomach is large compared to the prolroscis, narrow,

and has a small posteilor mixing area.

Remarks: McLean (1995) established tlie genus Rcti-

inohuia (tvy^e species by original designation, Mohuia

friclei (Dali, 1891) to incorporate several species previ-

ously assigned to the genus Mohuia Friele, 1878. Onr

analysis demonstrates that M. friclei belongs to the same

clade and is morphologically rathei' similar to 71. jessoeu-

sis, the ty|re species oi' Rctifusus. Thus, Rctiiuolmia is a

jimioi' subjective synonym ot Rctifusus. Rctifusus is often

considered to be a snbgenus of Plicifusus (e.g., Higo

et ah, 1999) but onr analysis demonstrates that it is not

closely related to the latter.

Genus Composition: We include the following spe-

cies in Rctifusus, although some others may belong to

this group as well:

Rctifusus jessocusis (Schrenck, 1863) [= Fusus (Sipho?)

mauchuricus E. A. Smitli, 1875; Chn/socloiuus hniiiueus

Dali, 1877; Mohuia okhotskaua Tiba, 1981 - .synomany

based on examination of the tspe specimens and anato-

mical studies .]

Rctifusus friclei (Dali, 1891)

Rctifusus virens (Dall, 1877)

Rctifusus ijanaiuH (Yokoyama, 1926)

Rctifusus laticiugulatus Golikov et Gulbin, 1977

Rctifusus roseus (Dall, 1877) [= Rctifusus semiplicatus

Golikov in Golikov and Scarlato, 1985; Plicifusus par-

vus Tiba, 1980; Plicifusus saginatus Tiba, 1980 - syn-

onymy based on examination of the tvpe specimens

and anatomical studies].

Rctifusus similis (Golikov et Gulbin, 1977)

Rctifusus attcnuatus (Golikov et Gulbin, 1977)

Rctifusus iturupus (Golikov et Sireiiko, 1998)

Type Speeies: Tritonium (Fusus) jessocusis Schrenck,

1863 (by original designation)-

Diagnosis: Tire genus is characterized by a snrall (on

average < 2.5 cm) shell, which has an axial and spiral

sculpture similar to that of Plicifusus (Figures 32, 34);

however, the radula has a different nrorphology (Figure

37, 38). The lateral teeth usually have three or four long

Rctifusus differs from Plicifusus in radnlar moiphologv;

from Mohuia in the form of its operculum, the pi'esence

of axial sculpture and in radnlar morphology; from Colas,

Aulacoftisus, and Latisipho in axial sculpture and radnlar

morphology'.

Glade 5 includes representatives ol Aulacofusus that are

rather uniform conchologically and morphologically'.

Figures 18-24. Anatomy. IS. Tran.sverse section of the proboscis wall oi Aulacofu.sus hereiicleeui. 19, 20. Transverse section of

the proboscis wall ot Aulacofusus hrevicauda. 21. Salivan' ducts of A. hrevicauda. 22. Transverse section of the proboscis wall ol

Rctifusus jessoensis. 2.3. Salivaiy duct of R. jessoensis. 24. Transverse section ol the proboscis wall of Plicifusus hastarius. Abbrevia-

tions: aoe, anterior esophagus; eni, circular inuscies; cut, connective tissue; crt, odontopfioral cartilage; ep, epitlieliinu; liii,

longitudinal nniscles; n, nen'es; r, radula; scl, salivan' duct.

Page 90

THE NAUTILUS, Vol. 123, No. 3

Figures 26-35. Shells. 26. Colus islmulicus. 27. Latisipho haJIii. 28. Pararefifiisiis tenuis. 29. Lafisipho Ju/polispus. 30. Pani-

niifusus kantori. .31. Plicifusus kroet/eii. .32. Rctifiisiis attenuatus. 3.3. Plicifiisus rht/.ssus. 34. Retifusus jessoensis. 3.5. Aulacofusus

Ijrevicaiichi .

Aulacofusus Dali, 1918

Aulacofusus Dali, 1918: 217.

Type Species: Fusus spitzbergeusis Reeve, 18.55 (by

original cle,signation).

Diagnosis: Tlie group i.s characterized by an elongat-

ed, inedinm-.sized fusilonn .shell sculptured with wide

spiral cords (Iroin 6 to 16 cords on the penultimate

whorl) (Figure .35). The axial sculpture is represented

only by incremental growth lines. Tlie radula structure

i.s in general the same as in Flicifisus (Figure 39).

The salivary ducts are thick-walled, with aii additional

external layer ol longitudinal muscles (Pdgure 21, lin).

The stomach is large, as comparetl to the proboscis.

and narrow, with a veiy long posterior mixing area

(Figure 16, pnia).

Remarks: The taxon was proposed as “group of spe-

cies, Aerified l)y Fusus spitzbergeusis Reeve that has a

special aspect due to the short canal and tlie promi-

nence of the spiral ribs. . .” Thus, the rank of the taxon

was not specified, but it is obvious, from the context of

the de.scription, that Dali (1918) considered it even

lower than that of a section of the genus Colus. Later,

Dali (1921) treated it as subgenus oi' Colus, a view that

has been followed by most recent authors (e.g., Higo

et ak, 1999), but not by some Russian researchers (e.g.,

Golikov and Gulbin, 1977; Kantor and Sysoev, 2()()5,

2006).

A. H. Kosyan and Yu. I. Kantor, 2009

Page 91

Figures 36—41. Radulae. 36. Plicifiisiis ki'oci/cri. .37. Puiifiisiis jcssocnsi.s. 38. Panmiifitsus hinlori. .39. Aiilacofiisii.s hn'i'icaiida .

40. Ldtisipho lii/polispiis. 41. Coins islandicns.

Species of AuJacofusus have a consideral)le coiieho-

logical similarity to species attrilnited to the genus

Coins, particularly in the shape and sculpture of the

shell (Figures 26, 35). Some anatomical characters, such

as the extremeh long, coiled proboscis Apical of Anlaco-

fiisiis (Figure 14), are also present in some species ol

Coins. Nevertheless, the presence of several autapomor-

phies ol Anhicofnsns. including stomach structure tliat is

Pa,<re 92

THE NAUTILUS, Vol. 123, No. 3

Table 1. Character coding (see Appendix 1).

perificclidiis

unique in the entire subfamily Colinae, and the histolog-

ical strnctnre of the wall of the salivai-y dncts, lead ns to

treat it as a separate genus.

Genus Composition: Alany species has been attribu-

ted to this group at various times. We inclmle the follow-

ing examined species in the snhgenns:

Aulacofusus brevicauda (Deshayes, 1832) { = Tiif()iiiin)t

schanlaricinn Aliddentlorff, 1849; Nepinnca (Sipho)

terchralis Cionld, I860)

Aidacofnsns heroidecni (Dali, 1899) (=Cohis (Aiilacofii-

siis) iiohilis Dali, 1919)

Aiilacofitsiis omhroniiis (Dali, 1919)

Auldcofiisits peiiscclidiis (Dali, 1891)

Glade 6 is the most basal clade in our study, and is

supported in only 53% of the trees. It includes three

species of the genus Neptunea Rikling, 1798: Ncptunea

anticpia (Linnaeus, 1758) (t)qre species of the genus by

subsefjuent desiguation of Sandberger, 1861), N. jag^udi-

nae Cowachev and Kantor, 1983, and N. gulhini Goiya-

chev and Kantor, 1983. The genus was included in the

analysis based on published data (Goryachev and Kan-

tor, 1983) and its detailed description is beyond the

scope of the current paper. Nevertheless, our analyses

suggest that the genus iu its conventional sense may be

parajibyletic.

Both known species pi'eviously referred to Lafisipho

(Ko.syan, 2()06b) (Figures 27, 29), do uot emerge as a

A. R. Kosyan and Yu. I. Kantor, 2009

Page 93

monophyletic group in our study, aud their taxonomic

position should be reconsidered.

Our study indicates that the anatomical characteristics

are important and suitable tor diiferentiating among tlie

genera of Colinae and Buccinidae. Despite the absence,

in many cases, of autapomorphies, many closely related

genera may be diagnosed by combinations of characters

through the use of phylogenetic techniques.

ACKNOWLEDGMENTS

Many thanks are due to Dr. M. G. Ilarasewych and

Dr. E. Strong for organizing the Neogastropod Sym-

posium during the World Congress of Malacology in

Airtwerp in 2007.

We are indebted to Dr. B.I. Sirenko and R.A.

Kormushkina from the Zoological Institute (Saint

Petersburg, Russia), Dr. A. V. Sysoev and Dr. D. L.

Ivanov from the Zoological Museum of the Moscow

State University (Moscow, Russia), Dr, A. V. Gebruk

from the P. P. Shirshov Institute of Oceanolog)' (Moscow,

Russia), Dr, D. G. Reid, Dr. f. D. Taylor, A. McLellan,

and K. Way from the British Natural Histoiy Museum

(London, UK) for help during work in the museum

collections. We are grateful to Dr. D. O. Alexeev

(VNIRO, Moscow, Russia) who placed his personal

collection of Buccinidae at our disposal. We also are

indebted to the anonymous reviewers for their valuable

comments.

LITERATURE CITED

Dali, W. H. 1902. Illirstrations and descriptions of new, iinli-

gnred or imperfectly known shells, chiefly American, in

the U. S. National Museum. Proceedings of the United

States National Museum 24(1264): 499-566, pis. 27-40.

Dali, W. H. 1918. Notes on Chn/sochvnus and other mollnsks

from the North Pacific Ocean. Proceedings of the United

States National Museum-54(2234): 207-2(34.

Dali, W. II. 1921. Summar)' of the marine shellhearing mol-

lusks of tlie northwest coast of America, from San Diego,

California, to the Polar Sea, mostly contained in the col-

lection of United States National Museum, wth illustra-

tions of hitherto unfigured species. United States

National Museum Bulletin 112: 1-217, pis, 1-22.

Fraussen, K. and Y. Ternm. 2007. The family Buccinidae, Ge-

nus Neptnnea. A Conchological Iconography. Co7i-

chbooks, Hackenheim, 166 pp., 154 plates.

Golikov, A. N. 1963. Gastropod molluscs of tlie genus Nej)tu-

nea Bolten. Fauna of USSR, Mollnsks. Leningrad, Nauka,

5(1): 1-217 [In Russian],

Golikov, A. N. 1980. Molluscs Buccininae of the World Ocean.

Fauna of USSR, Mollnsks. Leningrad, Nauka, 5(2): 1-466

[In Russian],

Golikov, A, N. and V. V. Gnlbin. 1977. Prosohranchiate gastro-

pods (Gastropoda, Prosohranchiata) of the shelf of Kurile

Islands. II. Ordo Ilamiglossa - Homoestropha. In: Fauna

of coastal zones of Kurile Islands. Nauka, Moscow, pp.

172-268.

Goiyachev, V. N. and Yu. I. Kantor. 1983. Two new Northern-

Kurile species of the genus Nepfiniea (Gastropotla. Bucci-

nidae). Zoologicheskii Zhnrnal 62(12): 1765-1774 [In

Russian],

Iligo, S., Callomon, R, and Y. Goto. 1999. Catalogue and bibli-

ography of the Marine shell-hearing Mollusca of Japan.

File Scientific Publications, Osaka, 749 pp.

Kantor, Yn. I. 1982. About the hyre species of tlie genus Volii-

iopsius (Gastropoda, Pectinihranchia). Zoologicheskii

Zliurnal 61(6): 843-850 [In Russian],

Kantor, Yu. I. 1988. Gastropod molluscs of the subgenus Aitcis-

trolcpis {Clinope^ma) (Gastropoda, Buccinidae) of the

Sea of Okhotsk. Zoologicheskii Zhnrnal 67(8): 1126-1140

[In Russian],

Kantor, Yu. I. and A.V. Sysoev. 2005. Gatalogue ol molluscs of

Russia and adjacent countries, Moscow: KMK Scientific

Press Ltd., 1-627.

Kantor, Yu. I. and A.V. Sysoev. 2006. Alarine and brackish

water Gastropoda of Russia aud adjacent countries.

KMK Scientific Press Ltd., Moscow, 1-371.

Kosiige, S. 1967. On the transfer of "Piti/morhi/nchii.s?'' tcnuis_

Okiitani, 1966 to the family Buccinidae. Wiiiis 25: 59-64.

Kosyan, A. R. 2006a. Two new species of the genus Pnraniifit-

sns Kosiige, 1967 (Buccinidae: Golinae), with notes on the

morpholog)' of Ptiraniifiisns feiuiis (Okutani, 1966).

Rutheniciri6(l-2): 5-15.'

Kosyan, A. R, 2006h. Anatomy and ta,xononiic composition ol

the genus Latisij)lio Dali (Gastropoda: Buccinidae) from

the Russian waters. Riithenica, 16 (1-2): 17-42.

McLean, J. H. 1995. Four new genera for Northeastern Pacific

prosohranch gastropods. The Nautilus 108: 39-41.

Swofford, D.L. 1998. PAUP: Phylogenetic Analysis Using Par-

simony and Other Alethods, v. 4, Sunderland: Sinaiier,

APPENDIX 1. List of characters and character states.

CEPIiALOPOlilUM

1. Operculum: 0 — wdth terminal nucleus (Figure 1), 1

— with spiral nucleus (Eigure 4), 2 — with terminal

nucleus displaced to the left.

Mantle

2. Mantle: 0 — square, 1 — length e.xceeds width.

3. Osphradium: 0 — symmetrical, I — asymmetrical.

4. Osphradium: 0 — short (< Vi of mantle length), 1 —

long (> Vi ol mantle length).

5. Ctenidium: 0 — lamellae of ctenidium wider than

lamellae of osphradium, 1 — lamellae of ctenidium

of the same width as lamellae of osphradium.

Reproductive System

6. Penis: 0 — with large seminal papilla (Figures 6, 8,

9), 1 — with small papilla (Figure 10), 2 — wdthout

papilla (Figure 5).

7. Seminal papilla: 0 — cone-shaped, encircled by fold

of skin (Figures 6, 8, 9), 1 — claw-like, 2 — absent.

8. Male genital opening: 0 — not surrounded by tiny

papillae, 1 — surrounded by multiple tiny papillae.

9. Vas deferens: 0 — thin, convoluted, not protruding

into body haemocoel, 1 — tliick, located in body

haemocoel.

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THE NAUTILUS, Vol. 123, No. 3

10. Cap.sule gland: 0 — with ventrally folded vagina, 1

— with terniinal vagina.

Dige.sti\’E Sy.stem

11. Proboscis: 0 — straight ( Figures 12-13, pr), 1 —

folded wdthin rhynchoeoel (Figure 14, pr).

12. Rhynchodeuni: U — thick-walled, everting, 1 —

thin-walled, non-everting.

13. Relative length of hnccal mass: 0 — equal in length

to contracted proboscis, 1 — half the length of the

contracted proboscis, 2 — less than half the length

of the contracted proboscis, 3 — longer tlian tlie

contracted proboscis.

14. Proboscis retractors: 0 — running along rh\nicho-

deuin and attached to roof and lateral walls of body

haeniocoel (Figure 12-13, prr), 1 — short, situated

at the base of the proboscis anti attached to the

bottom of body haemocoel (Figure 14, prr).

15. Sequence of layers in the proboscis wall [onter to

inner edges]: 0 — epithelium, circular muscles, lon-

gitudinal muscles, circular muscles, longitudinal

mnscles (Figure 19), 1 — epithelium, longitudinal

muscles, circular muscles, longitudinal muscles, cir-

cular muscles (if present) (Figure 24).

16. Salivaw glands: 0 — small and rounded (< 1/3 of

proboscis length) (Figure 12), f — beau-

shaped (> 2/3 of proboscis length) (Figures 13, 14).

17. Salivary ducts: 0 — wdthout additional longitudinal

muscle layer in the wall (Figures 22, 23), 1 — with

e.xternal layer of longitudinal mnscles in the wall

(Figures 20, 21).

IS. Salivary ducts: 0 — wdthout salivaiy sacs (Figures 13,

14), 1 — with salivary sacs (Figure 15).

19. Salivaiy tlucts: 0 — thin, convoluted (Figure 13), 1

— thick, straight (Figures 14, 15).

20. Gland of Leiblein: 0 — well tleveloped, 1 — thin,

poorly developed, 2 — absent.

21. Stomach: 0 — with small posterior mi.xing area

(Figure 17), 1 — with veiy long posterior mixing

area (Figure 16), 2 — \x4thout posterior mixing area.

22. Stomach: 0 — large (>l/3 whorl), 1 — small (<l/3

whorl).

Smell

23. Axial ribs: 0 — <14 axial ribs on last whorl, 1 — >

14 ribs on last whorl, 2 — axial ribs absent.

24. Axial ribs: 0 — s-shaped, 1 — straight, 2 — absent.

25. Spiral scnlptnre: 0 — numerons cords present (> 20

on penultimate whorl), 1 — few cords present (< 20

on penultimate whorl), 2 — cords absent.

26. Alicroscopic spiral threads: 0 — present, 1 — ab-

sent.

27. Spiral cords: 0 — absent, 1 — present, low, acnte

distally, 2 — present, rounded distallv, 3 — present,

llattened.

2S. Ratio, body whorl height / shell height: 0 — <0.7; 1

— >0.71.

29. Ratio, aperture length / shell length: 0 — <0.5; 1 —

>0.51.

R.vdula

30. Central tooth: 0 — with 3 cusps (Figures 36, 38—41),

1 — with multiple cusps, posterior tooth edge

rounded (Figure 37), 2 — with multiple cusps, pos-

terior tooth edge nearly straight.

31. Central tooth: 0 — with 3 cusps, all of equal size, 1

— with 3 cusps, medial cusp differing in size from

the marginal cusps, 2 — with more or fewer than 3

cusps.

32. Lateral teeth: 0 — with 3 cusps, 1 — with more or

fewer than 3 cusps.

33. Lateral teeth: 0 — medial cusps smallest, 1 — all

cusps equal in length.

34. Cnsps of the central tooth: 0 — do not overlap tooth

of followdng row; 1 — overlap tooth of following row.

THE NAUTILUS 123(3):95-J05, 2009

Pa«;e 95

The anatomy and relationships oi Trosclielia (Neogastropoda:

Bnceinidae): New evidence for a closer fasciolariid-bnccinid

relationship?

Alisa R. Kosyan

A.N. Seveitsox Institute of Ecoioo;\' and Evolution

Hnssian Academy ol Sciences

33 Leninski Prospekt

Moscow 119071, RUSSIA

kosalisa@ranil)ler,rn

Maria Vittoria Moclica

Marco Oliverio

Dipartimento di Bioloo;ia Aniinale e deH'Uomo

“La Sapienza" Rome Universih'

\dale deirUniversita 32

1-001S5 Roma. ITALY

[email protected]

marco.oliverio@uniromal .it

ABSTRACT

Analyses of new anat(5mical and molecular data confirm the

taxonomic position ot Troschclia hernicicusis (King, 1S46)

within the Bnceinidae and provide the framew'ork tor a review

of the relationships of the families Fasciolariidae and Bnccini-

dae. The morphology of TnisclicUa Mtirch, fS76, is similar to

tliat of other Northern Atlantic and Pacific hnccinid genera.

Anatomical examiiiation ol a nnmher ol lasciolariid species

rev^ealed only a single character, the structure ol tlie proboscis

retractor muscles, to be tliagnf)Stic of the Easciolariidae, while

other characters are shared witli the Bnceinidae. A molecular

phyiogeny also confirms a close relationship between the tvv'o

groups.

Additional kci/icords: Taxonomy, phyiogeny, anatomy, Bnccini-

dae, Easciolariidae

INTRODUCTION

Phylogenetic relationsliips among the more than 209

genera and .snhgenera included in the ga.stropod family

Bnceinidae remain quite amhiguons. The northeastern

Atlantic monotypic genus Trosclielia Mdrch, 1876, has

been classified by a number ol authors (e.g., G.O, Sars

(187S), J. Thiele (1929)) in the lamily Fasciolariidae, due

to the peculiar radnla of Trosclielia herniciensis (King,

1846). However, Bonchet and VVaren (1985) placed

Trosclielia within Bnceinidae, based on the moiphology

of its lateral teeth, whicli have multiple, uniform cusps

similar to tho.se of some otlier Bnceinidae (Thalasso-

planes Dali, 1908). To elucidate the jihylogenetic rela-

tionships ol Trosclielia with the families Bnceinidae and

Fasciolariidae, morpliological and anatomical featni'es

were studied, and partial secjnences from the mitochon-

drial 16S rllNA gene were analyzed for Trosclielia her-

niciensis as well as for a number of bnccinid and

lasciolariid taxa.

MATERIALS AND METHODS

Samples lor the present study were collected during field

work and expeditions to the West Pacific (P.yNcaw) 2004,

Philippines, and S.\nto 2006, Vannatn, organized by the

Alnsenm national d Ilistoire natnrelle, Paris), to Panama

(Neoga,stropod Workshop 2006, at the Smith.sonian Trop-

ical Research Institute, Panama), the Mediterranean Sea,

and at other localities, and supplemented by specimens

prov ided by VInsenms (BMNH - Natural Iliston' Mnse-

nm, London, UK, MNHN - Mnsenm National d'llistoire

natnrelle, Paris, France, BAU - Mnsenm of Biologia Ani-

male e deH'Uomo Department, University-’ "La Sapienza",

Rome, Italy) and colleagues. The taxa listed in Table 1

were used lor anatomical studies. Animals vv^ere dissected

and radnlae examined using an SEM. Sequence data w'as

newly generated lor several species, and supplemented

with additional taxa that were obtained from GeneBank

(see Table 2).

DNA Extraction, PGR, and Seouencing: Total DNA

wTis extracted following a standard Phenol/Ghloroform/

Ethanol protocol (Ilillis et ak, 1990) with slight modifi-

cation as prevhonsly described by Oliv'erio and Vlariottini

(2001) for mollnsks. QIAGEN QiAmp Extraction Kit

was used according to manufacturer’s instructions for

e.xtraction of DNA from difficult samples.

A region ol tlie gene encoding IbSrDNA encompass-

ing the domains IV and V (Gntell and Fox, 1988) wms

amplified using primers 16SA (5’-GGGGTGTTTATGA-

AAAAGAT-3') (Palnmbi et ak, 1991) and 16SH (5’-GG

GGTGTC;AAGTGAC;ATGAG-3’) (Espiritn et ak, 2001).

Amplification conditions w^ere as follovws (30-35 cycles):

94° for 30 sec, 45-50°G for 30 sec, 72°G for 60 sec.

When a single band vvyis olvtained the PGR product wyrs

purified using the Exo-Sap enzvmatic method. Purified

products were tlien double straml sequenced vv'itli Big-

Dye V'. 2.0 (Applied Bio.systems, Foster Gitv, GA, USA)

Pae;e 96

O

THE NAUTILUS, Vol. 123, No. 3

Table 1. Species used in the anatomical study.

using the PCR primers and sequences visualized on

automatic se(|uencer. Sequencing was performed by

Macrogen Inc. (Seoul, South Korea). Chromatograms

were analysed by Staden Package (Version 1.6.0, Staden

et ah, 199S, 2005). All setpiences have been deposited at

EVIBL (see Table 2 for accession numbers).

Sequences obtained were aligned using ChistalX

(Thompson et al., 1994; 1997) with the default settings.

The alignments obtained were manually edited. The

test implemented in PAUP* v. 4bl0 (Swolford, 2002)

was used to test for base composition homogeneity of

the sequence data aligned.

A Bayesian analysis of the aligned secpiences was

perlormed using MrBayes v. 3.1.2 (Bomjnist and Hnel-

senbeck, 2003), which sampled trees from posterior

densities using the Alarkov Chain Monte Carlo method

(Larget and Simon, 1999; Yang and Rannala, 1997),

44ie substitution model to be used in the Bayesian

analysis was chosen after evaluation by the solHvare

MrModeltest 2.2 (Nylander, 2004), wliile base frecjuen-

cies, relative rates ol the six substitution t\pes and

model parameters were estimated by MrBayes during

phylogenetic reconstruction. A four-chain metropolis-

coupled iVIonte Carlo analysis was run twice in parallel

for 10*^ generations, and trees were sampled eveiy 1000

generations, starting after a burn-in of 250000 genera-

tions. Stationarity was considered to be reached

when the average standard deviation of split frecpiencies

shown in AIrBayes was less than 0.01 (Ronquist and

llnelsenbeck, 2003). Bayesian posterior probabilities

(BPP) of a branch were estimated as the percentage of

trees (aftei' burn-in) which showed that specific node.

RESULTS

Anatomy of Troschelia berniciensis: External AIor

IMIOLOGY: Animal (Eignres 1-3) is uniform cream in col-

or. Eoot (Eignre 1, ft) partly contracted, \\4th deep

propodial groove (pi'pg) separating narrow propodinm.

Operculum oval, with terminal nucleus (Figure 1, op).

Head moderately large, bnxid (Figure 1, hcl) with pair

of long, thick tentacles, each wdth large black eye on

outer side of swelling at its base. Penis (Figure 1, p)

small, apparently underdeveloped.

M ANTLE: Mantle margin indented (Figure 3). Siphon

(Figure 3, s) moderately long, innscnlar. 0,spliradinm (os)

occupies ~ 14 mantle length, ~l/6 mantle widtli. Cteni-

dinm (ct) long, crescent-shaped, occupying 4/5 mantle

length. Hy|iohranchial gland (hg) not well developed.

Digestin'e Syste.m: Proboscis e.xtremelv long, narrow

(Figures 4-5, pr), compactly folded within rhynchotlenm

(Figure 4, rd). Buccal mass occupies ~ Vi proboscis

length. Badnla equal in length to odontophore, with

structure similar to that illustrated in Bonchet and Waren

(1985; 184, fig. 485). Proboscis attached to bottom ol

body haemocoel by proboscis retractors (Figures 4-5,

prr) emerging Irom its base, con.si.sting ~ 6 multiple

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THE NAUTILUS, Vol. 123, No. 3

Figurc.s 1-7. Anatomy ol Trosclwlia hcnticiciisis. 1. (Aphalopodinm, x'i.sceral mass removed. 2. section of esophagu.s with dnct of

gland of la'ildein. 3. Mantle. 4. Foregnt. 5. Proboscis. 6. Stomacli. 7. Internal strnctnre ol'the stomach. Ahhrexiations: ao, anterior

aorta, aoe, anterior esopliagns; cm, coinmella muscle; ct, ctenirliimi; II, loot; gl, gland ol: Leihlein; licl, head; hi, heart; kcl, kidney;

nr = ncMA’c ring; op, opercnhnn; o.s, osphradiniii; p, penis; pclg, posterior dnct ol digestive gland; poc, posterior esophagus; pr,

proboscis; prpg, propodial groove; prr, proboscis retractors; rcl, rhyncbodeum; re, rectum; .s, siphon; sg, saliwiiy glaiul; st, stomach;

vl, valve ol Leiblein.

A. R. Kosyan et al., 2009

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Figures S-17. Anatomy of Fasciolariiclae. 8—10. Pciisteniia ii.stulata. 8. Cephalopodinin, visceral mass removed. 9. Foregnt,

ventral view. 10. Foregnt lateral \iew. 11. Foregut of FnsUilafirus inediainericanus, ventral \iew. 12. Proboscis ol P. ii.stiilafa.

13—1.5. Pnstiilafirm mecliamericaiiiis. 1.3. Mantle. 14. Proboscis. 1.5. Internal structure ol the stomach. 16. Stomach ol Lalints

poUigOHUs. 17. Stomach of P ustulata. Abbre\iations: aoc, anterior esophagus; be, buccal eaxits': bni, buccal mass: cm, cohunellar

muscle; ct, ctenidiuni; clg, digestix'e gland; dgl, duct of gland ol Leibleim eye, eye; ft, loot; gl, gland ol Leiblein: gon, gonad; hd,

head; int, intestine; mo, mouth opening; mrr, medial retractor ol radula; nr, neiwe ring; odn, odontophore nen’cs; odr, odorito-

phore retractors: op, operculum; os, osphradiuni; poe, posterior esophagus; pr, proboscis: prg, prostate gland: prr, prolioscis

retractors; r, radula; rd, rhynchodeum; re, rectum; s, siphon; sd, salivaiw duct: sg, salivaiT gland; st, stomach; vl, x’alve ol Leiblein.

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THE NAUTILUS, Vol. 123, No. 3

Figures lS-29. Anatomy of Fasciolariidae. 18—21. Fiisiuus tenerifcnsis. 18. Foregut, 19. Proboscis. 20. Cephalopodium. 21.

Stomach. 22—24. Opeatosloma j)scitclodoii. 22. Mantle. 2.3. Soft parts. 24. Foregut. 25. Internal structure of the stomach. 26.

Proboscis. 27 — 29. Fasciolaria ligmiria. 27. Foregut. 28. Internal structure ot the stomach. 29. Proboscis. AbbreUations: adg,

anterior duct ol digestive gland; ao, anterior aorta; aoe, anterior esophagus; be, buccal caUty; bep, bursa copulatri.x; bin, buccal

mass; eg, capsule gland; ein, columella muscle; et, ctenidium; clg, digestive gland; eye, eye; ft, foot; gl, gland ol Leiblein; gon,

gonad; bd, bead; bg, hyjrobranchial glaml; int, intestine: mrr, medial retractor of radula; n, nei'ves; nr, nen'e ring; odr, odonto-

phore retractors; oeo, oesopliageal opening; op, operciilnni; os, osphradinni; p, penis; pdg, posterior duct ol digestive gland; poe,

posterior esophagus; pr, proboscis; prr, proboscis retractors; r, ratlula; rd, rhynchodemu; s, siphon; sd, salivaiy duct; sg, salivary

gland; spd, sperinoduct; si, stomach; vl, valve ol Leiblein.

A. R. Kosyan et al., 2009

Page 10]

Figure 30. Bayesian topoIog\' obtained for the inolecnlar

dataset. Numbers at nodes are the Bayesian Posterior Probabil-

ities.

muscle tufts. Anterior esophagus wide, convoluted. \'al\ e

of Leiblein small, pyriform, situated immediately anterior

to nerye ring. Saliyaiy' glands ol medium size (Figures 2, 4,

sg), tightly packed with newe ring (Figure 4, nr) by con-

nectiye tissue. Anterior aorta (Figure 4, ao) passes

through neiwe ring, runs parallel to posterior escjphagns.

Stomach narrow, occupying ~ 1/3 whorl (Figure 6, st).

Posterior mixing area is absent, stomach walls lined with

high, transverse folds of epithelium. Opening of posterior

duct of digestive gland large, situated above the oesopha-

geal opening (Figure 7, pdg).

Anatomy of FasciolariitI Species: Eight species of

Fasciolariidae, representing seven genera from three sub-

families, were studied anatomically (Table 1). Main external

moiphological features included a folded, large muscular

foot (Figures 8, 20, 23, ft), a broad head with relatively

short tentacles (Figures S, 20, 23, hcl), and an operculum

with a terminal nucleus (Figures 8, 23, op). The mantle

(Figures 13, 22) has a moderately large, muscular siphon,

a ctenidium occupying '/4-1/3 of the mantle \\4dth, and

an aswnmetrical o.sphradium that may be large (Fusi)}tis

tencrifensis Fladorn and Rolan, 1999, Latinis poh/gonits

(Gmelin, 1791), Pustulatinis incdimneiicaniis (Hertlein

and Stnmg, 1951a), Fasciolaiia lignana (Linnaeus, 1758),

Peristemia nassatula (Lamarck, 1822), P. iishtlala (Reeve,

1847) - Figure 13, os) or small {0))catosfoiita pficudodou

(Burrow, 1815) - Figure 22, os).

Digestive System: The proboscis is straight and never

coiled within the rhynchodeum. It may be rather shoi't

in Pustulatinis mcdiomericanus (Figure 14), Latinis

pohpgoiiiis, Opcatostoma pscudodou (Figure 20), Fusi-

ims teuerifensis (Figure 19), aud Fasciolaria liguaria

(Figure 29), \\4th the length of the buccal mass and