1VIBL/WH0I

THE NAUTILUS

Volume 110

1996-1997

AUTHOR INDEX

Ahlstedt, S. a 97 Kkausk, M. K 31, 42

Arnold. VV. S 31. 42 Lavzek. J. B 102

Beis, 1 1 Lyons, W. G 31. 42

DEL Rio. C. J 77 Marelll D. C 31. 42

Dillon. R. T. ]R 97 Petuch. E. J 17, 122

E\'ERsoLE. A. G 107 Sabei.i.i. B 94

Graf, D. L 45 Sysoev, A. V. 22

Harasewtch. M. G 55 Tamam. M 94

HouART, R 12 Theler, J. L 94

Kaloyianni, M 1 Wise, J. B 76

Kirk, S. G 102 Xii\i\i,i, R 1

NEWTAXA PROPOSED IN N'OLUME 110 (199fi-!997)

GASTROPODA

Astriwa (Astraliiiin ) polkcnsis Petiicli, 1997, new .specie.s (Tnrbinidae) 125

Pijrazishms ki'itdrcui Petuch, 1997. new species (Potiimididae) 126

Ci'stiiinccrithiniu Pefiicli, 1997, new genus (Cerithiitlae) 126

Prismaccrithium Petuch, 1997, new genus (Geritliiidae) 127

Prismacerithium prisma Petuch, 1997, new species (Cerithiidae) 127

Pachijcroinmium dalli Petuch, 1997, new species (Naticidae) 127

Pachi/rrommiiim mansfieldi Petuch, 1997, new species (Naticidae) 127

Cahisacyprnca Petuch. 1997. new species (Cvpraeidae) 17

Cahisdciiprncn liriani Petuch. 1997. new species (Cvpraeidae) 18

Cnhisiuiipniea cliierri Petuch. 1997. new species (Cvpraeidae) 19

C.nlitsdnipraca sdraaotiwnsis Petuch. 1997. new species (C\praeidae) 19

Ciiprarorhis kenilmii Petucli. 1997. new species (Cvpraeidae) 129

Iphitus rohertsi .SabeHi & Taviani, 1997, new species (Epitoniidae) 94

Ergalatax obscura Houart. 1996. new species ( Muricidae ) 13

Spiniful<s,ur ncmmulattim Petuch. 1997. new species ( Busvconidae) 131

Solcnostcira f/uicnnuccn.sis Petuch. 1997. new species ( Bucciinidae) 131

Vasuin snudnnccnsis Petuch, 1 997, new species (Turhinehidae) 131

Falsilijiid kcndrcwi Petuclu 1997, new species (Volutidae) 131

Conomitni kcndrcwi Petncli, 1997. new .species (Volutidae) 131

Dcniiiiuiriifi dtilli Petuch. 1997. new species (Marginellidae) 134

Persicuhi dockcnji Petuch. 1997. new species (Marginellidae) 134

Persicida nuicncili Petuch. 1997, new species (Marginellidae) 134

Persicuhi suwannccnsis Petuch. 1997. new species ( Marginellidae) 134

Pniiuim ciictic Petuch. 1997. new species ( Marginellidae ) 134

Pninum jcssicac Petuch. 1997. new species (Marginellidae) 135

Pniiuun sdiidmc Petuch, 1997. new species (Marginellidae) 135

Ahi/ss(iili(iiiiiui .Svsoev, 1996, new genus (Conidae) 24

Conns (Asprcll(i) kcndrcwi Petuch, 1997, new species (Conidae) 136

Plcurofnsia dowlingi Petuch, 1997. new species (Turridae) 136

Suwannccscaphri Petuch. 1 997. new species (Cyclichnidae) 136

Suwnnnccscapha lindric Petuch. 1997. new species (( A'cliehnidae) 136

Petitilln Wise. 1997, mw name (Pvraniidelli(iae) 76

BI\.\I.\I.\

Rctrotapcs del Hfo, 1 997, new germs (N'eneridae) 80

Rctrotaprs fnci^ocnsis del Ri'o, 1997, new species (Veneridae) 89

Rcfrolapcs ninfasicnsis del Rfo. 1997, new species (Wneridae) 82

THE NAUTILUS

Voluiiw 110, Number 1

August 13, 1996

ISSN 0028-1344

A quarterhj devoted

to malacology.

Marine Biological Laboratory/

Woods Hole Oceanographic Institution

Library

AUG 2 2 1996

Woods Hole, MA 02543

EDITOR-IN-CHIEF

Dr. M. G. Harasewych

Division of Mollusks

National Mnseuni of

Natural Historv'

Smithsonian Institution

Washinj^ton, DC 20560

CONSULTING EDITORS

Dr. Riicliger Bieler

Department of Invertebrates

Field Museum of

Natural Histor)'

Chicago, IL 60605

Dr Arthur E. Bogan

Freshwater Molluscaii Research

36 Venus Wav

Sewell, NJ OSOSO

Dr. Robert T. Dillon, Jr

Department of Biology

College of Charleston

Charleston, SC 29424

Dr. William K. Emerson

Department of Living Invertebrates

The American Museum of Natural

History

New York, NY 10024

Mr. Richard I. Johnson

Department of Mollusks

Museum of Comparative Zoology

Harvard University

Cambridge, MA 02138

Dr Aurele La Rocque

Department of Geology

The Ohio State University

Columbus, OH 43210

Dr James H. McLean

Department of Malacolog)'

Los Angeles Count)' Museum of

Natural History

900 Exposition Boulevard

Los Angeles, CA 90007

Dr Arthur S. Merrill

% Department of Mollusks

Museum of Comparative Zoology

Harvard University

Cambridge, MA 02138

Dr Paula M. Mikkelsen

Department of Malacology

Delaware Museum of Natural History

P.O. Box 3937

Wilmington, DE 19807

Dr Donald R. Moore

Division of Marine Geology

and Geophysics

Rosenstiel School of Marine and

Atmospheric Science

University of Miami

4600 Rickenbacker Causeway

Miami, FL 33149

Dr Gustav Paulay

Marine Laboratoiy

University of Guam

Mangilao, Guam 96923

Mr Richard E. Petit

PO. Box 30

North Myrtle Beach, SC 29582

Dr. Edward J. Petuch

Department of Geology

Florida Atlantic University

Boca Raton, FL 33431

Dr David H. Stansbeiy

Museum of Zoology

The Ohio State Universitv

Columbus, OH 43210

Dr Ruth D. Turner

Department of Mollusks

Museum of Comparative Zoology

Har\'ard University-

Cambridge, MA 02138

Dr Geerat J. Venneij

Department of Geology

University of California at Davis

Davis, CA 95616

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THE€7NAUTILUS

Volume 110, Nti tuber 1

Auo^ust 13, 1996

ISSN U028-1344

CONTENTS

Rallio Xoniali Time Course of Tissue Specific Metabolism of the Suhtidal

Martha kaIo\ianni Castrojiod Hcxaplcx tninruliis uiulcr Anaeroliic Conditions 1

Isidoros Beis

Roland Houart On tin- Identih ol Morula nwiicnsi Dall, 1923 and

Description ot a New Species of Eriifilatax from the Red

Sea (Gastropoda: Muricidae: Ergalataxinae) 12

Edward J. Petuch Cdlusiictipnica. A New. Possibly Neoti'nic CJenus ol

Cowries (CJastropoda: Cvpraeidae) Ironi the Pliocene ot

Southern Florida 17

Alexander V. Sysoev Tiixononiic Notes on South African Deep-Sea Couoitlean

C;astro[)ods (Gastropoila: Conoidea) described bv K. H.

Barnard, 1963 ' 22

Notice 30

Marine Biological Laboratory /

Woods Hole Oceanographic Institution

Library

AUG 2 Z 1996

Woodi Molt, MA 02S43

THE NAUTILUS 110(1):1-11, 1996

Page 1

Time Course of Tissue Specific Metabolism of the Subtidal

Gastropod Hexaplex trunculus under Anaerobic Conditions

Rallio Xomali

Martha Kaloyianni'

Isidores Beis

Laborator\ of Animal Plnsiology,

Department of Zoolog>', School of

Science, Aristotle University of

Tliessaloniki. Thessaloniki 54006,

Greece

ABSTRACT

Anaerobic energy metabolism \\ as investigated in foot, mantle

and hepatopancreas of the subtidal gastropod Hexaplex trim-

ciilu-s (Linne, 1758) {=Murex tnineulu.s). C^hanges in the le\els

of adein lates, substrates and end products were monitored over

a time course of 16 hours of anoxia. It is suggested that unlike

intertidal species, the succinate pathway is a metabolic route

of minor importance during anoxia in subtidal gastropods. The

existence of an alternative electron acceptor, apart from Oj

and fumarate, in the tissues of the subtidal gastropods under

anaerobic conditions is discussed.

Key M'ords: metabolism, anaerobiosis, subtidal, gastropods, foot,

mantle, hepatopancreas, Hexaplex trunculus.

INTRODUCTION

It is generally accepted tfiat within the animal kingdom

the ability to sustain periods of oxygen deprivation varies

among the different phyla. MoUusks and particularly

bivalves present well developed capacities for the sur-

vival of environmental anoxia and for this reason have

been extensively studied in terms of their anaerobic en-

ergy metabolism (de Zvvaan, 19<S3).The available liter-

ature on molluscan metabolism during anaerobiosis deals

with metabolic responses of separate tissues, as well as

with comparative differences and similarities of carbo-

hydrate catabolism of various organs. The data on met-

abolic tissue specific differences derive mainly from stud-

ies on the bivalve Mytilus edulis Linne, 1758 (Li\ing-

stone & Bayne. 1976; Kluytmans et a!., 1977; Zurburg

& Kliixtmans, 1980), on the cherrystone clam Merce-

iiaria mercenaria Linne, 1758 (Korycan & Storey, 1983),

on the oyster Crassostrea virginica (Gmelin, 1791)

(Eberlee et al.. 1983), on the common cockle Cardium

edule Linne, 1758 (= Cerastodernia e£/w/p)(Meinardus

Author to whom correspondence should be addressed.

& Gade, 1981) and on the marine whelk Busycotypus

canaliculatum (Linne, 1758) (Storey, 1988; Eberlee &

Storey, 1988; Whitman & Storey, 1989; Storey et al.,

1990), where it is suggested that the metabolic response

of different tissues during anaerobiosis and recovery dif-

fers quantitatively but not qualitatively. Succinate and

alanine were found to be the main end products of an-

aerobiosis in all tissues studied (Kluytmans et al., 1977;

Zurburg & Kluytmans, 1980; Meinardus & Gade, 1981;

Eberlee et al.. 1983; Korycan & Storey, 1983; Eberlee &

Storey, 1988; Storey et ai, 1990), whereas glycogen and

aspartate served as the major substrates of anaerobic fer-

mentation (Korvcan & Storey, 1983, Eberlee & Storey,

1988),

The above pattern of tissue specific response refers to

studies made mainly on species living in the intertidal

zone, in which e.xposure to air is a relatively common

occurence. Consequently very little information is avail-

able on metabolic tissue specificities of subtidal species

(Wieser, 1980; Carpene et ai. 1981; GSde et ai. 1984)

concerning mainly some end products of their anaerobic

metabolism, Hexaplex trunculus is reported to be ca-

pable of sustaining oxygen depletion for a period up to

100-200 hours (Zs Nagy, 1971), This species normally

lives in the subtidal zone, where it may be exposed to

anoxic conditions caused by a variety of factors, includ-

ing for instance severe water pollution combined with

high water temperatures. Despite the variety of studies

on anaerobic metabolism of intertidal species, no sub-

stantial information is available on the potential mech-

anisms of anoxia tolerance of subtidal species. Therefore,

we examined metabolic patterns used under anaerobic

conditions in different tissues of Hexaplex trunculus.

Comparison of the overall anaerobic metabolism of sub-

tidal and intertidal species show substantial tissue specific

differences in the time course of metabolic changes dur-

ing anaerobiosis, as well as in the products accumulated

and substrates utilized. In addition, the comparison of

the general response between intertidal and subtidal gas-

tropods revealed a different metabolic scheme concern-

Page 2

THE NAUTILUS, Vol. 110, No. 1

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Duration ofanaerobiosis (h)

Figs 1-3. Changes in the concentrations of aspartate, glycogen

and glucose in the hepatopancreas of Hexaplex tninculus dur-

ing various periods of anaerobiosis. 1) aspartate, 2) glycogen,

■3) glucose. * Significantly different from the corresponding

aerobic condition (t-test P<0.05). ** Significantly different

from the corresponding aerobic condition (t-test P<0.00.5)

ing the use of the succinate pathway by Htxuplcx tnin-

culus tissues during anaerobiosis.

MATERIALS AND METHODS

Animals and Chemicals: Specimens of Hexaplex trun-

culus were obtained from the shores of Thessaloniki

through a local fisherman. They were kept in circulating

seawater at about 2()°C and were used two to three days

after arrival. Aerobic animals were sampled directly from

the seawater tank. To impose anoxia the animals were

placed in a vessel of seawater with an air tight cover,

containig 1 L of seawater that had been previously de-

o.xygeiiated by bubbling a steady stream of nitrogen gas

through it for 1 h to give zero oxygen tension as moni-

tored w ith an ox\ gen electrode (Digital Oxygen System,

model 10, Rank Brothers Manufacture). Animals were

kept under anoxic conditions for 2, 4, 8, 12 and 16 h at

20°C. At the end of the anoxic incubation the shells were

quickly opened and the tissues were rapidly excised,

blotted and immediately frozen in liquid nitrogen. Tis-

sues were stored at -80°C until assayed.

Substrates, enzymes and coenzymes were purchased

from Sigma Chemicals Co. (St Louis, L^SA). All other

chemicals were purchased from Serva (Heidelberg, Ger-

many).

Preparations of Tissue Extracts: Samples of frozen tissue

were ground to a powder under liquid nitrogen using a

mortar and a pestle. The powdered tissue was extracted

by adding three volumes of cold perchloric acid (HCIO4,

10% w/v). The precipitated protein was removed b\

centrifugation at 4000 x g lor 10 minutes and the su-

pernatants were neutralized with 3M potassium bicar-

bonate (KHC03).The precipitated potassium perchlorate

was removed by centrifugation as above and the super-

natants were taken for determination of metabolite con-

centrations.

Determination of Metabolites: Glycogen was deter-

mined by the method of Keppler and Decker (1974),

aspartate by the method of Bergmeyer et al. (1974a) and

glucose by the method of Bergmeyer et al. (1974b). Al-

anine was assayed according to Williamson D.H.(1974)

and succinate according to Williamson JR. (1974).

L-Lactate was measured according to Gutmann and

Wahlefeld (1974) and octopine according to Storey et

al. (1979). ATP was quantified bv the method of Lam-

precht and Trautschold (1974), ADP and AMP by the

method of Jaworek et al. (1974).

Statistics: Results are presented as means ± SEM (n=5).

Tests for significant differences between groups used the

Student's t-test.

RESULTS

Hepatopancreas: Figures 1-3 show the changes in as-

partate, gl\ cogen and glucose content of the hepatopan-

creas of Hexaplex trunculus during 16h of anaerobiosis.

Aspartate content decreased throughout all the periods

of anoxia tested. Aspartate levels measured in the tissue

were lowest alter 12h of anaerol)iosis. (iKcogen and glu-

cose levels tended to increase throughout anaerobiosis.

(ilucose concentration increased 10-told during 16h of

anoxia.

Figures 4-7 illustrate the time course of the changes

R. Xomali et al., 1996

Page 3

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Duration of anaerobiosis (h)

Figs 4-7. Changes in the concentrations of lactate, octopine, alannie and succinate in the hepatopancreas of Hexaplex trunculiis

during various periods of anaerobiosis. 4) lactate, 5) octopine, 6) alanine, 7) succinate. * SignificantK different from the corresponding

aerobic condition (t-test P<0.05). ** Significantly different from the corresponding aerobic condition (t-test P<0.00.5)

in levels of potential end products of Hexaplex trunculus

hepatopancreas during anoxia. Alanine accumulated

during the first hours of anoxia. Thereafter its levels

started to fall, but remained significantK increased com-

pared to the aerobic state. Succinate accumulated after

the 4th hour of anoxia. The maximum alanine accu-

mulation (2.9 ^mol/g wet mass) observed during the first

two hours of anoxia exceeded that of succinate (0.8 ^mol/

g wet mass) observed after 16 h of anaerobiosis. Octopine

accumulated significantly after 12h of anoxia, while no

significant lactate accumulation was observed through-

out the 16h anaerobiosis.

From the data shown in figures 8-11 a sharp decline

in ATP content can be detected, while levels of ADP

and AMP are not significantly changed. The energy

charge shifts from 0.61 to 0.49 after 4h of anoxia. There-

after, it slightly increases to 0.5.3 after 16h of anoxia.

Foot Muscle: Figures 12-14 show changes in concentra-

tions of aspartate, glycogen and glucose in the foot muscle

of Hexaplex trunculus during 16h of anoxia. As in the

hepatopancreas, aspartate levels had fallen significantly

by the 4th hour of anoxia. At that time its levels are

reduced by 5.5 ^mol/g wet mass. Thereafter, its con-

centration remained almost constant. Contrary to the

results in hepatopancreas, glycogen levels had fallen by

the 4th hour of anoxia. After 8h of anaerobiosis a sharp

reduction in glycogen content was observed (44.2 nvao\/

g wet mass). Glucose levels were significantly increased

after 8 hours, after 12h of anaerobiosis its concentration

was almost double that of the aerobic state.

Changes in the concentrations of lactate, octopine, al-

anine and succinate during 16h of anoxia in the foot

muscle of Hexaplex trunculus are shown in figures 15-

18. Onh alanine levels ro.se significantly during anoxia.

The maximum alanine accumulation was observed after

12h of anoxia and was about 3.7 ixmo\/g wet mass. No

significant changes in octopine. lactate and succinate lev-

els were obser\ed.

In figures 19-22 the changes in adenylates and in the

energy charge in the foot muscle during 16h of anoxia

of Hexaplex trunculus are presented. A significant de-

Page 4

THE NAUTILUS, Vol. 110, No. 1

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Duration of anaerobiosis (h)

Figs 8-11. Changes in the concentrations of ATP, ADP, .■\MP and energy charge in the hepatopancreas of Hcxaplex trunculus

(hiring various periods of anaerobiosis. S) ATP, 9) ADP, 10) AMP, 1 1 ) energy charge. * Significantly different from the corresponding

aeroljic condition (t-test P<0.0.5). ** Significantly different from the corresponding aerobic condition {t-test P<0.005)

dine in ATP content together with a signiBcant increase

in ADP level were observed after 4h of ano.xia. AMP

levels remained almost unaltered. A decline in energy

charge was also evident. Energy charge shifted from 0.87

to 0.72 during the first four hours of ano.xia. Thereafter,

it remained almost constant.

iMantlc: Figures 23-25 show changes in the concentration

of aspartate, glycogen and glucose in the mantle of Hex-

aplex trunctiluH during various [periods of anaerobiosis.

Among the substrates measured only aspartate showed

a significant decline, being reduced to 3.4 ^imo]/ g wet

mass during the 8 first hours of anoxia. Glycogen levels

remained almost constant throughout anoxia. Glucose

concentration in the mantle increased and showed a sig-

nificant rise after 12 and 16h of anoxia exposure, com-

pared to the aerobic state.

Among the metabolites presented in figures 26-29, a

significant rise in lactate content was observed after 8h

of an(jxia. During the first four hours alanine levels de-

creased. Thereafter an apparent increase in alanine con-

tent v\as observed. This increase was not significantly

different from the value of the aerobic control, owing to

high variation in the alanine content of different animals.

There were no significant differences in octopine and

succinate content among the difierent periods tested.

Concerning the changes of the adenylates shown in

figures 30-33, a significant decrease in ATP content was

seen until the fourth hour of anoxia. Levels of ADP

increasexl during the first eight hours, but declined there-

after again, reaching control values. No significant change

in AMP concentration was observed. Energy charge de-

clined, mainly during the first eight hours, decreasing

from 0.9 to 0.73.

DISCUSSION

The ability of in\ertel)rate anaerobes to sustain long pe-

riods of anoxia is niainlv based on the metabolic adap-

tations of their tissues. These species are able to reduce

their metabolic rate and at the same time to maintain

R. Xomali et al., 1996

Page 5

their energy reserves so as to confront the energetic needs

of their tisssues. The ATP production is generally based

on the anaerobic breakdown of glycogen. In contrast to

mammalian systems, this anaerobic breakdown of gly-

cogen does not lead to lactate production. Various end

products accumulate during anaerobiosis depending on

species, as well as on the duration of anaerobiosis. In

intertidal bivalve mollusks, the main end product of the

first stages of anaerobiosis is alanine, while at later stages

succinate, propionate, acetate and fatty acids can be de-

tected (de Zwaan, 1983). Hochachka and Mustafa (1972)

proposed that amino acids should be an important an-

aerobic fuel in intertidal bivalve tissues and they put

forward a metabolic map integrating the metabolism of

glucose, aspartate and glutamate during ano.xia. The

metabolic behaviour of subtidal species during anaero-

biosis is almost unknown.

The results of the present study revealed that foot

muscle, mantle and hepatopancreas of Hexaplex trun-

ciilits catabolize aspartate and accumulate alanine in re-

sponse to 16h of ano.xia. The pattern of metabolism differs

among tissues. In the hepatopancreas, aspartate declines

during the earlv stages of anaerobiosis and is accompa-

nied b\ a simultaneous rise in alanine content (figs, 1

and 6). After this initial response, aspartate declines more

slowly, while alanine accumulates up to the 16th hour

of anaerobiosis, compared to the aerobic state. Gluta-

mate, a possible end product of anaerobiosis in intertidal

species, is not accumulated in the hepatopancreas of Hex-

aplex trtinculus during anoxia (results not shown). A

significant rise in glucose content was observed, starting

from the very beginning of oxygen deprivation and rose

by a factor of ten till the 16th hour of anaerobiosis (fig.

3). It could be suggested that Hexaplex trtinculus he-

patopancreas responds rapidly to anoxia b\ catabolising

mainh aspartate and accumulating initialK alanine and

subsequently succinate (fig. 1,6,7). Elevation of tissue glu-

cose could not be attributed to glycogen degradation

since glycogen levels showed no signs of decreasing

throughout anoxia. Similarly no significant decrease in

glycogen content was found in hepatopancreas, foot, and

phasic adductor muscle of the cherrystone clam Mer-

cenaria mereenaria (Korycan & Storey, 1983), No change

in the percentage of phosphorvlase in the active a form

was found in the adductor muscle of Mytilus edulis over

a time course of up to 18h of anoxia (Ebberink & Sali-

mans, 1982).

In the foot muscle of Hexaplex Iruneulus, the response

to anoxic stress is less rapid than in the hepatopancreas.

This agrees with the findings of Whitman and Storey

(1989), who suggested that the processes involved in an-

oxia induced modification of givcolv tic enzymes are more

rapidh initiated in non muscular than muscular tissues.

A sharp decline in aspartate content is obvious after four

hours of anoxia, while alanine accumulation is maximal

after twelve hours of anoxia (fig. 12 and 17). The low

levels of succinate did not change throughout anoxia (fig.

18). This could either suggest that the succinate pathway

is not functioning in the foot muscle of Hexaplex trnn-

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Figs 12-14. Changes in the concentrations of aspartate, gly-

cogen and glucose in the foot muscle of Hexaplex trunculus

during various periods of anaerobiosis. 12) aspartate, 13) gi\-

cogen, 14) glucose. * SigiiificaiitK different from the corre-

sponding aerobic condition (t-test P<0.05)

cuius, or that succinate is rapidly metabolized to another

end product (i.e. propionate or acetate). Similarly, it was

reported that in the subtidal marine gastropod Nassarius

reticulatu.s only alanine and no succinate accumulated

throughout anoxia (Wieser, f980). The Mediterranean

bivalve Scapharca cf. cornea Reeve, 1843 also produced

no succinate during 72h of anoxia (Carpene et al., 1981).

However, in the foot muscle of the subtidal whelk Nassa

Page 6

THE NAUTILUS, Vol. 110, No. 1

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Duration of anaerobiosis (h)

Figs 15-18. Changes in the concentrations of lactate, octopine, alanine and succinate in the foot muscle of Hexaplex trunculus

during various periods of anaerobiosis. 15) lactate, 16) octopine, 17) alanine, 18) succinate. * SignificantK different from the

corresponding aerobic condition (t-test P<0.05).

mutalnlis succinate was produced, but only in low con-

centrations (Gade et ai, 1984). In contrast to hepato-

pancreas, glycogen content falls sharply after four hours

of anoxia (fig. 13). The observed glycogen degradation

in the foot muscle could be correlated with the contin-

uous effort of the animal to move away from areas de-

prived of oxygen. The elevated glucose levels, probably

derived from glycogen catabolism, could be the main

metabolic substrate used to support muscular work under

anaerobic conditions. No lactate and octopine accumu-

lation was found {fig. 15, 16). Our results concur with

the data presented by Kluytmans et ai (1983) and Gade

et al. (1984). It seems, therefore, that the foot muscle of

Hexaplex truncitlus shows no signs of energ\' stress dur-

ing the first two hours of anoxia. Later, it responds to

anoxia by catabolizing aspartate and glycogen and ac-

cumulating mainly alanine.

Due to the declining levels of aspartate in the mantle

during anaerobiosis, aspartate appears to be the main

source of energy production during anoxia. Alanine ac-

cumulation presented a somewhat different pattern of

accumulation compared to the two other tissues exam-

ined (fig. 28). There was an initial decline in its level,

followed by a subsequent increase up to the 16th hour

of anoxic stress. However, owing to great variation in

mantle alanine content among different individuals, the

values did not differ from the control. Similarly, alanine

did not accumulate in the mantle tissue of Mytilus ediilis

at reduced oxygen tensions, but it accumulated in the

posterior adductor muscle under the same conditions

(Livingstone & Ba\iie, 1976).

Glucose levels increased during anoxia in all three

tissues examined. Glucose levels in the hepatopancreas

rose from the very begining of anoxia, reaching its max-

imum value after 16h of oxvgen deprivation where a

tenfold increase in its content could be observed. How-

ever, glucose increases in loot muscle only after 8h of

anoxia and in mantle ti.ssue after 12h of anoxia. This

delayed increase could be correlated w ith the time need-

ed for glucose residues to be mobilized from the hepa-

topancreas and transferred to the foot muscle and the

mantle via the hemolv mph. While glucose levels in the

R. Xomali et al., 1996

Page 7

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16

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4

— I —

12

— I —

16

Duration of anaerobiosis (h)

Figs 19-22. Changes in the concentrations ot ATP, ADP, AMP and energ\ charge in the foot muscle of Hexaplcx trunculus during

various periods of anaerobiosis. 19) ATP, 20) ADP, 21) AMP, 22) energ\ charge, * Significant!) different from the corresponding

aerobic condition (t-test P<0,U.5). ** Significant!) different from the corresponding aerobic condition (t-test P<000.5)

hepatopancreas increased by tenfold, only a t\\ ofold in-

crease was observed in foot muscle and mantle. This

suggests that glucose was synthesized in the hepatopan-

creas to supply other tissues and help them meet their

metabolic demands during conditions of prolonged ox-

ygen deprivation. A similar rise in glucose concentration

during anoxia was observed in various tissues of the cher-

rystone clam, Mercenaria mercenaria where glucose lev-

els doubled throughout anoxia (Kor\can & Storey, 1983).

In the fresh water snail Lijmnaea stagnalis Linne, a

strong and rapid hyperglycaemic response is evoked dur-

ing anoxia (VVijsman et al., 1988). Hemminga et al.

(1985a) were able to prove that in L. stagnalis a h\per-

gKcemic factor is released by the central nervous system

during conditions of glycogen breakdown. The cerebral

ganglia contain the main release sites of this factor. The

ganglionectomised animals showed reduced levels of hy-

pergKcemia associated with exposure to anaerobic con-

ditions (Hemminga et al.. 1985b).

Al! three tissues showed a significant decrease in ATP

content (figs. 8, 19, 30). In hepatopancreas and mantle

tissue this decline is evident after the first two hours of

anoxia, while in foot muscle a significant decline is ob-

vious after the fourth hour. The sharp reduction in ATP

levels of the hepatopancreas may in part be attributed

to gluconeogenic procedures that continually consume

ATP which is bareK replenished, owning to the low levels

of glycoK'sis. Only the hepatopancreas shows a significant

accumulation of succinate throughout anoxia. Therefore,

the succinate pathway may be activated in the tissue

after the initial drop in ATP reserves. Although ATP

levels decrease in foot muscle and mantle tissue, these

organs still maintain a relatively high energy charge, so

activation of succinate pathwa\' ma\ not be necessary.

In contrast to intertidal species studied so far (Zurburg

& Kluytmans, 1980; Michaelidis & Beis, 1990), the suc-

cinate pathway is a route of minor importance for an-

aerobic energy production in mantle and foot muscle of

I'age 8

THE NAUTILUS, Vol. 110, No. 1

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c

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Duration of anaerobiosis (h)

Fips 23-25. Changes in the concenlrations of aspartate, gl\-

cogen and ghieose in the mantle of Hcxaplcx truiiculii.s during

various periods of anaerobiosi.s. 23) aspartate, 24) glycogen, 25)

glucose. * Significantly different from the corresponding aer-

obic condition (t-tesl P<0.05). ** Significantly different from

the corresponding aerobic condition (t-test P<0,00.5),

the subtidal species Hexaplex tnniculns. This observa-

tion tan he correlated with its habitat. In general, most

sublittoral inollusks can tolerate anoxia less well than

intertidal molhisks. There seems to be a relationship be-

tween the habitat (and its oxygenation) and the animals

anoxia tolerance. (Gade, 1983). The production of suc-

cinate during anaerobiosis provides the animals with ex-

tra ATP. ilue to the coupling of the fumarate reductase

system w ith the first phosphorylation site of the electron

transport chain (Holwerda & de Zwaan, 1979; 1980; Koh-

ler, 1976; 1980). The enhancement of the energetic ef-

ficiency of the anaerobic metabolism in these animals

enables them to withstand long periods of anoxia. It

seems that subtidal species have not adapted a function-

ing succinate pathway.

However, the question of the final electron acceptor

under conditions of ox\gen depri\ation remains anan-

swered. Hexaplex trunculiis survives anaerobiosis forlOO-

200h (Zs Nagy, 1971). Furthermore, the lack of succinate

accumulation in its tissues after several hours of anoxia

suggests that another electron acceptor, apart from O2

and fumarate may exist. In 1971, Zs Nag\' suggested that

energy production in mollusks during prolonged anoxia

may be realized by means of a mechanism called "anoxic

emlogenous oxidation '. This mechanism is localized in

the cvtosomes and its essential point is that molecular

oxygen is replaced by an internal electron acceptor, a

lipochrome pigment located in the cytosomes. Further-

more, heavy pigmentation was found in the central ner-

vous s\ stem of Hexaplex truneiilus (Zs Nag\-, 1971 ). Con-

sequently, the latter process, which takes place within

special cell organelles, may function for a certain period

of time, in the tissues of this subtidal gastropod for pro-

duction of biologicalK utilizable energ\- under conditions

of oxygen depri\ation. The biochemical mechanisms for

such control processes warrant further investigation.

LITERATURE CITED

Bergnie>er, H. U., E Bernt, H Miillering and G Phleiderer.

1974a. L-aspartate. In: Bergine>er, H. L'.(ed.). Methods

of enz\matic anahsis .\cademic Press, New York, 4:1696-

1700. '

Bergmeyer, H. I , E. Bernt, F, Sclimidt and H Stork 1974b,

D-glucose, /;i. Bergmeyer, H. U.(ed.). Methods of enzy-

matic analysis, .Academic Press, New York. 3:1196-1201.

Carpene, E., W. Cortesi and VV. Zurburg. 1981. Anaerobic

energy metabolism in the bivalves Scap/iarca inaeqtiivalvis

(Bruguiere) and \'(';h« gallina L. .•Abstracts of the Third

ESC^PB Congress. Pergamon Press, Oxford, pp. 118-119.

de Zv\aan, A, 1983. Carbonh\drate catabolism in bivalves.

In: Hochachka, P. W (ed ). The Moliu.sca, .Academic Press,

New York. 1:137-175.

Ebberink, R. H. M. and M. Salimans. 1982. Control of gly-

cogen phosphorylase activit\ in the posterior adductor

muscle of the .sea nuissel Mytilus cclulis. Journal of Com-

parative Physiolog) 148:27-33.

Eberlee. J. C. and J .M Storey. 1988. Tissue-specific bio-

chemical responses tluring anoxia and reco\er\ in the

channelled whelk. Journal ol Experimental Marine Biol-

og\ and Ecology 121:l(i5-171

Eberlee, J. C, J. M. Storey and K. B. Store). 1983. .Anaero-

biosis, recovery from anoxia, and the role of strombine

and alanopine in the oyster Crassustrea virginica. Cana-

dian Journal of Zoology 61: 2682-2687,

Glide, G, 1983, Energ\ metalxilism (}f arthropods and mol-

lusks during en\ionniental and lunctioiial anaeroljiosis

Journal oi Experimental Zoolog\ 228415-429

Gade, G.. K II Carlsson ami (.', Mcinardus, 1984 Energy

metabniisni ui the loot of the marine gastropod \assa

R. Xomali et al., 1996

Page 9

V>

(0

E

w

«

o

E

o

c

(0

0)

o

0 75

0 50

025

0 00

0 50n

01

c

■q.

o

O

0 25

27

0 00

Duration of anaerobiosis (h)

Figs 26-29. Changes in the concentrations of lactate, octopine, alanine and succinate in the mantle of Hcxaplex ininculus during

various periods of anaerobiosis. 26) lactate, 27) octopine, 2S) alanine, 29) succinate. * Significantly different from the corresponding

aerobic condition (t-test P<0.05). ** Significantly different from the corresponding aerobic condition (t-test P<0.05).

nmtabilis during environmental and functional anaero-

biosis. Marine Biolog> 80:49-56.

Gutmann, I. and W. Wahlefeld. 1974. L- Lactate. In: Berg-

meyer, H. U.(ed.). Methods of enzymatic analysis. .Aca-

demic Press, New York. .3:1464-1468.

Henmiinga, M. A., J. J. Maaskant, \\ Koomen and J Joosse.

1985a. Neuroendocrine control of gKcogen mobilization

in the freshwater snail Lymnea stagnalis. General Com-

parative Endocrinology 57:117-123.

Hemminga, M. A., J. J. Maaskant, J. van der Plas and P.A.

Gabbott. 1985b. The hyperglycemic factor of the CNS

of the freshwater snail Lymnea stagnalis: interaction with

glucose stimulation of glycogen s> nthesis and evidence for

its release during anaerobiosis. General Comparative En-

docrinology 59:301-307.

Hochachka. P. W and T. Mustafa 1972 Invertebrate Fac-

ultative .Anaerobiosis. Science 178:1056-1060.

Holwerda DA. and A. de Zwaan. 1979. Fumarate reductase

of Mijtilus edulis L. Marine Biological Letters 1:33-40

Holwerda, D. A. and A. de Zwaan. 1980. On the role of

fumarate reductase in anaerobic carbohydrate catabolism

of Mytilus ednlis L. Comparative Biochemistrv and Phys-

iology 67B:447-453.

Jaworek, D., \V. Gruber and H U. Bergmeyer. 1974. Aden-

osine-5-diphosphate and Adenosine-5-monophosphate. In:

Bergmeyer, H. U.(ed.). Methods of enzymatic analysis.

Academic Press, New York. 4:2127-213l'

Keppler, D. and K Decker 1974. Glycogen. In: Bergmeyer,

H. U.(ed.). Methods of enzvmatic anaKsis. .Academic Press,

New York. 3:1127-1131.

Kluytmans, J. H., A. M. T. de Bont, J. Janus and T. C. M.

Wijsman. 1977. Time dependent changes and tissue

specificities in the accumulation of anerobic fermentation

products in the sea mussel Mytilus edulis. Comparative

Biochemistry and Physiolog) 588:81-87.

Kluytmans, J H.] A. M. T. de Bont, E. C. J. Kruitwagen, H J.

L. Ravenstein and P. R. V'eenhof. 1983. Anaerobic ca-

pacities and anaerobic energy production of some medi-

terranean bivalves. Comparati\e Biochemistrv and Phys-

iology. 75B:171-179.

Kohler, P 1976. Hydrogen transport in the muscle mito-

chondria of Ascaris suum. In: Bossche, H. V. D. (ed.).

Biochemistry of Parasites and Host-Parasite Relationships.

North-Holland, Amsterdam, pp. 125-132.

Kohler, P. 1980. The function of mitochondrial enzsmes in

parasitic helminths. /;i: \itale, L. and \'. Simeon (eds).

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THE NAUTILUS, Vol. 110, No. 1

(0

re

E

■«->

o

E

zl

c

re

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0,030

0015

0000

u

uJ

05-

00

12

16

-I—

8

12

16

Duration of anaerobiosis (h)

Figs 30-33. Changes in the concentrations of ATF, .ADP, AMP anil energy charge in the foot muscle of Hcxaplcx tninculus during

various periods of anaerobiosis. 30) ATP, 31) ADP, 32) AMP, .33) energy charge. * SignificantK- different from the corresponding

aerobic condition (t-test P<0.05).

Proceedings of the FEBS Special Meeting on Enzymes:

Industrial and Clinical Enzvmologv. Pergamon Press, Ox-

ford. 61:24.3-256.

Korycan, S. A. and K. B Storey, 1983, Organ specific me-

tabolism during ano.xia and recovery from anoxia in the

cherrystone clam, Mcrcenaria mercenaria. Canadian

journal of Zoology 61: 267-4-2681,

Lamprechi, VV. and 1. Trautschold. 1974. Adenosine-5-tri-

phosphate. In: Bergmeyer, H. U.(ed.). Methods of enzy-

matic analysis. Academic Press, New York. 4:2101-2110.

Livingstone, D R. and B. L. Bayne. 1976. Responses of My-

lilus cdnlis L to low oxygen tension: Anaerobic metabolism

of the posterior adductor mu.scle and mantle tissues Jour-

nal of Comparative Physiology 114:143-15.5

Meinardus, G. and G. V.Mv. 1981. Anaerobic luelabolisni ol

the common cockle Cardium edule — IV. Time dependent

changes of metabolites in the foot and gill ti.ssue induced

by anoxia and electrical stimulation Comparative Bio-

chemistry and Physiology 70B: 271-277,

Michaelidis, B and I. Beis, 1990 Studies on the anaerobic

energy metabolism in the loot muscle of marine gastropod

Patella cacrulea L. Coniparati\'e Biochemistr\ and Phvs-

iology 95B:493-500.

Storey, K. B. 1988. Mechanisms of glycolytic control during

facultati\e anaerobiosis in a marine mollusc: tissue-specific

analysis of glycogen phosphorylase and fructose- 2,6-bis-

phosphate, Canadian Journal of Zoolog) 66:1767-1771.

Store) , K B,, J. M. Storey, J. Kjell and P. \V. Hochachka. 1979,

Octopine metabolism in Sepia officinalis, effect of h\ poxia

and metabolite loads on the blood levels of octopine and

related compounds C'anadian |ournal of Zoologv 57:2331-

2336

Storey, K, B,, D, A, Kelly, J. A. Duncan and J. M Storey 1990,

,\naerobiosis and organ specific regulation of glycoKsis in

a rji.ii iiir u liclk C>anadian Journal of Zoology 68:974-980.

\\ luljiian, K, E and K. B, Storey, 1989. Organ-specific reg-

ulation of phosphofructokinase during facultative anaero-

biosis in tile marine whelk Busycotijpu.s canaliciilatiiin.

(Canadian Journal of Zoolog\ 6970-75.

\\ ieser, VV. 1980. Metabolic end products in three species of

marine gastropoils Journal oi Marine 15iological Associa-

liciii nl the I riUril Kuii;d(.iii 60 175-1.80,

R. Xomali et al., 1996

Page 11

Wijman, T. C. M , J J Maaskant, P Balm and J FClijnstra

1988. H\ pergKcaeniia in relation to anoxia and stress in

the freshwater snail Lynuiat'a stagnalis. (:oniparati\e Bio-

cheniistr\ and Ph>'siolog\ 89B:.55-63.

Williamson, D, H, 1974. L- alanine. /(i: Bergme\er, H. U.(ed.).

Methods of enz\ niatic anaK sis. .\cadeniie Press, .\e\v York.

4:1679-1682.

Williamson, J. R. 1974. Succinate /n; Bergmeyer, H. U.(ed.).

Methods of enz\ matic anal\ sis. .Academic Press, New York.

3:1616-1621.

Zs Nagy 1971 The lipochrome pigment of molluscan neurons

as a specific electron acceptor. Comparative Biochemistry

and Physiology 40A:595-602.

Zurburg, W., and J. H. Kkntmans. 1980. Organ specific

changes in energy metabolism due to anaerobiosis in sea

mussel Mtitilus cilulis L Comparative Biochemistrv and

Physiology 67B:317-322.

THE NAUTILUS 110(1):12-16, 1996

Page 12

On the Identity of Morula martensi Dall, 1923 and Description

of a New Species of Ergalatax from the Red Sea

(Gastropoda: Muricidae: Ergalataxinae)

Roland Houart

Research associate

Institut Royal des Sciences Natiirelles

de Belgique

Departement des Invertebres Recents

Rue \'autier, 29

B-1000 Brussels, Belgium

ABSTRACT

Morula martensi Dall, 1923 differs from the similar species

Ergalatax margariticula (Broderip, 1833) by having a smaller,

narrower and white shell The species erroneously identified as

Ergalatax or Cronia martensi (Dall, 1923) in recent literature

is named here as Ergalatax obscura n.sp.

Key Words. Gastropoda, Muricidae, Morula martensi, new

species

INTRODUCTION

Martens (1874:95, pi. 5, fig. 49) identified and illustrated

a shell from the Hausknecht collection as Purpura (Ri-

cinula) siderea Reeve, 1846. Examination of the type

materia! of Ricinula siderea (3 syntypes BMNH 1968478,

type locality: Philippines, e.\ coll. H. Cuming), revealed

that it is a buccinid, and not the species illustrated by

Martens, which Dall (1923:304) subsequently named

Morula martensi. The seven specimens examined by

Martens, all from the Persian CJulf, are stored in the

Museum fiir Xaturkunde, Berlin. Martens' original figure

is here reproduced (fig. 2), and the actual specimen fig-

ured (figs. 12, 13).

Martens' description, figure, and comparisons with

Purpura iindata (Chemnitz, 1795:pl. 192, fig. 1852) [ =

Ergalatax margariticola (Broderip, 1833)], my re-e.\-

amination of the specimens of the Hausknecht collection

and comparison with specimens from Qatar (coll. R.

Houart), reveal that this species is similar to E. margari-

ticola. The shells from Oman and the Persian Gulf are

white with black blotches on the spiral cords (figs. 12-

15), and differ from E. margariticula. which has a broad-

er, more darkly pigmented shell. Specimens from Oman

were illustrated by Boscii and Bosch (1982:95, 1989:60)

as Cronia konkanensis (Melvill, 1893), which, however,

is probably a distinct species, more closely related to

Thais muricina (Blainville, 1832) and T. marginatra

(Blainviile, 1832), both classified in Rapaninae. II the

populations from the Persian Gulf prove to be only an

ecological form of £. margariticola. then E. martensi

(Dall, 1923) will become a junior synonym of tliat species.

Ergalatax n^artensi was reported to be a s\non\ m of E.

contracta (Reeve, 1846) by Houart (1995:251). It is here

considered a distinct species.

Three distinct species with similar gross morpholog\

have been confused in the recent literature. Ergalatax

martensi (Dall, 1923), from Oman and the Persian Gulf,

is closely related to but probably distinct from E. mar-

gariticola (Broderip, 1833); Ergalatax obscura, new spe-

cies, from the Gulf of Aden, the Red Sea, the Gulf of

.Acjaba and the Mediterranean coast of Turke\ , is a dis-

tinct species identified incorrectly as Cronia (or Erga-

latax) martensi (Dall) in recent literature; and Thais

konkanensis (Melvill, 1893), from the Indo-West Pacific,

which is closely related to T. muricina (Blainville, 1832)

and T. marginata (Blainville, 1832).

My researchs were incomplete when I received pho-

tographs and material for identification from Turkey,

from G. Buzurro and VV. Engl. .\t the time, I provision-

ally identified the new Mediterranean immigrant as Er-

galatax martensi, and it was reported as such by Buzurro,

Engl and Tiimtiirk (1995) and by Engl (1995). It is now-

evident that E. martensi is confined to Oman and the

Persian Gull, and tiiat the Turkish species is £. obscura

n.sp.

SYSTEMATICS

Order Neogastro()oda Thiele, 1929

SuperfamiK Muricoidea Rafinesque, 1815

Family Muricidae Rafinesque, 1815

SubfamiK Ergalataxinae Kuroda & Habe, 1971

(Jenus Ergalatax Iredale, 1931

Ergalatax Iredale, 1931:231. T\pe species (by original

R. Houart, 1996

Page 13

designation): Ergalatax rcciinens Iredale, 1931; oft Syd-

ne\' Harbour, New South Wales, Australia.

Ergalatax martensi (Dall, 1923)

(figs. 2, 12-15)

Morula nujrtcnsi Dall, 1923:304, new name for Morula aiderea

von Martens,1874, not Reeve, 1846.

Cronia konkanauis—V.O'iCH & BOSCH, 1982:95, te.xt fig.;

SMYTHE, 1982:60, pi 1, fig i: B0SC:H & BOSCH, 1989:

60, text fig. (not Ricinula hjiikanensis Melvill, 1893).

NOT Cronia mar(e;!si— SHARABATI, 1984:pl. 19, fig, 9;

SINGER & MIENIS, 1991b:58, fig. 19 (in part); COU-

LOMBEL, 1994:73, text figs {==Ergalatax olxcura n.sp).

NOT Ergalatax martensi BUZl'RRO, ENGL & TiiMTiiRK,

1995: 17, text fig,; ENGL,1995: 46, fig. 10 (= Ergalatax

obscura n.sp).

Description: Shell up to 26.5 mm in length at maturity,

slender, squamous. Spire high with 6 weakly convex,

shouldered teleoconch whorls. Suture adpressed. Proto-

conch eroded in all specimens examined. Axial sculpture

of teleoconch whorls consisting of high, strong, nodose

ribs, producing blunt knobs at intersection with spiral

cords. 9 or 10 ribs on first and second whorls, 8-10 on

third, 9-12 on fourth and fifth, 7-10 on last whorl. Ribs

more strongK developed at shoulder. Other axial sculp-

ture of numerous growth lamellae. Spiral sculpture of

low, squamous cords of various strength, lisually 13 or

14 cords on last whorl, and 6 additional, smaller cords

on shoulder. Aperture narrow, ovate. Columellar lip

smooth, occasionalK with 1 or 3 weak folds abapicalK .

Lip adherent. Anal notch narrow, deep. Outer lip cren-

ulate, with 6-8 weak, elongate denticles within. Siphonal

canal short, broad, open. White, with primary spiral cords,

or knobs only, topped with dark brown or black. Aperture

glossv white. Operculum dark brown with subterminal

nucleus, l^adula unknow n.

Type material: Lectotxpe (here selected), the specimen

figured by Martens (1874) as Purpura (Ricinula) siderea,

not Reeve, 1846, and 6 paralectotypes ZMB 21596.

Type locality: Persian Gulf, here designated.

Other material examined: QATAR: Doha, landing stage,

coll. R. Houart (10).

Distribution: Oman and the Persian Gulf.

Remarks: .\s stated above, the shell is close to Ergalatax

margariticola (Broderip, 1833), but differs in being usu-

ally smaller relative to the number of teleoconch whorls,

narrower, and in having a white ground color and a

white, glossy aperture, instead of cream or light brown

in E. margariticola.

Ergalatax obscura n.sp.

(Figs 1, 3-8)

Cronia mar/fri.si— SHARABATI. 1984; pi 19, fig. 9, 9a, 9b;

49

Figures 1.2. 1. Protoconch of Ergalatax obscura n.sp. (scale

bar: 0.5 mm), 2. Fig. 49 of Martens (1874), as Purpura siderea

(Reeve).

SINGER & MIENIS, 1991b: 58, fig. 19; COULOMBEL,

1994: 73, text figs) (not Morula martensi Dall, 1923).

Drupella rugosa SINGER & MIENIS, 1991a: 18, fig. 6 (not

Murex rugosus Born, 1778),

Cronia cf konkanensis—GlVf<iCm & TISSELLI, 1995: 8, text

figs.

Ergalatax martensi— Bill RRO. ENGL & TuMTiiRK, 1995:

(no pag), text fig; ENGL, 1995: 46, fig. 10 (not Morula

martensi Dall, 1923),

Description: Shell medium sized for the genus, up to

25.5 mm in length at maturity, heavy, stout. Spire high

with 3-1- protoconch whorls (partially broken), and up

to 7 broad, strongly shouldered teleoconch whorls. Suture

adpressed. Protoconch conical, acute, whorls smooth. Ax-

ial sculpture of teleoconch whorls consisting of high,

roundeci, nodose ribs: 10 or 11 from first to penultimate

whorl, 6 to 8 on last whorl. Spiral sculpture of high,

strong, primary, secondary, and tertiary cords. Sculpture

forming high, nodose knobs at intersection of spiral cords

and axial ribs. .Aperture relatively small, ovate. Golu-

mellar lip with 2 or 3 weak knobs abapically, rim ad-

herent. .\nal notch broad, moderately deep. Outer lip

weaklv crenulate, with 7 strong, elongate denticles with-

in. Siphonal canal short, broad, broadly open. Milky-

white, creamy-white, or tan, usually with some light to

dark brown coloured spiral cords on shoulder and on

more prominent nodes. .Aperture cream or pale vellow

within. Operculum ergalataxine, with subterminal nu-

cleus. Ratlula unknown.

Type material: RED SE.A: Djibouti, Perim, Strait of Bab

el' Mandeb. Holotvpe MNHN; paratypes: MNHN (4),

coll. R. Houart (1); GULF OF ADEN: Djibouti, Obock.

Paratvpes: MNHN (9); Yemen, Aden. Paratvpes: MNHN

(4), AMS C. 309658 (1), NMNZ M.270540 (1), NM L1647/

T1336 (1), coll. R. Houart (1).

Type locality: Red Sea, Djibouti, Perim, Strait of Bab el

Mandeb.

Other material examined: GULF OF ADEN (no other

data): coll. R. Houart (3); RED SEA: Massawa, MNHN

(2); no other data, coll. R. Houart (1 ); GULF OF AQABA:

Egypt, Sharm El Sheik, coll. R. Houart (2); Israel, Eilat,

coli. R. Houart (3).

Page 14

THE NAUTILUS, Vol. 110, No. 1

Figures 3-8. Ergalatax ohscura n.sp. 3-4. Holotype MNHN, Djibouti, Perim, Strait of Bab el Mandab, 24 2nim 5. Paratspe

MNHN, .Aden, Yemen. 24.2 mm, 6. Sliarm El Sheik, 20 mm. coll. R, Houart. 7. Parat\'pe MNHN, Aden, Yemen, 19.2 mm. 8.

Paratype MNHN, Gulf of .\den. Djibouti. Obock. 2.3 mm. Figures 9, 10. Ergalatax cuntracta (Reeve, 1846). 9. New Caledonia,

31.9 mm, coll, R, Houart 10. Sri Lanka. 22,8 nnn, coll, R. Houart.

R. Houart, 1996

Page 15

Figure 11. Ergalatax heptagonalis (R.^eve, 1846), 31 mm, syntype BMNH 1968476. Figures 12-15. Ergalatax nmrtensi (Dall,

1923). 12-13. Persian Gulf, Hausknecht coll., illustrated by Martens (1874), 21.4 mm. Museum fiir Naturkunde, Berlin n° 21.596.

14-15. Persian Gulf, Qatar, 23.2 mm, coll. R. Houart.

Distribution: Gulf of Aden, Red Sea, and Gulf of Aqaba.

Mediterranean: Turkey, from Iskenderun toTasucu (Bu-

zurro, Engl & Tiimtiirk, 1995; Engl, 1995; Giunchi &

Tisselli, 1995).

Etymology: obscurus — (Latin) enigmatic, [indistinct is a

more literal translation].

Remarks: Ergalatax ohscura n.sp. differs from E. mar-

tcnsi in having a smaller, more strongl\' shouldered shell

with broader, less numerous axial ribs, and more ad-

pressed suture. It is milky white, cream or tan with light

or dark brown blotches, with a cream or pale yellow

aperture, while E. martcnsi is white with dark brown or

black blotches, with a glossy white aperture. Ergalatax

ohscura n.sp. differs consistantly from the similar E. con-

tracta (Figs 9-10) in its stouter shell with stronger, more

pronounced, and more nodose axial ribs, broader spiral

cords, shorter and relatively broader siphonal canal, and

more adpressed suture.

I am not aware of intermediate forms between £.

obscura and £. contracta, and no specimen of £. ohscura

has been found elsewhere in the extensive material of

£. contracta examined from localities throughout the

Indo-West Pacific. However, there are two typical spec-

imens of £. contracta in the lot containing the holotype

of £. ohscura.

From £. heptagonalis (Reeve, 1846) (Fig. 11), £. oh-

scura differs in its higher, more pronounced, and more

nodose axial ribs, and in its broader spiral cords.

Although the shell of £. ohscura seems to be close to

Page 16

THE NAUTILUS, Vol. 110, No. 1

some forms of Drupella rugosa (Born, 1778), a Rapan-

inae, it differs in being stouter, in hav ing a lower spire,

fewer, broader spiral threads, and a more adpressed su-

ture. The operculum of E. obscura is typically ergala-

ta.xine with a subterminal nucleus, compared to the typ-

icalK rapanine "D shaped operculum of D. rugosa, with

lateral nucleus.

ACKNOWLEDGEMENTS

I am very indebted to Prof. R. Kilias (Museum fiir Na-

turkunde der Humboldt Universitat zu Berlin) for loan-

ing material, to G. Buzurro (Monza, Italy), and G. Engl

(Diisseldorf, Germany), for information on the Turkish

specimens of E. obscura n. sp., to the staff of the Natural

History Museum, London, for their collaboration during

my visit, to Dr. B. A. Marshall (Museum of New Zealand,

Wellington) and to Prof. E. H. Yokes (Tulane University)

for their remarks and comments on the manuscript.

LITERATURE CITED

Bosch, D. and E. Bosch, 1982. Seasheils of Oman. Longman

Group, England. 206 pp.

Bosch, D. ii E. Bosch, 1989. Seasheils of Southern Arabia. Mo-

tivate, Dubai. 95 pp

Buzurro, G., W. Eiigi & I. Tiimtiirk, 1995. Bivalven und

gastropoden dereuropaischen meere (4): Ergalatax mar-

tensi (Dall, 1923) (Muricidae), ein neuer Lesseps'scher

einwanderer von der Tiirkischen Siidkiiste. Club Conch\ -

lia informationen 27(1): 17-18.

C;heninitz, J. H , 1795. NeuessystematischesConchylien-Cab-

inet. G. .\. Raspe, Niirnberg 11:.312 + 124 pp.

Goulonibel, A., 1994. Coquiliages de Djibouti Edisud, La

Calade, Aix-en-Provence. 143 pp.

Dall. W. H., 1923. Notes on Drup« and A/ort^/a. Proceedings

of the Academy of Natural Sciences of Philadelphia

75(1 923): 303-306.

Engl, W., 1995. Specie prevalentementeLessepsianeattestate

lungo le coste Turche. Bollitino Malacologico 31(l-4):43-

50.

Giunchi, L. & M, Tisselli, 1995. Cronia cf. konhanensis (Mel-

vill, 1893), new Indo-Paciiic host in the Mediterranean

Sea. La Conchiglia 27(275):8-9.

Houart, R., 1995. The Ergalataxinae (Gastropoda, Muricidae)

from the New Caledonia region with some comments on

the subfamily and the description of thirteen new species

from the Indo-West Pacific. Bulletin du Museum national

d'Histoire naturelle, Paris, 4e ser., 16.-\(2-4): 197-245.

Martens, E., von, 1874. L'eber vorderasiatische Conchylien

nach den sammlungen des Prot. Hausknecht, Fischer, Cas-

sel. 127 pp.

Sharabati, D., 1984. Red Sea Shells, KPI, London. 128 pp.

Singer, B S. & H K. Mienis, 1991a. Shells of the Red Sea.

The familv Thaidididae (sic) (I). La Conchiglia 27(260):

16-19,

Singer, B, S. & H. K. Mienis, 1991b. Shells of the Red Sea.

Thf family Thaididae (II). La Conchiglia 27(261 ):54-60.

Smythe, K., 1982. Seasheils of the .Arabian Gulf. George Allen

& L'uwin, London. 123 pp.

THE NAUTILUS 110(1):17-21, 1996

Page 17

Cahisacijpraea, A New, Possibly Neotenic Genus of Cowries

(Gastropoda: Cypraeidae) from the Pliocene of

Southern Florida

Edward J. Petuch

Department of Geology

Florida Atlantic University

Boca Raton, Horida 33431 USA

ABSTRACT

The new cypraeid genus Calu^acypraca n, gen. is described

from the Okeechobean Sea region of Pliocene southern Florida

This new genus is unusual among American fossil cowries in

that it bears many juvenile shell characters into adulthood and

possibly represents a neotenic evolutionary offshoot of Sipho-

cypraea. Based on paleoecological inferences, Calusacypraea

was an inhabitant of shallow water lagoonal and estuarine en-

vironments and became extinct at the end of the Pliocene \\ ith

the obliteration of the Okeechobean lagoonal svstenis. Three

new Calusacypraca species are described; C. briani n. sp, (from

Petuch Unit 7), C. diierri n. sp. (type of the genus, from Petuch

Unit 10), and C. tequesta nsp. (from Petuch Unit 4). The

previously described species, C. sarasotaensis (Petuch, 1994),

is included in this new genus.

Key words: C\praeidae, Calusacypraca. Pliocene, Florida.

INTRODUCTION

Southern Florida has long been known to house the larg-

est Pliocene cov\rie fauna found anywhere in the United

States. The works of Heilprin (1886), Mansfield (1931),

Olsson and Petit (1964, 1968), Parodiz (1988), and Petuch

(1986, 1991, 1994), altogether, show that the ecosystems

of the Okeechobean Sea region of Pliocene Florida (Pe-

tuch, 1994:26) contained at least 23 different cowrie spe-

cies. With the exception of one species of Pseudozonaria,

all were thought to belong to a single endemic genus,

Siphocypraea Heilprin, 1886. The southern Caribbean

Gatunian cow ries previously placed in Siphocypraea are

now placed in Barycypraea (Muracypraea). This classic

Okeechobean genus radiated into numerous neritic hab-

itats during the late Pliocene, but became extinct in tlie

early Pleistocene. The genera Macrocypraea and Luria

later replaced Siphocypraea in the local reef commu-

nities (Petuch, 1994:11).

Recently, studies of new collections of fossil cowries

from the Quality .Aggregates, Inc., quarries at Sarasota

(housed at Florida Atlantic University) have shown that

there existed a second endemic cowrie genus in the Plio-

cene Okeechobean Sea. This previously-overlooked ge-

nus, here named Calusacypraea, bears many juvenile

cliaracters into adulthood (so much so that I discarded

numerous choice adult specimens, thinking that they

were mereK' juvenile or subadult Siphocypraea species!).

This new, possibly neotenic, genus and its new species

are described and discussed here.

SYSTEMATICS

Superfamily Cypraeoidea Rafinesque, 1815

Family Cypraeidae Rafinesque, 1815

Genus Calusacypraca new genus

Diagnosis: Cvpraeid shells of average to large size, rang-

ing from 35 to 70 mm in length, extremely inflated, thin,

delicate, bulliform; apices characteristically very de-

pressed, forming distinct apical pit; posterior edge of lip

typically highly-developed, greatly-projecting beyond

plane of spire; columellar side of apical area poorly-

developed, only slightly projecting beyond plane of spire;

anterior extremities poorly developed, without distinct

projecting beaks; ends of anterior extremities do not ex-

tend beyond end of anterior canal, producing rounded,

blunt, sheared-off appearance; in some species, anterior

extremity on labial side almost absent, with extremity

blending into rounded end of lip and giving shell juvenile

appearance; bases of shells very rounded; margins of

shells poorly-developed, rounded, with only thin enamel

deposits; anterior canal thickened, bordered by promi-

nent enamel collar; apertures proportionally narrow,

variabK w idening tow ard anterior end; columellar teeth

proportionally small, thin, and narrow, extending along

length of columella; some species with row of obsolete,

secondar\ teeth developed along anterior end of colu-

mella; labial teeth well-developed, confined to edge of

lip; tossula nonexistent; color pattern (when preserved)

composed of numerous small, closely-spaced dots on dor-

sum.

Type Species: Calusacypraea duerri Petuch, new spe-

cies, lower Pinecrest Beds (Petuch Unit 10), Tamiami

Page 18

THE NAUTILUS, Vol. 110, No. 1

Formation, Piacenzian Pliocene ot southern Florida,

U.S.A..

Other Species .4ssigned to Calusacypraea: Cahisacy-

praea hriani Petuch, new species, Pinecrest Beds (Petuch

Unit 7), Tamiami Formation; Calusacypraea saraso-

taensis (Petuch, 1994), Pinecrest Beds (Petuch Unit 3),

Tamiami Formation; Calusacypraea tequesta Petuch,

new species, Pinecrest Beds (Petuch Unit 4), Tamiami

Formation.

Stratigraphic Range: .\t present, known oni\ from the

Piacenzian Pliocene of southern Florida (Pinecrest Beds,

Tamiami F"ormation).

Etymology: Named for the Calusa Indian tribe of pre-

Columbian southern Florida.

Remarks: Calusacypraea is most closely related to the

s\ mpatric genus Siphocypraca and probably represents

a neotenic evolutionary offslioot of this sister group. The

new genus differs from Siphucypraca in consistently hav-

ing much thinner, more delicate, and much more inflated

shells with a distinctly juvenile appearance (an example

of neoteny?). The anterior extremities of Siphocypraca

are well-developed and flange-like, with most species

having the extremities formed into large, greatly-pro-

jecting beaks that extend beyond the end of the siphonal

canal (Figure 9). In Calusacypraea the anterior extrem-

ities are distinctly truncated and poorK developed, never

extending beyond the end of the siphonal canal, a mor-

phology generally seen only in juvenile cowries. In some

species of Calusacypraea (such as C. sarasotacnsis) the

extremities are asymmetrically developed, with the labial

extremity being virtualK' unformed and blending into

the curvature of the lip, much as in a juvenile.

The greatest differences between Calusacypraea and

Siphocypraca are seen in the posterior area of their shells.

In Siphocypraca, the posterior extremities are well-de-

veloped and symmetrical, with the columellar extremity

projecting as much or nearly as much as the labial ex-

tremity and characteristically forming a deep, narrow,

and curved apical sulcus (F"igure 11). In Calusacypraea

the columellar posterior extremity is nearly obsolete, be-

ing represented by only a thin, angled, bladelike keel.

Unlike Siphocypraca, Calusacypraea species lack the

curved, narrow apical sulcus, having, instead, a propor-

tionally small, sharply-defined, deeply-impressed apical

pit (Figure 10).

Ecologically, Calusacypraea species appear to have

occupied a different niche than did Siphocypraca species.

Without exception, all lour known species of the new

genus have only ever been found in muddy-sand, ex-

treme nearshore facies of the Pinecrest Beds. These de-

positional environments imply estuarine conditions, and

Calusacypraea ma\ ha\ e represented a neotenic offshoot

of Siphocypraca that had evolved to live in the extensive

brackish water lagoons (such as the paleoestuary of the

proto-Myakka River) that fringed the entire western coast

of the Pliocene Floridian peninsula (Petuch, 1994, Fig.

10). With the onset of the late Pliocene glacioeustatic

regressive sequences, these estuarine environments were

obliterated and the ecologicalK -narrow genus became

extinct.

The descriptions of three new Calusacypraea species

are given here, along with the description of the only

previously-known species, C. sarasotacnsis (Petuch,

1994). The holotypes are deposited in the Invertebrate

Paleontology collections of the Florida Museum of Nat-

ural History , L ni\ersit\ of Florida, Gainesville, Florida,

and bear LF numbers.

Calusacypraea hriani new species

(figures 1, 2)

Description: Basic morphology as for genus; shell large

for genus, elongated and subcylindrical; aperture nar-

row, widening slightly at anterior end; columellar lip

with 22 small, narrow teeth; columellar teeth sub-obso-

lete tow ard posterior end, becoming stronger and better-

developed toward anterior end; outer lip with 26 small

but well-developed teeth; base of shell roimded; margins

poorly developed; apical pit deep, surrounded b\' raised

apical collar; posterior columellar extremity small but

well-developed, narrow and bladelike.

Type Material: Holotype, UF 6824S, length 73 mm,

width 43 mm; 2 paratypes, lengths 75 mm and 72 mm.

Graves Museum of Archaeology and Natural History,

Dania, Florida; 3 paratypes, lengths 75 mm, 73 mm, and

68 mm, collection of the author, Florida Atlantic Uni-

versity.

Type Locality: Pinecrest Beds Fauna (Petuch I'nit 7),

Tamiami Formation, in Phase 7 pit. Quality Aggregates,

Inc., Sarasota, Sarasota County, Florida.

Remarks: Calusacypraea hriani is the most elongated

and cylindriform species in Calusacypraea. It is most

similar to the older C. duerri from Petuch Unit 10, but

differs in having a much narroyver aperture, in having

proportionally larger and coarser teeth on the outer lip,

and in having smaller and less elongated teeth on the

columellar lip. The posterior columellar extremity of C.

hriani is also better-developed and larger than that of C.

duerri, producing a much more distinct posterior sulcus.

Calusacypraea hriani yvas collected trom sediments

that indicate a very shallow, quiet, intertidal mud flat

Figures 1-1 I. Species ol Calusacypraea, iitw genus, Irom the coastal lagoonai environments of the Pliocene Okeechobean Sea.

1.2. Calusacypraea briani, new species, holotype (length 7.3 mm), Pinecrest Beds Fauna (Petuch Unit 7), Tamiami Formation. 3.4.

Calusacypraea duerri, neyv species, holotype, length 64 mm. Pinecrest Beds Fauna (Petuch I'nit 10), Tamiami Formation, 5.6.

Calusacypraea sarasotacnsis (Petuch, 1994), length 42 mm, specimen from the Pinecrest Beds Fauna (Petuch L'nit .3), Tamiami

E. J. Petuch, 1996

Page 19

Formation, APAC pit, Sarasota. 7.8. Calusacypraea tequesta, new species, holot\pe, length 73 mm, Pinecrest Beds Fauna (Petuch

Unit 4), Tamiami Formation. 9. Siphucypiaca problematica Heiiprin, 1886, length .56 mm, specimen from the Caloosahatchee

Fauna, Okeechobee Formation, in the Cochran Pit, La Belle, Hendry County. Type species of Siphocypraea Heiiprin, 1886. 10.

Apical view of Calusacypraea duerri, new species (type species of Calusacypraea), showing details of apical pit morphology. 11.

Apical view of Siphocypraea problematica. showing details of apical sulcus morphology.

Page 20

THE NAUTILUS, Vol. 110, No. 1

environment and w as louiul u ith other estuarine species

sucli as Echinojulfiiir dalli Petucli, 1994, Cymatophos

lindae Petuch, 1991, and Bttsycun Irupicalia Petuch, 1994.

Etymology: Named for my son, Brian N. Petuch, one of

my best field assistants.

Calwiacijpraea ducrri new species

(figures 3, 4)

Description: Basic morphology as for genus; shell very

inflated, rounded, ovate; aperture wide, flaring toward

anterior end; columellar lip with 24 small, narrovw and

elongated teeth; posterior columellar teeth less developed

than anterior teeth, frequently bifurcated; outer lip with

23 small teeth; base of shell rounded; margins poorly

developed; apical pit deep; posterior columellar extrem-

ity small, low, poorly-developed.

Type Material: Holotype, UP 68249, length 64 mm,

width 40 mm; 3 paratypes, 72 mm, 66 mm, and 65 mm,

Graves Museum of Archaeology and Natural History,

Dania, Florida; 3 paratypes, lengths 70 mm, 65 mm, and

61 mm, collection of the author, Florida Atlantic Uni-

versity; 1 parat\ pe in the collection of Mr. Richard Duerr,

Hollywood, Florida.

Type Locality: Pinecrest Beds Fauna (Petuch Unit 10),

Tamiami Formation, in Phase 6 pit. Quality Aggregates,

Inc., Sarasota, Sarasota County, Florida.

Remarks: Calusacypraea diteni. the type species of the

genus, is most similar to the younger and stratigraphi-

calK -higher C. sarasotaensis (Petuch, 1994), but differs

in being a much larger shell with more numerous and

better-developed columellar and labial teeth. The apical

pit of C. duerri is deeper and more impressed than that

of C. sarasotaensis. and the posterior columellar extrem-

ity of the new species is much lower and more poorly-

developed than that of the younger, descendant species.

Calusacypraea duerri was an inhabitant of a muddy-

sand substrate, shallow water ecosystem dominated by

the large bivalves Mcrccnaria iridacnoides (Lamarck,

1818), M, ochlochoneensis (Mansfield, 1932), and M.

rileyi (Conrad, 1838), and classic estuarine gastropods

such as the melongenids Tropochasca pctiti Olsson, 1967,

Ecltinojulgur cannoni Petuch, 1994, and Mcluitgena

taurus Petuch, 1994, and large busyconids such as Bu-

sycon filosum Conrad, 1862, B. tritonis (Conrad, 1867),

B. pachijus Petuch, 1994, Sinistrofiilgur hollisieri Pe-

tuch, 1994, and S. grabaui Petucli, 1994

Etymology:The new taxon honors Mr. Richartl Duerr of

Hollywood, Florida, who collected the holol) pe in Qual-

ity Aggregates Phase 6 pit.

Calusacypraea sarasotaensis (Petuch, 1994)

(figures 5, 6)

Original DeM-ription (from Petuch, 1994:271, as Siph-

ocypraea sarasotaensis): "Shell small for genus, rounded,

very inflated and bulliform; base of shell rounded, mar-

gins not produced; aperture proportionally very wide

and open, becoming widest at anterior end; columellar

lip with 14 small, poorly-developed teeth; outer lip w ith

19 small teeth; posterior sulcus poorly-developed, open,

curving slightly to the left; extremities poorK -developed,

very rounded; fossula nonexistent."

Type material: Holotype, UF66226, length 40 mm, w idth

18mm.

Type Locality: Pinecrest Beds Fauna (Petuch Unit 3),

Tamiami Formation in APAC pit, Sarasota, Sarasota

County, Florida.

Remarks:

Although originally described as coming from Petuch

Unit 6, Calusacypraea sarasotaensis is now known to

occur much higher stratigraphically, in Petuch Unit 3.

The genus Calusacypraea becomes extinct after Unit 3

time, and C. sarasotaensis is the last-living member of

its genus. It is also the smallest Calusacypraea species,

averaging a length of only one-half that of the other

three congeners. Calusacypraea sarasotaensis most closely

resembles the much older, stratigraphically lower C.

duerri from Petuch Unit 10, but differs in having a

smaller and more rotund shell, fewer and coarser labial

and colimiellar teeth, and a better-developed and more

prominent posterior columellar extremity.

Colonies of C. sarasotaensis have been collected in

muddy-sand facies of Ihiit 3 along with characteristic

estuarine species such as Calophos nanus Petuch, 1994,

Echinofulgur lielenae Olsson, 1967, and Littorina lin-

dae Petuch, 1994.

Etymology: Named for Sarasota County, the general area

of the type locality.

Calusacypraea tequesta new species

(figures 7, 8)

Description: Basic morphology as for genus; shell very

inflated, iMilliform, large for genus; aperture proportion-

all\ wide, becoming wider and more open toward an-

terior end; columellar lip v\ ith 19 large, prominent teeth;

some columellar teeth bifurcate; outer lip with 22 large

and prominent teeth; base of shell rounded; margins

poorly -developed accept in area bordering anterior end,

where they become flattened and sharp-angled; apical

pit very deep; posterior columellar extremit\ large and

well-developed, thin and bladelike.

Type Material: Holotype, L'F68250. length 73 mm, w idth

47 mm; 1 paratype, length 75 mm. Graves Museum of

Archaeolog\ and Natural Histor\, Dania, Florida; 2

parat\ pes, lengths 75 mm and 72 mm, collection of the

author, Florida .Atlantic University.

Type Locality: l^inecrest Beds Fauna (Petuch Unit 4),

Tamiami Formation, in APA(^ pit, Sarasota, Sarasota

County.

Remarks: Calusacypraea tequesta is most similar, es-

pecialK in its inflated, bulliform shell shape, to C. duerri

E. J. Petuch. 1996

Page 21

from Petuch Unit 10. The new species differs from the

older, stratigraphically lower C. ducrri in being a larger

shell, in having fewer, larger, and coarser labial and

columellar teeth, and in having better-developed and

more flattened margins bordering the anterior end. Cal-

usacijpraea tequesta has the coarsest dentition of the four

known species of Calusacypraca. This new species is a

component of the rich estuarine fauna found in Petuch

Unit 4 in the Sarasota quarries ("Black Layer"), where

it is collected with Siphocypraea inetae Petuch, 1994,

Pijrazisintis scalimis (Olsson, 1967), P. lindae Petuch,

1994, Ccrithidea lindae Petuch, 1994, Btdia sarasoiaen-

sis Petuch, 1994, Mclongcna draperi Petuch, 1994, and

Echinofulgur jonesae Petuch. 1994.

Etymology: Named for the Tequesta Indian tribe of pre-

Columbian southern Florida.

ACKNOWLEDGMENTS

I thank Mr. Richard Duerr, Hollywood, Florida, for as-

sisting me in field collecting at Sarasota, Mrs. Cynthia

Mischler, Department of Geologv , Florida Atlantic I'ni-

versity, for t\ ping the manuscript, and Mr. Hugh Can-

non, President, Quality Aggregates, Inc., Sarasota, Flor-

ida, for allowing me to collect in the Phases 6 and 7 pits.

LITERATURE CITED

Heilpriii, .A. 1886. Explorations on the West Coast of Florida

and in tiie Okeecliobee Wilderness, with special reference

to the Geologv and Zoology of the Floridian Peninsula.

Transactions of the Wagner Free Institute of Science of

Philadelphia. l;.365-50(5. (often incorrectly dated as 1887).

Mansfield, W. C;. 1931. Some Tertiary Mollusks from Southern

P^lorida. Proceedings of the L'.S. National Museum of Nat-

ural History 79(21):1-12.

Olsson, A. A. and R. E. Petit. 1964. Some Neogene Mollusca

from Florida and the Carolinas. Bulletins of American

Paleontology 47(217):509-575.

Olsson, A. A. and R. E. Petit. 1968. Notes on Siphocypraeo-

Bulletins of American Paleontology 54(242): 279-289.

Parodiz, J. J. 1988. A New Species of Siphocypraea (Gas-

tropoda: Cvpraeidae) from the Neogene of Southwest

Florida. Annals of the Carnegie Museum 57(3):91-97.

Petuch, E. J. 1986. The Pliocene Reefs of Miami; Their Ge-

omorphological Significance in the Evolution of the At-

lantic Coastal Ridge, Southeastern Florida, U.S.A.. Journal

of Coastal Research 2(4):391-408.

Petuch, E. J. 1991. New Gastropods from the Plio-Pleistocene

of Southwestern Florida and the Everglades Basin. Special

Publication No. 1, W.H. Dall Paleontological Research

Center, Florida Atlantic University. 8.5pp.

Petuch. E. J. 1994. Atlas of Florida Fossil Shells (Pliocene and

Pleistocene Marine Gastropods). The Graves Museum of

.Archaeology and Natural History, Dania, Florida. 394 pp.

THE NAUTILUS 110(l):22-29, 1996

Page 22

Taxonomic Notes on South African Deep-Sea Conoidean

Gastropods (Gastropoda: Conoidea) described by

K.H. Barnard, 1963

Alexander V. Sysoev

Zoological Museum of Moscow State

Universit)

Hertzen st. 6

Moscow 113009, RUSSIA

ABSTRACT

Type specimens of ten species of deep-sea conoidean gastropods

from South Africa described by Barnard in 1963 are described

and illustrated. Lectotypes are designated for those species rep-

resented by syntype series. A new genus, Ahyssothauma, is

established for Moniliopsis pstlarosis Barnard, 1963.

Key words Cnnnidca. South ,\frica, deep-sea, t\pe specimens.

INTRODUCTION

In 1963, K. H. Barnard described a collection of South

African deep-sea mollusks obtained at lower bathyal to

upper abyssal depths. Most of the new species described

were conoidean gastropods. Unfortunately, the new spe-

cies were rather sketchily and often inadequately illus-

trated, which hampers their identification and compar-

ison with otiier deep-sea species. None of Barnard s spe-

cies of deep-sea conoidean gastropods were subsequentK

figured, and most were not subsequently mentioned by

other authors.

There are no reviews of the deep-sea conoidean gas-

tropod fauna of the Cape Point region, and Barnard's

work still remains the primary reference. The Cape Point

region is of particular interest due to its geographical

position, because it is situated at the border between two

oceans, and its deep-sea fauna may represent a mixture

of Atlantic and Indo-Pacific elements.

'ihrough the courtesy of Mrs. Michelle G. van der

Merwe from the South Alricaii Museum, (Jape Town

(SAM), I was able to examine the type specimens of

Barnard's species of deep-sea conoidean gastropods. The

purpose of the present paper is to illustrate these species

and to revise their taxonomic position

In his pul)licution, Barnard (1963) designated sotiie

specimens from his material as "types ". Sometimes, when

a species was represented by a few specimens, he did

not mention the presence of types at all. In these cases,

all such specimens are considered syntypes.

Specimens of tour of Barnard s species were sent by

him to the British Museum (Natural History) as "para-

types ". However, the designation of paratypes without

a designation of holotype is invalid. Moreover, almost all

these specimens are not even syntypes, because they were

not originally included into "types" (= type series) (ICZN

Art. 27(b)VI).

SYSTEMATICS

Superfamily CONOIDEA Fleming, 1822

Familv TURRIDAE H. & A. Adams, 1853

SubfamiK Tl'RRINAE H. & A. Adams, 1853

Cemmiila (Ptijchosijrinx) lobatopsis (Barnard. 1963)

(figures 1-3)

Tiirris lolxila (Sowerb) ) [pdiiuii] Barnard, 19.58:107, fig, 3j.

Clavatulci lobatopsis Barnard, 1963:420-421, fig. 2a.

Ccmnuila (Ptychosyrinx) lobatopsis (Barnard) Kilburn, 1983:

.582.

Material examined: Three specimens (SAM A9730) in-

dicated by Barnard (1963) as "Types".

Type locality: "Africana 11 ', sta. A191, 33°36'S, 16°15'E,

1520-1570 fms (2780-2918 m).

Description: Two of the three syntypes are heavily cor-

roded, though the presence of opercula indicates that

thev were collected alive. The best preserved specimen

is here designated as lectoty pe (figs. 1-2). The dimensions

are: lectoty pe — 32.4 x 13.6 mm at 6.5 preser\ed whorls;

paralectotypes — 34.6 x 12.9 mm and 33.4 x 13.7 mm

at 5,5 and 6+ [)reserveil whorls, respecti\el\ . The upper

w horls are eroded and partly missing in all shells Lec-

totype strongly angulate at periphery. Suture slightK

wavy, clear, despite Barnard's statement, visible on all

whorls. Subsutural cord thin, narrow, less distinct on body

whorl, with small tubercles at points of intersection with

growth lines passing through peri|)heral gennnnles The.se

growtli lines strong, raised, especialK on body whorl.

A. V. Svsoev, 1996

Page 23

Figures 1-10. 1-3. Gf »U7!)(/(j (P/ yr/ias i/rin.v) /otefopsis, lectotype ( 1,2) and paralectotype, 33.4 mm (3). 4-5,9-10. Abi/ssof/iauma

psilarosis, lectot>pe (4. 5) and paralectot\ pe (9. 10). 6. Cymnobela (Theta) chrysopelex. holot>pe. 7. 8. Typhlonmngelia polythele,

lectotype (7) and paralectotype (8).

Four weak, narrow, widely spaced cords between sub-

sutural cord and row of peripheral gemmules, 3 distinct

cords below gemmuiate periphery on last whorls. Shell

base and canal covered by 17 narrow, widely spaced

cords, which gradually weaken towards the canal end.

Peripheral gemmules on spire whorls mostly eroded, but

preserved ones strong and acute, becoming progressively

weaker, longitudinally elongate, arcuate towards the body

whorl. They are numerous, irregularly placed on last half

of the body whorl, almost obsolete near aperture. Pen-

ultimate whorl bears 20 gemmules, the body about 33.

Paralectotypes have less angulate whorls with less

prominent peripheral gemmules. Subsutural ramp may

be concave, especially on last half of the body whorl.

Page 24

THE NAUTILUS, Vol. 110, No. 1

Siphonal canal varies in length and nia\ be almost straight

or turned to left, as in iectot\ pe. The pattern of sculpture

is not similar to that shown in Barnard s drawing (Bar-

nard, 1963:£ig. 2a), i.e., with very strong and closely set

bilobed tubercles forming the subsutural cord.

Remarks: This species was also recorded in the area from

33°26'S to 34°37'S and from 16°23.5' E to 17°03'E, at

2378-3036 m (Barnard, 1963). The shells found in the

John Murray E.xpedition, sta. 118, off Zanzibar, 1789 m,

and identified as G. teschi (Powell, 1964) (Sysoev, in

press, figs. 68-70), are G. lobatopsis.

Family CONIDAE Fleming, 1822

Subfamily CLATHURELLINAE H. & A. Adams, 1858

Typhlomangelia polijthele Barnard, 1963

(figures 7-8)

Typhlomangelia (?) polylhele Barnard, 196.'3:422, fig. .Se, i.

Material examined: Two s\ ntypes (SAM A9802).

Type locality: "Africana 11", sta. A317, 33°50'S, 16''30'E,

1480-1660 fms (2707-3036 m).

Description: Largest (12.3 x 5.4 mm) and better pre-

served of two syntypes here designated as lectotype. Its

shell rather thick, consisting of 5,5 remaining whorls.

Protoconch missing, first teleoconch whorl eroded. Whorls

angulate at periphery. Subsutural ramp slightly concave,

with narrow subsutural cord and numerous, regular, nar-

row, arcuate folds formed by thickened growth lines.

Axial sculpture represented by numerous, very short ribs

below subsutural ramp. Ribs rapidly weaken, do not reach

lower suture or peripherv of body whorl. Ribs number

about 22 on penultimate whorl (its dorsum broken, ap-

parently during extraction of soft body) and 26 on body.

Spiral cords (7 on penultimate whorl and 32 on body

whorl) are narrow, separated by wide interspaces (2-4

times wider than cords). Spiral cords are more closely

set on siphonal canal. Canal short, straight. Svmmetrical

anal sinus occupies entire subsutural ramp, being deepest

in its middle. Paralectotype (11.5 x 5. l mm at 5+ pre-

served whorls) has the same conchological characters.

Remarks: Barnard provisionalK placed this species in

Typhlomangelia. \\ hich seems to be appropriate for the

species, despite its small and broad shell lacking spiral

sculpture on the subsutural ramp. Typhlomangelia ad-

enica Sysoev, a very similar species both conchologically

and in radular characters, is known from the Gulf of

Aden, John Murray Expedition sta. 26 and 185, 200-

2312 m (Svsoev, in press).

Abyssothauma Sysoev, 1996, new genus

Type species: Moniliopsis psilarosis Barnard, 1963.

Diagnosis: Shell large, to 60 mm, lusiiorm, w ith rounded

and vseakly convex whorls, covered b\ thick olivaceous

periostracum. Spire more than 30% of shell height. Earlv'

v\li()rLs wcaklv angulate. Axial sculpture of numerous

oblique narrow folds below shoulder in early whorls, or

absent. Spiral sculpture of low , rather wide cords belov\

subsutural ramp. Shell base weakly convex, not diff^er-

entiated from canal. Aperture narrowly oval, siphonal

canal wide, long. Anal sinus moderately deep, rounded,

occupv ing the subsutural ramp, with deepest point in its

middle. Operculum large, leaf-shaped, with terminal nu-

cleus. Radular teeth hollow, long, slightK curved, not

barbed, with rounded base and terminal opening of tooth

cavity.

Remarks: The new genus resembles the genera Typhlo-

syrinx Thiele, 1925 and Belaturricula Powell, 1951. From

the former genus it differs in having a stronger shell with

better developed spiral sculpture, thick dark-colored

periostracum, different form of axial sculpture on early

whorls, and presence of operculum. This new genus dif-

fers from Belaturricula in having a narrower shell with

stronger sculpture, pronounced siphonal canal and deep

anal sinus. The new genus includes the type-species and

a new species from the Tasman Sea, 3610-3688 m that

will be described elsewhere. Pontiothauma ergata Hed-

ley, 1916, resembles A. psilarosis in general shell outline,

predominance of spiral sculpture, structure of radular

teeth (see Egorova, 1982, figs. 63a. b), presence of oper-

culum with terminal nucleus, but differs in having a

shallow anal sinus (holotype illustrated b\ Dell, 1990:

figs. 418-419) and very thin light-colored periostracum.

The taxonomic position of P. ergata is uncertain, but it

obviously does not belong to Pontiothauma, differing in

the shell form, sculpture, the presence of operculum and

different radular teeth indicating the placement in the

Clathurellinae, not Raphitominae. Probably P. ergata

belongs to a separate genus, closely related to Abysso-

thauma.

Abyssothauma psilarosis (Barnard, 19(i3)

(figures 4-5, 9-10)

Moniliopsis psilarosis Barnard, 1963:421-422, fig. 4a.

IVIaterial examined: Two specimens (SAM A9789) in-

dicated by Barnard (1963) as "Types".

Type locality: Barnard (1963) did not designate the type

localitv nor indicate which of his two samples contained

'tvpes '. However, judging from the indicated number

and cjualitv of specimens from '.\fricana 11 sta. A316,

34°42'S, 16°54'E, 1725-1780 fms (3155-3255 m), cor-

responding to specimens from the type series, the t\pe

localitv is restricted to these coordinates.

Description: Largest of two tvpe specimens is here des-

ignated as lectotvpe. Dimensions: 47.2 x 16.0 at 6.5

preserved whorls (lectotype) and 40.8 x 14.4 at 6 pre-

served whorls (paralectotype). Lectotype shell strong,

solid, covered by thick dark-olivaceous periostracum.

Protoconch and at least one teleoconch whorl missing.

Upper whorls distinctiv angulate above peripherv, an-

gulation weakening in subsequent whorls, becoming ob-

-solete in penultimate whorl. Subsutural ramp weakly

A. V. Sysoev, 1996

Page 25

concave. Upper whorls with strong, obhque axial ribs

below subsutural ramp U8 ribs on first well-preserved

whorl). Ribs form small knobs in upper part. Ribs mark-

edly weaken towards lower suture, becoming shorter in

successive whorls. Last 3.5 whorls devoid of a.\ial sculp-

ture e.xcept for indistinct, irregular folds formed b> growth

lines. Spiral sculpture of initial w horls consists of wide,

very low cords separated by narrow grooves. Cords be-

come stronger and more widely spaced on two last whorls,

but interspaces do not exceed cord width. Subsutural

ramp lacks spiral sculpture except of 3-4 feeble striae

on lower part. Aperture narrow, inner lip evenly curved,

covered by a smooth callus. Siphonal canal moderately

long, narrow. Paralectotype smaller, with only 2.5 whorls,

lacking axial ribs. Spiral cords slightly weaker than those

of lectoty pe. Outer lip intact, evenly projecting forward

below anal sinus. Sinus rounded, moderately deep, slight-

ly asymmetrical, with its deepest point just above middle

of subsutural ramp.

Remarks: Barnard (1963) placed his species in Moni-

liopsis Conrad, 1865. However, this Eocene genus con-

taining few species from southern USA and Peru, is char-

acterized by a strong sculpture of very wide spiral cords

cut into subrectangular nodes that are present on the

subsutural ramp. In Barnard's time, Moniliopsis was used

to include some Recent West American species which

are now included in the genus Ophioclennella Bartsch,

1944. This genus, in turn, cannot accomodate the Bar-

nard's species either, because it is characterized by a

much lower position and different type of anal sinus.

Subfamily RAPHITOMINAE Bellardi, 1875

Gymnobcla glaucocreas (Barnard, 1963)

(figures 15, 19)

Cijthara (.'■") glaucocreas Barnard, 1963:422-42.3, fig, 2cl.

Material examined: One specimen (SAM A9821) indi-

cated by Barnard (1963) as "Type ' (holotype).

Type locality: "Africana 11", sta. A318, 33°52'S, 16°51'E,

1380-1520 fms (2524-2780 m).

Description: Shell 26 0 x 11.8 mm, with 7.5 preserved

whorls, greyish-white, covered by thin periostracum.

Protoconch whorls eroded, probabK partK missing. Ear-

ly teleoconch whorls with only a slight angulation at

periphery. Towards body whorl, whorls become more

angulate, subsutural ramp becomes concave. Sutures

shallowl) channeled. Axial ribs begin in lower part of

subsutural ramp, numerous (27 on body and penultimate

whorls), narrow, oblique, with sharp crests. Two ribs may

rarely merge together near whorl shoulder. Subsutural

ramp covered by thin arcuate folds following traces of

anal sinus. Folds are not always continuations of axial

ribs, and are more numerous than latter (41 on bod\

whorl, plus several similar short folds that do not reach

lower part of subsutural ramp). Spiral sculpture of nar-

row, flattened, widely spaced cords covering entire shell

surface. Cords on subsutural ramp more closely spaced.

Interspaces between cords bear thinner secondary thread.

Remarks: This species has all the characteristics of Gym-

nobcla \errill, 1884. It is most similar to the North At-

lantic G. engonia N'errill, 1884 (t\ pe species of the genus)

and differs from the latter mainly in having less angulate

whorls, less oblique axial ribs, and fainter spiral sculpture.

The radula of G. glaucocreas. described by Barnard as

similar to that of G. verecunda (Barnard, 1963), also has

much in common with that of G. engonia (see Bouchet

& Waren, 1980, fig. 26).

Gymnobcla dagama (Barnard, 1963)

(figures 11-14)

Cythara ('.■') dagama Barnard, 1963:423, fig. 2g, h.

Material examined: Four specimens (SAM A9731) out

of 5 indicated by Barnard as "Types ',

Type locality: "Africana 11 ", sta. A191, 33°36'S, 16°15'E,

1520-1570 fms (2780-2871 m).

Description: Largest synt\pe (38.4 x 16.3 mm) with

heavily eroded surface. Second largest specimen (32.8 x

13,8 mm) with well preserved shell (fig. 12) is here des-

ignated as lectotype. Shell of 6.5 whorls. Protoconch miss-

ing, upper whorls eroded. Sutures deeply channeled.

Whorls almost evenly convex, with only slight angulation

at about 1/4 of whorl height, Subsutural ramp narrow,

slightK concave. Axial ribs weak, narrow, with rounded

or flattened crests, numerous (19 on penultimate whorl

and 27 on bod\ whorl) and somewhat irregular in strength,

especialK' on bod\ whorl. Ribs begin at border of sub-

sutural ramp and extend to lower shell base, Subsutural

ramp devoid of sculpture. Spiral sculpture on spire whorls

appears to consist of wide, flattened cords separated by

narrow grooves due to the erosion of surface. Cords well

preserved and are narrow and widely spaced (interspaces

1.5-2 times wider than cord) only in some shell areas,

particularly on body whorl. Aperture oval, with short,

wide siphonal canal poorly difterentiated from aperture.

Judging from growth lines, outer lip very slightK projects

forward below anal sinus. Anal sinus narrow, with its

deepest point at middle of subsutural ramp. Two smaller

paralectot\pes (31.3 x 1,3.1 mm and 23.6 x 11.5 mm

at 7-1- and 6-1- whorls, respectively) have more angulate

whorls with more concave subsutural ramp, especially

in smallest, best preserved specimen. This specimen has

weak spiral cords on lower part of subsutural ramp, cov-

ered In numerous thin arcuate folds formed by thickened

growth lines.

Remarks: The radula (Barnard, 1963, fig. 2h) is not char-

acteristic of Gymnobcla and more closeK resembles that

of Bcnthomangclia Thiele, 1925. However, the concho-

logical characters indicate the assignment of this species

to Gymnobcla. Besides the t\pe locality, the species was

found in the area from 33°36' to 34°37'S and from 16°15'

to 17°03'E, at 2707-3219 m (Barnard, 1963).

Page 26

THE NAUTILUS, Vol. 110, No. 1

Figures 11-20. \l-l'i. Cymnobcla dagama, Iw-totvpo (12) ami paraU-dotypes ( 1 1, 13, 14; .38.4, ;31.;3, ami 2.3,6 mm, respectively).

1.5. 19. Gyrmwhcla glaucocrcas. hololype. 16-1«, 20. Xanlhodaphnc pyrropelex, lectotype (17-18) and paralectotypes {16. 20;

49.3 and 44 7 mm, respectively).

A. V. Sysoev, 1996

Page 27

Gi/tnnobcla verccunda (Barnard, 1963)

(figures 21-24)

Daphnella {?} verecunda Barnard, 1963:427-428, fig. 2e.

Material examined: Three syntypes (SAM A9697).

Type locality: "Africana 11". sta. A189, 33°50'S, 17°21'E,

600 1ms (1097 m).

Description: Largest (shell 22.4 x 10.8 mm) ol three

syntypes best preserved, and here designated as lectotype

(figs. 21-22). Sliell white, consisting of 7 preserved whorls

(uppermost broken). Protoconch missing. Spire whorls

angulate at periphery, subsutural ramp slightly concave.

Axial ribs begin in lower part of subsutural ramp, very

rapidly become prominent at shoulder, and gradually

weaken below, not reaching canal. Ribs strong, oblique,

narrow, 15 ribs on penultimate whorl, 16 on body whorl.

Some growth lines on subsutural ramp thickened near

upper suture. Spiral sculpture on subsutural ramp con-

sisting of weak, narrow threads, which appear at some

distance from upper suture and become stronger towards

shoulder. Spiral sculpture on subsutural ramp not always

visible due to shell erosion. Spiral cords below the shoul-

der rather wide, flattened, sometimes with a narrower

secondary cord in interspaces. Siphonal canal short,

slightly twisted. Anal sinus, judging from growth lines,

is asymmetrical, with its deepest point in lower half of

subsutural ramp. Paralectotypes are 19.5 x 9.5 and 16.6

X 8.3 mm in size. Larger paralectotype better preserved,

with 15 axial ribs on body whorl and wider spiral cords

than in lectotype. All three shells have a characteristic

light-orange staining on columella.

Remarks: Besides the type localitv , the species was also

found off northern Namibia (19°45'S), at 1600 m, R/V

"Professor Shtokman", sta. 1563 (Sysoev, unpublished

data).

Gijmnohela (Theta) chrysopelex (Barnard, 1963)

(figure 6)

Typhlusyrinx chrysopelex Barnard, 1963:425, fig, 3g, h.

Material examined: One specimen (SAM A9857) (ho-

lotype).

Type locality: "Africana 11", sta. A322, 34°36'S, 17°00'E,

1500-1760 fms (2743-3219 m).

Description: Shell 18.8 x 8.6 mm, white, with glossy

surface. Protoconch of three preserved whorls (at least

1-1.5 initial whorls missing), diameter 1.25 mm. Pro-

toconch sculpture not well preserved, consisting of ar-

cuate axial riblets passing from suture to suture. Riblets

on lower half of whorl diagonalK crossed by oblique

riblets begiiming at lower suture. Teleoconch of 4.75

whorls. First whorl angulate slightly above periphery,

penultimate whorl at about 1/3 of whorl height. Oblique

axial ribs beginning just above whorl angulation, slightly

weaken below, and reaching lower suture and upper shell

base. Ribs 15 on penultimate whorl, 18 on body whorl.

Spiral sculpture of obscure striations below whorl an-

gulation, feeble threads on lower part of shell base and

canal. Based on growth lines, anal sinus very shallow as

compared to congeners.

Remarks: This species agrees well with the subgenus

Theta Clarke, 1959 in the shell outline and sculpture.

The protoconch is very similar to that of G. (T.) chariessa

(Watson, 1881) (see Bouchet & Waren, 1980: figs. 254,

255), but larger than in that species. The radular teeth

(Barnard, 1963, fig. 3h) are typical of Gymnobela. Gijm-

nohela (T.) chrysopelex is closely related to G. (T.) char-

iessa. but differs in having a shallower anal sinus, larger

protoconch, and less obliciue axial ribs.

Xanthodaphne pyrropelex (Barnard, 1963)

(figures 16-18, 20)

Pletirotoma (Surcida)dissimili.s (nun Watson) — Barnard, 1958:

147-148, fig. 23a,

Typhlusyrinx pyrropelex Barnard. 1963:423-425, fig, 2c.

Material examined: Four specimens (SAM A1643, SAM

A1644, SAM A1645)out of 5 indicated by Barnard (1963)

as "Types".

Type locality: Barnard did not designate the type lo-

cality, and the type series came from three different

localities described as "Cape Point N. 77°E. 650-700

fathoms; NE. x E. 1/2 E. 43 miles, 900 fathoms; NE.

X E 3/4 E. 38 miles, 750-800 fathoms". The syntypes

were collected by the Cape Goverment trawler s.s. "Pie-

ter Faure , but the respective station numbers were not

indicated in the publications (Barnard, 1958, 1963) or

on labels. Assuming that Barnard (1963: 424) listed lo-

calities and museum numbers in the same order, the

lectotype was found in the second of above-mentioned

localities.

Description: Largest syntype (SAM A1643, 49.3 x 18.5

mm at 7 teleoconch, 1.5 preserved protoconch whorls)

with slightly broken canal, eroded parts of body whorl.

Another syntype (SAM A1645, 45.0 x 15.3 mm) strongly

eroded, with broken aperture. Largest of two remaining

syntypes designated here as lectotype (figs. 17-18). Shell

48.2 X 17.2 mm, white, thin, fragile. Protoconch of 1.5

preserved brown whorls w ith thin oblique arcuate axial

riblets. Ribs more crowded and irregular near the tran-

sition to teleoconch, less inclined, occasionally bifurcat-

ing in lower part. Teleoconch whorls (6.8 in number)

almost evenl\- rounded, with well-marked subsutural

ramp, slightK concave in earl> whorls, almost flat in last

ones. Border of subsutural ramp defined by shallow

groove. Some growth lines on subsutural ramp thickened

below suture, which is more pronounced in early whorls.

Subsutural ramp devoid of spiral sculpture except for

indistinct striation on its lower part. Spiral sculpture be-

low shoulder consisting of very low, flattened, wide,

slightly wavy cords with serrated edges, separated by

narrow grooves. Cords becoming stronger, narrower be-

low bod\' whorl periphery. Based on growth lines, the

Page 28

THE NAUTILUS, Vol. 110, No. 1

Figures 21-29. 21-2J-. C.ymmjhcki icmiinda. Ii-etot) pi' (21-22) and paralectotypes (23, 24-, 195 and 16.6 mm. respectively).

25-27. XaulIuHlaphnr aubruscu. lectotype (25-26) and paraleetolvpe (27). 28-29. Fanwiica hilntdis. lectotvpe (28) and par-

alectotype (29).

anal sinus subsutural, wide, deep. Its margin almost ver-

tical below suture, nearly horizontal at border of sub-

sutural ramp. Outer lip very strongly projecting forward

below sinus. Paralectotypes very similar to lectotype,

varying insignificantly in details of spiral sculpture. None

have more intact protoconch than on lectotype.

Remarks: The species has all the essential characters of

XanlluHlapInw Powell, 1942. Protoconch sculpture is

sinular to that oi the type species oi the genus, Plcuro-

toma membranacea Watson, 1886. However, the shell

of X. pyrropclcx is more slender than that of most other

repre.sentati\es of the genus. Xanthodaphne pyrropclcx

A. V. Sysoev, 1996

Page 29

species was also recorded in the area from 33°36'S to

34°36'S and from 16°15'E to 17°00'E, at 2524-3219 m

(Barnard, 1963).

Xanthodaphne suhrosea (Barnard. 1963)

(figures 25-27)

Typhlosijrinx subrosea Barnard. 1963:425-426 , fig.3a-d.

Material examined: Two syntypes (SAM A9822).

Type locality: "Africana II", sta. A318, 33°52'S, 16°51'E,

1380-1520 fms (2524-2780 m).

Description: Larger of two syntypes (36.0 x 16.2 mm)

designated here as lectotype. Shell white, semi-transpar-

ent, fragile. Protoconch of at least 3.5 brown whorls

(upper whorls eroded) sculptured b\ numerous arcuate

a.xial riblets on upper half, diagonally cancellated on

lower half. Protoconch diameter 1.45 mm. Teleoconch

of 4.75 evenly convex, rapidh' expanding whorls. Whorls

smooth initially, then obscure spiral striations appear,

turning into rather strong and flattened cords (about 60

on body whorl) co\ering the entire whorl except for

narrow area just below upper suture. Strongest, most

widely spaced cords situated on whorl shoulder. These

cords appear first on shell. Interspaces between cords

usually do not exceed the cord width at whorl periphery.

Siphonal canal rather long, slightly twisted. Inner lip

without callus. Based on growth lines, anal sinus subsu-

tural, outer lip greatK projecting forward below sinus.

Paralectotype with much smaller, proportionally broader

shell (20.3 X 11.0 mm), its protoconch consisting of about

4.5 whorls. Upper whorls eroded, others with sculpture

similar to that of lectotype. Protoconch diameter 1.35

mm. Spiral sculpture similar to that of respective whorls

of lectotype. Sculpture irregular on most of body whorl,

due to scar of heavy damage to shell. Inner lip covered

by very thin callus.

Remarks: The species corresponds well to the diagnosis

of Xanthodaphne, and is quite similar to its type-species,

A', memhranacea. The only difference is that the pro-

toconch sculpture in X. memhranacea is mainly axial.

However, the protoconch sculpture of X. subrosea falls

within the range of variability characteristic of the genus

in general (see Bouchet & Waren, 1980: figs. 234-237).

Famelica bitrudis (Barnard, 1963)

(figures 28-29)

Daphnella (.^) bitrudis Barnard, 1963: 428. fig. lb.

Material examined: Two s\nt\pes (SAM A9754).

Type locality: "Africana 11", sta. A193, 33°49'S, 16°30'E,

1500 fms (2743 m).

Description: Larger s>nt\pe (15.2 x 3.4 mm) with bro-

ken aperture, partly broken penultimate whorl. Smaller

synt\pe (13.6 x 3.3 mm) here designated as lectotype.

Shell very narrow, semi-transparent, white, very fragile,

consisting of 7.5 whorls, uppermost vvhorl(s) missing. Up-

per 3 w horls with eroded surface, differing in their pro-

portions from subsecjuent whorls, apparentK' represent-

ing a protoconch. Teleoconch w horls rather evenly con-

vex, with clearly defined subsutural ramp, bordered by

spiral cord corresponding to lower, almost horizontal

branch of anal sinus. Spiral sculpture developed below

subsutural ramp, consisting of weak narrow cords sep-

arated by wide (2-3 times wider than cords) interspaces.

Cords weaken on shell base, with smooth band at tran-

sition of shell base to canal. Sculpture resuming on canal,

but with very weak cords. Siphonal canal long, narrow,

twisted. Anal sinus subsutural, not very broad. Outer lip

projects forward below anal sinus. Paralectotype differs

onl\ in having slightly more numerous and more closely

spaced spiral cords (12 on the penultimate whorl vs. 11

in the lectotype).

Remarks: In their original description of Famelica,

Bouchet and Waren (1980), the authors included Daph-

nella bitrudis in their new genus.

ACKNOWLEDGEMENTS

The author is greatly indebted to Mrs. Michelle G. van

der Merwe, Collection Manager, South African Museum,

for the loan of the material.

LITERATURE CITED

Barnard, K. H. 1958 Contributionsto the knowledge of South

African marine Moliusca. Part I. Gastropoda: Prosobran-

chiata: Toxoglossa. Annals of the South .African Museum

44:73-163,

Barnard, K. H. 1963, Deep sea Moliusca from west of Cape

Point, South Africa. Annals of the South African Museum

46:407-452.

Bouchet, P. and A. Waren. 1980. Revision of the North-East

Atlantic bathyal and abyssal Turridae (Moliusca, Gastrop-

oda), Journal of Molluscan Studies, supplement 8:1-119,

Dell, R, K, 1990, Antarctic Moliusca: with special reference

to the fauna of the Ross Sea, Bulletin of the Royal Societ\

of New Zealand 27:1-311,

Egorova, E. N. 1982. Biological results of the Soviet Antarctic

expeditions, 7. MoUusks of the Davis Sea. Issledovaniya

fauny morei 26(34):3-142 (in Russian).

Kilburn, R. N. 1983. Turridae (Moliusca: Gastropoda) of

southern Africa and Mozambique, Part 1, SubfamiK Tur-

rinae. Annals of the Natal Museum 25(2):549-585.

Sysoev, A. \'. 1996. Deep-sea conoidean gastropods collected

b\ the Jcihn Murra\ Expedition, 1933-34, Bulletin of the

Natural History Museum, London (Zoology), in press.

THE NAUTILUS 110(1):30, 1996 Page 30

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THEt^NAUTILUS

CONTENTS

Volume 110, Number 2

Fehmanj 19, 1997

ISSN 0028-1344

Dan C. Marelli

Maureen K. Krause

William S. Arnold

William G. L^•ons

Systematic Relationsliips Among Florida Populations of

Aroopcctcn irmdidiis (Lamarck. 1819) (Bi\al\ia: Fectinidae) 31

Subspecific Status oi Araopccten irradiaiis coiiroithcns

(Say, 1S22) and of the Bay Scallops of Florida 42

Dan C. Marelli

William G. Lyons

William S. Arnold

Maureen K. Krause

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THE NAUTILUS 110(2):31-41, 1997

Page 31

Systematic Relationships Among Florida Populations of

Argopecten irradians (Lamarck, 1819) (Bivalvia: Pectinidae)

Dan C. Marelli

Maureen K. Krause' -

William S. Arnold

William G. Lyons

Florida Department of

Environmental Protection

Florida Marine Research Institute

100 8th Avenue SE

St. Petersburg, FL 33701-5095 USA

ABSTRACT

Morphometric and genetic examinations (using allozyme elec-

trophoresis) were conducted on two Florida populations of bay

scallops, Argopecten irradians (Lamarck, 1819), to investigate

the status of the subspecies A. i. taijlorae Petuch, 1987. One

other Florida population [A. i. coiicentricus (Say, 1822)] was

examined morphometrically. Morphometric examinations em-

phasized mensural and meristic characters used in previous

systematic diagnoses, Morphometric data were analyzed using

anal) sis of variance and principal component analysis. Scallops

taken from Florida Bay (putatively A. i. taijlorae Petuch, 1987)

were smaller but otherwise not morphologically distinct from

populations of A. i. concentricus (Say, 1822) from Pine Island

Sound and Homosassa Bay, Florida. Mean Nei s modified ge-

netic distance value shows a close relationship between Florida

Bay and Homosassa Ba\ scallops. Neither morphometric nor

genetic e\idence supports the proposed status of A. i. taijlorae

as distinct from Florida populations of A. i. concentricus.

Key words: Pectinidae, Argopecten irradians. morphometries,

svstematics.

INTRODUCTION

Scallops are often distributed in patchy or contagious

patterns, and, although disjunct local populations may

be joined by variable degrees of lar\ai exchange to form

metapopulations (Levins, 1970; Andrewartha k Birch,

1984; Roughgarden et ai, 1985; Roughgarden & Iwasa,

1986; Simberloff, 1988; Hanski, 1989; Orensanz et a!.,

1991), local populations along a species' range are prob-

ably self-sustaining (Sinclair ct ai. 1985). ExpectedK',

' Department of Ecology and Evolution, State University of

New York at Stony Brook, Stony Brook, NY 1179-4-5245 USA.

^ Current address: Southampton College, Long Island Univer-

sity, Southampton, NY 11968 USA.

reproductive isolation may contribute to genetic drift

that can produce locally distinctive morphologies. The

bay scallop, Argopecten irradians (Lamarck, 1819), which

inhabits semi-enclosed coastal bays, sounds, and estuaries

from Cape Cod, Vlassachusetts, to Tampico, Mexico

(Clarke, 1965; Abbott, 1974), exemplifies the results of

such isolation (Clarke, 1965; Waller, 1969; Kraeuter et

ai, 1984).

Three extant subspecies ot A. irradians have custom-

arily been recognized: Argopecten irradians irradians

(Lamarck, 1819), distributed from Cape Cod to New

Jersey; A. i. concentricus (Sa\-, 1822), distributed from

New Jersey to South Carolina and from Palm Beach Inlet,

Florida, to Louisiana; and A. i. aniplicostatus (Dall, 1898),

distributed from Texas into Mexico (Clarke, 1965; Wal-

ler, 1969, 1991; Abbott, 1974). A fourth subspecies, A. i.

sablensis (Clarke, 1965), described from shells collected

in Nova Scotia, apparently became extinct during the

Holocene (Waller, 1969). These subspecies have been

distinguished morphologically by plical number and

shape, valve size and shape, and valve coloration (Clarke,

1965; Waller, 1969; Abbott, 1974). Bricelj et at. (1987),

however, cautioned that subspecific distinctions within

A. irradians have not yet been substantiated by bio-

chemical techniques.

Recently Petuch (1987) described an additional sub-

species of ba\' scallop, Argopecten irradians taijlorae,

based upon specimens from Rabbit Key Basin, Florida,

and reported it to be restricted to Florida Bay and the

western sides of the middle and upper Florida Keys

(figure 1). Petuch (1987) reported that plicae of A. i.

taylorae were more numerous than those of the other

subspecies and that plicae were lower, wider, and more

flattened than in A. i. concentricus. The valves of A. i.

taylorae were also said to be much smaller, more fragile,

and much more colorful than those of A. i. concentricus

(table 1).

To emphasize the geographic isolation of Argopecten

irradians taylorae, Petuch (1987) gave the distributional

Page 32

THE NAUTILUS, Vol. 110, No. 2

ST. JOSEPH BAY

GULF OF

MEXICO

MIAMI

Figure 1. Florida: (.■ollt'ttiiig localions ot Argiipcctrn irrcididiis piip\ilati(ins ust'tl in aiial\st>s and additional localities relevant to

bay scallop populations. Inset shows location of collection uilliin Hahhil Ke\ Dasin.

range of .A. i. concent ricus as "New Jerse\ to Georgia,

and. . .Tampa, Florida to Louisiana." In iloing so, lie

overlooked earlier Florida records of bay scallops at Boca

Grande and Sanibel Island (Clarke, 1965); at Whitewater

Bay, Sandy Ke) Basin, and south of I'laniiiigo in Florida

Bay (Tabb & Manning, 1961); and at several locations

from Vaca Key in the middle Keys northward through

Biseayne Bay to Palm Beach Inlet on the Florida east

coast (Waller, 1969). Clarke (1965) and Waller (1969)

had pre\iousl\ reported specimens from all of these lo-

cations as A. i. concent ricus. but at least some of the

specimens came from within the range of A. i. tatjlorae.

An investigation of the s\ stematic status of Argopecten

irradians taylorae was prompted b\ the need to address

resource management issues. Ba\ scallops have histori-

cally supported small-scale commercial fisheries at var-

ious locations between Pensacola Ba\ and Pine Island

Souiul along Florida s west coast. .Although the com-

mercial fishery has decreased substantially in recent years,

the species continues to be the target of an important

recreational fisherv throughout much of that region (Ar-

nold, 1990), The presence of an isolated subspecies of

ba> scallop in Florida Ba\ , as reported b\ Petuch (1987),

could have ramifications for resource management. The

Dan C. Marelli et a I., 1997

Page 33

Table 1. Characters of subspecies of Argopeclen irradians reported by previous authors.

Subspecies

Clarke, 19te

Waller, 1969

Abbott, 1974

Petuch, 1987

Argopecten irradians

irradians

Argopecten irradians

concenlricus

Argopecten irradians

amplicostatus

Argopecten irradians

taylorae

16-20 (usuall) 16-lS

ribs; ribs rounded;

valve length and

height nearl) equal,

L/H ratio approx.

1.06; W/H ratio ap-

pro.\. 0.21 (left) and

0.24 (right); right

valve color as in Ab-

bott (1974) but com-

monly white (or

nearly so),

17-23 ribs; ribs round-

ed or with flattened

tops; L/H ratio ap-

prox. 1.03 (1) and

1.02 (r); valves some-

what inflated, W/H

ratio approx. 0.26 (1)

and 0.32 (r); 75-

100'^ of specimens

with v\ hite or nearly

white right valves,

color otherwise like

left valve except

lighter.

12-18 (usually 14-16)

ribs; ribs rounded to

slightly flattened;

L/H ratio approx.

1.0(1) and 1.01 (r);

shell more inflated

than other 2 subspe-

cies, W/H ratio ap-

prox. 0.29 (1) and

0.34 (r); color of

right valves usually

white, sometimes

with slight color

tinge.

15-20 ribs; ribs low and 17-18 ribs; ribs low and

rounded; L = H;

valves thinner and

flatter and ribs lower

than A. i. concentri-

cus.

15-22 ribs; ribs high,

sharplv rounded, be-

coming semi-hexago-

nal distally; differs

from A. i. irradians

by having thicker

and more convex

valves and higher

ribs; differs from A. i

amplicostatus by

having thinner and

less convex valves.

13-17 ribs; ribs high,

sharply rounded

proximally, becoming

low and trapezoidal

to semi-hexagonal

distally; differs from

A. I. concentricus by

having thicker, more

convex valves with

fewer ribs.

roundish; valves most

compressed of the 3

subspecies; right

valve color only

slightlv lighter than

left.

19-21 ribs; ribs squar-

ish; right valve much

more convex than

left; right valve light-

est in color of the

subspecies, common-

Iv all white.

12-17 ribs; right valve

with high, squarish

ribs; shell more gib-

bose than other sub-

species; right valve

commonly white.

23-25 ribs; ribs flatter

and wider when

compared with A. i.

concentricus. shell

smaller, more fragile

and more colorful

than A. i. concentri-

cus; right valve gen-

erally yellow rather

than white, with

more brown mot-

tlings than A. i. con-

centricus.

population might require separate management because

of the increasingly popular recreational fishery or might

require protection as a rare and isolated subspecies.

We used both allozyme electrophoresis and morpho-

metric analyses to investigate the relationship of the Flor-

ida Bay population of Argopecten irradians taylorae to

the Homosassa Bay population of A. i. concentricus. Valve

characters were analyzed morphometrically for two

Florida populations of A. i. concentricus and one of A.

i. taylorae. as well as for the t\ pe series of A. i. taylorae.

Page 34

THE NAUTILUS. Vol. 110, No. 2

Although subspecific morphometric variation within A.

irradians has been investigated in other studies (Clarke,

1965; Waller, 1969), no effort had been made to char-

acterize corresponding genetic variation. This study is

the first to examine both morphological and genetic vari-

ation within a putative subspecies of Argopecten irra-

dians.

MATERIALS AND METHODS

Bay scallops (n = 66) were collected from Rabbit Key

Basin, the type locality of Argopecten irradians taylorae,

on 29 August 1990. The scallops were located in the

southeastern portion of the basin, in a turtle grass (Thal-

assia testudinum Banks e.\ Konig) bed, at a depth of

appro.ximately 2 m. At that time we also collected empty

valves (n of right valves = 13) of A. i. taylorae from that

locality. Scallops were returned to the Florida Depart-

ment of Environmental Protection s Keys Marine Lab-

oratory on Long Key, where sections of adductor muscle

and mantle tissue were dissected from live animals and

stored in liquid nitrogen for subsequent genetic analysis.

Shells were disarticulated, numbered, and returned to

the Florida Marine Research Institute at St. Petersburg.

Additional samples of bay scallops were collected from

Homosassa Bay (n = 60), just west of the mouth of the

Homosassa River, on 24 July 1990 and from Pine Island

Sound (n = 56) on 28 June 1991. The specimens from

Homosassa Bay and Pine Island Sound were also collected

from shallow (z < 2 m) T. testudinum beds; Homosassa

Bay scallops were also examined both genetically and

morphometrically, but Pine Island Sound scallops were

examined only morphometrically.

Electrophoresis. Tissue samples of scallops from Hom-

osassa Bay and Rabbit Key Basin were analyzed for ge-

netic composition using methods reported in Bricelj and

Krause (1992). Eight polymorphic loci (frequency of the

most common allele < 0.99) were examined, represent-

ing the following enzymes: phosphoglucomutase (PGM,

EC 2.7.5.1), glucose-6-phosphate isomerase (GPI, EC

5.3.1.9), octopine dehydrogenase (ODH, EC 1.5.1.11),

superoxide dismutase (SOD-1, EC 1.15.1.1), a-amino acyl

peptide hydrolase (LAP, EC 3.4.1.-), alanyl aminopep-

tidase (AAP, EC 3.4.1.-), dihydrolipoamide dehydroge-

nase (DHLD, EC 1.8.1.4), and nonspecific aminopepti-

dase (AP, EC 3.4.1,-). SOD-1 was only weakly poly-

morphic (three alleles, H = 0.03), and DHLD was in-

consistently resolved; thus these two loci were not in-

cluded in the data analysis.

The following enzymes examined were monomorphic:

superoxide dismutase (SOD-2), /i-galactosida.se (/i-GAL,

EC 3.2.1.23), isocitrate dehydrogenase (two loci, IDH-

1 and IDH-2, EC 1.1.1.42), mannose phosphate isom-

erase (MPI, EC 5.3.1.8), 6-phosphogluconate dehydro-

genase (PGD, EC 1.1.1.44), catalase (CAT, EC 1.11.1.6),

glycerol-3-phosphate dehydrogenase (GPDH, EC

l.i.1.8), trio.se phosphate isomerase (TPl, EC 5.3.1.1),

esterase (two isozymes, EST-1 and EST-2, EC 3.1.-.-),

arginine kinase (ARK, EC 2.7.3.3), L-iditol dehydroge-

nase (IDDH, EC 1.1.1.14), and malic dehydrogenase iso-

zymes (MDH-1 and MDH-2, EC 1.1.1.37).

Alleles were designated using standard notation: the

most common allele at each locus was assigned a value

of 100 and other alleles were defined based on mobility

relative to the most common allele.

Observed genotypic frequencies were compared with

those expected under Hard> -Weinberg equilibrium us-

ing the G-test and, when necessary, Williams correction

for small sample size (Sokal & Rohlf, 1981). Rare alleles

were pooled with the electrophoretically closest common

allele to obtain genotypic class frequencies >5. Hetero-

zygote deficit or excess was determined from the D sta-

tistic where D is the percentage deficit (Selander, 1970);

negative D values indicate a deficit of heterozygous ge-

notypes. Allele frequencies among populations were

compared using the R x C test of independence and the

G-test (Sokal & Rohlf, 1981). Nei's modified genetic dis-

tance (Nei, 1978) was calculated for each population pair

surveyed.

Morphometries. Morphological characters of scallops in

populations from Homosassa Bay, Pine Island Sound, and

Rabbit Key Basin and of the holotype (USNM 859901)

and paratypes (USNM 859902) of Argopecten irradians

taylorae were examined following the methods of Waller

(1969). We also collected and examined a sample of

recently emptied valves of scallops (putatively A. i. tay-

lorae) from Rabbit Key Basin that appeared morpholog-

ically identical to living scallops from that area. For each

scallop we counted right valve plicae and measured plical

height, plical width, and 13 other valve characters di-

mension that have been applied in morphological com-

parisons of bay scallops (Clarke, 1965; Waller, 1969; Ab-

bott, 1974; Petuch, 1987) (Appendix 1).

Mensural characters used by Petuch (1987) in his di-

agnosis were examined using analysis of variance (ANO-

VA) for their relationship to valve size in the same four

populations. Data from the type series of A. i. taylorae

from Rabbit Key Basin were deleted from the analyses

of population effects because of the small sample size (n

= 9) and the resultant undue influence on the statistical

models.

Principal component analysis (PCA) was performed

using only mensural variables. Marcus (1990) suggested

that measurement error can unduly influence variability

in the PCA model, so standard errors of measurement

were calculated for each mensural character to estimate

measurement error. Each character was measured 10

times on each of six specimens, two from each population,

and standard errors were calculated for each specimen-

by-character combination. Standard errors were then ex-

pressed as a percentage of the mean. L'sing all fifteen

variables we log transformed Uogio) the \alues and per-

formed a standard PCA (PRINCOMP procedure, SAS

Institute, Inc., 1985). Burnab\ s (1966) method of size-

corrected PCA was performed separateK on all data

[size-corrected PCA (N. Macleod, personal communi-

cation)]. Burnaby s method removes most of the influence

of size on the PCA by constructing variables orthogonal

Dan C. Marelli et a/., 1997

Page 35

Table 2. Allele frequencies for polymorphic enzymes from

populations of Argopecten irradians from Rabbit Key Basin

and Homosassa Bay, Florida, n = number of alleles sampled

(2 X number of individuals). N^ = effective number of alleles

(after Hartl & Clark, 1989).

Locus

AP

AAP

LAP

ODH

PGM

GPI

Allele

108

104

100

96

94

n

Ne

102

100

98

97

96

94

n

N.

100

96

94

n

N,

106

104

102

100

96

94

90

n

N.

98

96

95

94

92

91

90

n

N.

150

106

104

100

99

98

n

Frequency

RKB

HOM

0

0,38

0.53

0.08

0.01

132

2.32

0

0.35

0.17

0.13

0.19

0.15

132

4.37

0.62

0.37

0,01

132

1,92

0.01

0.08

0

0.77

0.07

0.01

0.05

132

1.63

0.02

0.49

0.1-4

0.13

0.11

0.03

0.07

132

3.36

0

0.06

0.61

0.30

0.02

0.01

132

2.14

0.02

0,23

0.51

0.24

0

118

2.70

0.03

0.40

0.24

0.04

0.17

0.13

112

3.79

0.85

0.15

0

118

1,35

0.02*

0,13

0.01

0.80

0

0.04

0

118

1.52

0.01*

0.56

0.18

0.19

0.01

0

0.05

lis

2.61

0,01

0.02

0.62

0.33

0.01

0

118

2()2

* Genotype frequencies not in Hardy-Weinberg equilibrium.

to a variable that is considered to represent size (Rohlf

& Bookstein, 1987). The relative contribution of each

variable to the variation represented by PC2 and PC3

was determined, and Cattell s (,1966) scree test was used

to identify and eliminate variables that failed to add

RKB-USNM RKB-FMRI PISL

LOCALITY

HOM

Figure 2. Valve heights of Florida Gulf coast Argopecten ir-

radians populations. Plots show mean, range, and ± 1 standard

error. RKB-USNM: Rabbit Key Basin, type seriesoi Argopecten

irradians taylorae. USNM 859901 & 859902; RKB-FMRL Rab-

bit Ke\ Basin, collection made for this study, FSBC I 41441;

PISL: Pine Island Sound, FSBC 1 44439; HONI: Homosassa Bay,

FSBC 1 40337.

CD

s

3

Z

-I

<

o

RKB-USNM RKB-FMRI PISL

LOCALITY

HOM

Figure 3. Numbers of right valve plicae for Florida Gulf coast

Argopecten irradians populations. Plots show mean, range, and

± 1 standard error RKB-L SNM: Rabbit Ke\ Basin, type series

of Argopecten irradians taylorae. USNM 859901 & 859902;

RKB-FMRL Rabbit Key Basin, collection made for this study,

FSBC I 41441; PISL: Pine Island Sound, FSBC I 44439; HOM:

Homosassa Bav, FSBC I 40337.

substantially to the variance in the analysis, leaving 9

variables: AM, AD, DG. AK, GP, BJ, IW, ad, and ce (see

Appendix 1 for explanations). Using only these variables,

data were reanalyzed, and size-adjusted shape variation

among populations was examined using PC2 and PC3

generated b\ the Burnaby technique.

Page 36

THE NAUTILUS, Vol. 110, No, 2

Figures 4-7. Left valves of Argopeclen irradians from Florida. 4. A. i. concentricus. height 60.0 mm. Rabbit Ke> Basin, FSBC I

41441. 5. A. i. taylorae. holotype, height 44.0 mm. Rabbit Key Basin, USNM 859901. 6. A. i. concentricus. height 60.1 mm,

Homosassa Bay, FSBC I 40337. 7. A. i. concentricus, height 57.9 mm. Pine Island Sound, FSBC I 44439. Scale bar = 10 mm

Growth relationships of \alve characters within and

between populations were examined using the method

of Jolicoeur (1963). Eigenvectors (coefficients) of PCI

from the standard PCAs using log-transformed data for

all populations combined and separately on populations

from the three localities were compared with (l/p)"^,

where p = the number of variables and therefore (1/

p)0o = 0.333. Coefficients for valve characters that varied

above or below 0.333 represented factors that reflected

positive or negative allometry.

Valve color and pattern were examined following the

criteria of Elek and Adamkewicz (1990), with the ex-

ception that the relationship between color pattern and

population was examined using the nonparametric Krus-

kal-W'allis procedure.

Disposition of specimens: Except for the t\pe material

of Argopectcn irradians tatjlorac, valves of all specimens

examined have been deposited in the Florida Marine

Research Institute Invertebrate Collection, lot numbers

FSBC I 40337 (Homosassa Bay), FSBC I 41441 (Rabbit

Key Basin), and FSBC I 44439 (Pine Island Sound).

RESULTS

Electrophoresis. Among the poK morphic loci no sig-

nificant differences were found bet\veen populations for

the loci GPI, ODH, AAP, or PGM. Allele frequencies

between Homosassa Bay and Rabbit Key Basin popu-

lations were significantly different for amino peptidase

(AP, p < 0.05) and leucine amino peptidase (LAP, p <

0.001) (table 2).

The mean percentages of heteroz) gous loci per indi-

vidual (MLH) for Homosassa Bay and Rabbit Key Basin

populations were 43^7 and 44*7 using the six polymorphic

loci. Homo.sassa Bay scallops fit Hardv -\\ einberg expec-

tations for heterozygosit) at all loci, but the Rabbit Key

Basin population had heterozvgote deficits at the PGM

(p < 0.05) and ODH (p < 0.01) loci.

Dan C. Marelli et a/„ 1997

Page 37

Table 3. Growth relationships for valve characters from Rab-

bit Key Basin (RKB), Pine Island Sound (PISL), and Homosassa

Bay (HOM), Florida Argopecten irradians populations ana-

lyzed using principal component analysis. Numbers are eigen-

vectors of principal component 1 and indicate positive or neg-

ative allometric growth where eigenvectors var>' above or below

l/Vp. where p = the total number of characters, following

Jolicoeur (1963).

The mean genetic distance between the Homosassa

Bay and Rabbit Key Basin populations was 0.035.

Morphometries. Standard error estimates for measure-

ments of morphometric characters were universally low,

averaging 0.64% of the mean for all variables, suggesting

that measurement error did not overly influence the vari-

ance in subsequent morphometric analyses.

Scallops from the Pine Island Sound collection were

significantly larger than those from Homosassa Bay, and

both Pine Island Sound and Homosassa Bay scallops were

much larger than those from our Rabbit Key Basin col-

lection (AXOVA, all p < 0.05) (figures 2-3). The type

specimens of ArgopfTfen irradians fay/oraf were slightly

larger than the live specimens from our Rabbit Key Basin

collection and similar in size to the empty valves we

collected from Rabbit Key Basin, although two pairs of

empty valves exceeded 50 mm in height. Representative

valves from the sample populations are illustrated in

figures 4-7.

Numbers of right-valve plicae were similar for all sam-

ple populations that we examined morphometrically, and

plical number was not significantly related to valve height

(0.05 < p < 0.06). However, the effect of population on

plical number was significant (p < 0.0001), and right-

valve plicae were significantly more numerous on Hom-

osassa Bay scallops (p < 0.05) than on Rabbit Key Basin

and Pine Island Sound scallops. Plical numbers were not

significantly different between Rabbit Key and Pine Is-

land Sound populations. Width of plicae relative to valve

size was positively related to valve height (p < 0.0002).

Patterns of growth for individual mensural characters

varied greatly and inconsistently among populations (ta-

ble 3). Although some characters varied isometrically,

there was no consistent trend among populations.

Coloration of valves from all populations was similar;

more than 90% of specimens from each population ex-

hibited the color pattern of t\ pe E of Elek and Adamke-

wicz (1990). Variations in color pattern were not signif-

icantly different among populations. More than 95% of

the valves from each scallop population had white back-

grounds, and 100% of the specimens had mottled left

valves (table 4). Excluding one yellow and two orange

scallops from Rabbit Key Basin and one orange scallop

from Pine Island Sound, right valves in all populations

were consistently all white except in the early juvenile

region of the shell.

The first principal component (PCI) from the standard

PCA of all data accounted for 83.3% of the total variance,

and the second and third principal components account-

ed for 7.7%' and 4.1%, respectively, of the variance. Bur-

naby size-corrected PCA showed that principal com-

ponents 1 through 3 accounted for 59.2%-, 14.9%, and

9.8% of total variance, respectively. Two variables in the

Burnaby PCA accounted for most of the variance in PC2

and PC3: width of plical interspaces (IW) represented

28.5% of PC2 variance and 69.5% of PC3 variance, and

ce (length of ligament insertion) accounted for 33.6% of

PC2 variance and 20.4% of PC3 variance (table 5). Plots

of PC2 and PC3 from the size-corrected PCA of the data

from all 3 populations and including the separate col-

lection of empty valves from Rabbit Key Basin (figure

8) demonstrate that scatterplots from each population

overlap with those from all other populations. Particu-

larly extensive overlap occurs between the Rabbit Key

Basin and Homosassa Bay populations, and less overlap

is seen in plots representing the Pine Island Sound pop-

ulation and those of all other populations.

DISCUSSION

The genetic distance (D = 0.035) between the Homosassa

Bay and Rabbit Key Basin populations of Argopecten

Table 4. Summar\' of

Island Sound (PISL, n =

color and color pattern on valves of Argopecten irradians from Rabbit Key Basin (RKB, n — 66),

= 56), and Humusassa Ba\ iHOM, n = 60), Florida. Numbers expressed as percentages.

Pine

Page 38

THE NAUTILUS, Vol. 110, No. 2

Table 5. Percentage of variance in principal components 2

and 3 from Burnaby PCA attributable to individual valve char-

acters for Argopecten irradian^ populations from Rabbit Key

Basin, Pine Island Sound, and Homosassa Bay. Character ab-

breviations are defined in the appendi.x.

0.4

irradians s.l. is on the order of distances between local

races (Nei, 1976, 1987). This value is comparable to those

between disjunct but apparently recently separated pop-

ulations of the Baltic clam Macoma balthica (Linnaeus,

1758) (Nei's D = 0.058) (Meehan et ai, 1989) and be-

tween Great Barrier Reef and Enewetak Atoll popula-

tions of the giant clam Tridacna maxima (Nei's D =

0.033) (Ayala, 1975). Electrophoretic data from five ad-

ditional populations of Argopecten irradians s.l. exam-

ined by Krause (1992) (Martha's Vineyard, Massachu-

setts; Niantic River, Connecticut; Orient Harbor, Long

Island, New York; Core Banks, North Carolina; and St.

Joseph Bay, Florida), treated identicalK , were used along

with data from the Rabbit Ke) Basin and Homosassa

Bay populations to create a genetic distance matrix. A

dendrogram was constructed from the genetic distance

matrix using the unweighted pair-group method with

arithmetic mean (UPGMA Sneath & Sokal, 1973), pro-

vided by the computer program NTSYS (Applied Bio-

statistics, Inc.) (figure 9). Standard errors of tree branch-

ing points were estimated using the procedure of Nei et

al. (1985) who point out that, when using electrophoretic

data and less than 30 loci, the topology of a reconstructed

tree is subject to a large stochastic error. The size of the

errors allows us little confidence in the dendrogram, but

two factors may contribute to an inflation of the esti-

mated standard errors. When genetic distance values are

very low, less than 0.105, the value of 1 in the equation

D = -log,, I exceeds 0.9. When I >0.9 and average het-

erozygositv (VILH) is >0.2 serious overestimation of the

variance may occur (Nei et ai, 1985). Twelve of 21

pairwise I values in the scallop genetic distance matrix

exceed 0.9 and MLH = 0.44 p. Despite the size of the

standard errors, the genetic distance data indicate a close

relationship between the Rabbit Key Basin and Homo-

sassa Bay scallops within Argopecten irradians.

The small but significant differences in allele fre-

quency between Homosassa Bay and Rabbit Key Basin

populations at the AP and L.'\P loci might be evidence

for localized selection among genotypes between these

sites (McMillen et ai, 1994), for reduced gene flow be-

0.3

CO

H

u 0.2

Z

o

a

o

u

<

a.

O 0.1

z

oc

a.

-0.1

1.3 1.4 1.5 1.6 1.7

PRINCIPAL COMPONENT 2

1.8

1.9

Figure 8. Bivariate scatterplot of principal component 2 and

3 scores generated using Burnabv 's (1966) size-corrected prin-

cipal component analvsis on morpliometric measurements of

Argopecten irradians from Rabbit Key Basin (D), Pine Island

Sound (■). and Homosassa Bav (A) populations. Population

indicated by (•) represents empty valves collected from Rabbit

Kev Basin localitv ,

tween the populations (Beaumont & Zouros, 1991). or

some combination of these processes. The overall mag-

nitude of genetic variation among the Florida popula-

tions is onl\ slightK greater than that found among

Krause's (1992) northern populations of A. i. irradians.

Therefore A. i. taylorae appears, based on genetic evi-

dence, not to differ at the subspecific level from other

Florida populations examined.

X'alues for the mean percentage of heterozygous loci

per individual (MLH) in the Homosassa Ba\ (43?c) and

Rabbit Ke\ Basin (44%) populations are relativeK high

but similar to the 45.3% MLH reported by Bricelj and

Krause (1992) for Argopecten irradians irradians from

the Niantic River estuary, Connecticut. These values in-

dicate that high MLH values may be characteristic of

Argopecten irradians s.l., but Wall et al. (1976) reported

a MLH of 11.6% for A. irradians from Bogue Sound,

North Carolina. Values of MLH reported for 5 other

pectinids range from 9.4% to 32.1% (Nikiforov & Dol-

gonov , 1982; Beaumont & Beveridge, 1984).

Color in .\rgopecteri irradians is genetically controlled

Dan C. Marelli et al.. 1997

Page 39

(0.026)

(0.014)

[(oToos)

— (0.04)

(0.019)

(0.021)

MASS

CONN

LINY

CBNC

SJFL

HOM

RKB

0.15

0.125

0.1

0.075

0.05

GENETIC DISTANCE

0.025

-I

0

Figure 9. I'PGMA tree constructed trom tlie Nei (1972) genetic distance matrix for populations of Argopecten irradians. Population

abbreviations are MASS: Martha's Vineyard, Massachusetts; CONN: Niantic River, Connecticut; LINY: Orient Harbor, Long Island,

New York; CBNC: Core Banks, North Carolina; SJFL: St. Joseph Bay, Florida; HOM: Homosassa Bay, Florida; RKB: Rabbit Key

Basin, Florida. Numbers in parentheses are standard errors of branching points estimated using the procedure of Nei et al. (1985).

and may sometimes be useful in distinguishing individ-

uals and populations (Kraeuter et al.. 1984; Adamkewicz

& Castagna, 1988). Nevertheless, we found no substantial

differences in valve coloration or color pattern between

the Homosassa Bay, Pine Island Sound, and Rabbit Key

Basin scallop populations to support Petuch's (1987) con-

tention that shells of A. i. taylorae are much more colorful

than those of A. i. concent ricii.s. Most scallops in each of

the populations had a rayed pattern on the left valve,

and additional color was generalK expressed as mottling

and banding, somewhat obscuring the rayed pattern.

Variations in the intensity of color were affected by the

nature and e.xtent of fouling on the left \alve and possibly

by ontogenetic change.

Petuch (1987) identified the small size of scallops from

the Rabbit Key Basin population as an important char-

acter that distinguishes Argopecten irradians taylorae

from A. i. concentricus. Although heights of live scallops

from our Rabbit Key Basin collection were significantly

smaller than those in all other collections, the empty

valves that we collected from the same locality were

comparable in size to those in Petuch's type series. This

suggests that the scallops in Rabbit Ke\' Basin may indeed

be distinctively smaller than other Florida scallops, al-

though the close genetic similarity between the Rabbit

Key Basin and Homosassa Ba\ populations indicates that

scallop size is readily influenced by local conditions. The

small size of our live Rabbit Key Basin collection coupled

with the heterozygote deficits at the loci ODH and PGM

that occurred among the scallops from this collection

[similar to deficits at these loci that have been reported

for juveniles of other species of Pectinidae (Volckaert &

Zouros, 1989; Bricelj & Krause, 1992)] may indicate that

the scallops we collected from Rabbit Key Basin were

juveniles.

Discontinuities in plical number occur between pop-

ulations of Argopecten irradiana along the geographic

range of the species, and these differences have been

emphasized ta.xonomically (Clarke, 1965). The number

of plicae ranges from 12 to 25 in the described subspecies

(Clarke, 1965; Waller, 1969; Abbott, 1974; Petuch, 1987)

and is reportedly under genetic control (Kraeuter et al.,

1984). We substantiate the elevated plical numbers that

have been reported for Florida A. i. concentricus. but

we did not find even greater plical counts in the Rabbit

Key Basin population as reported b> Petuch (1987). In

fact, the mean plical number for our sample of scallops

from Rabbit Key Basin was statistically indistinguishable

from that of scallops from Pine Island Sound. Higher

counts of plicae for the type series reported by Petuch

(1987) suggest that he may have misidentified as plicae

some of the "riblets" that occur on the disk flanks of this

species; such confusion has been previously reported by

Clarke (1965) and Waller (1969). Our analysis of right

valve plical numbers rejects the conclusion that scallops

from Rabbit Key Basin represent a separate and unique

subspecies.

Width of the plicae at the ventral margin was iden-

tified b\ Petuch (1987) as a character that can be used

to separate Argopecten irradians taylorae from A. i.

concentricus. However, because plical width relative to

valve size is positively correlated with valve height, its

use in a univariate comparison of scallops of different

sizes without accounting for allometry is invalidated.

Moreover, our data indicate that Rabbit Key Basin scal-

lops have relatively narrower plicae than do scallops from

Homosassa Bay or Pine Island Sound — not wider as re-

ported by Petuch.

Inconsistencies in the relationship between growth and

valve characters between populations indicate that mor-

phometric characters do not vary with growth in a simple

manner; variability due to small sample sizes and mea-

Page 40

THE NAUTILUS, Vol. 110, No. 2

surement error may also affect the allometric coefficients

(Marcus, 1990). Regardless of the source, the inconstancy

of shape variables and the presence of allometric rela-

tionships among mensural characters, morphometric ra-

tios, and size cast doubt upon the use of univariate char-

acters in describing morphometric differences between

scallop populations, adding empirical emphasis to warn-

ings b\ previous authors (summarized in Humphries et

ai. IQ'sI).

Burnab)' size-corrected PCA failed to distinctly sep-

arate the Rabbit Key Basin, Pine Island Sound, and Hom-

osassa Bay scallop populations. Although morphometric

overlap among populations is e\ ident in the plots of PC2

and PCS, scatterplots of the individual populations form

distinct clusters and indicate that the more geographi-

calK distant populations (Rabbit Ke> Basin and Hom-

osassa Bay) in our samples were morphometrically more

similar. Variation in shape, and hence some separation

in the plots of PC2 and PC3, may be influenced by

ontogeny of individuals, because the Burnaby technique

does not completeK remove the effect of size (Humphries

etal., 1981), but separationon the basis of shape variables

indicates that scallop shape is heavily influenced by local

conditions and may simply reflect ecophenotvpic vari-

ation.

We assessed alleged differences between typical Ar-

gopecten irradians taylorae and representatives of other

bay scallops from peninsular Florida. Based on the results

of our electrophoretic and morphometric examinations,

we refute those differences and conclude that bay scal-

lops from Rabbit Key Basin do not represent a subspecific

taxon distinct from Florida populations of A. i. concen-

tricus.

ACKNOWLEDGEMENTS

Clarita Lund, Brenda Hedin, Richard Darden, Yantian

Lu, Don Hesselman, and James Seagle assisted in col-

lecting scallops, and C. Lund also performed data entry.

Robert McWilliams and Charlotte LaTorre measured

valve characters. Catherine Bray assisted in data analysis

and figure preparation. Ll\ n French prepared figures.

Dr. Theresa Bert and Hector Cruz-Lopez helped with

interpretation of electrophoretic data. Everglades Na-

tional Park biologist Daniel Foxen facilitated our Florida

Bay collection. This project was partially supported b\

funds created by the Florida Saltwater Products License

and the Florida Saltwater Fishing License. Electropho-

retic anaKses were supported by a National Science

Foundation dissertation improvement grant (BSR-

9015991) to MKK

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2: 66-85.

Nikiforov, S. M. and S. M. Dolganov. 1982. Genetic variation

of the Japanese scallop Patinopecten yessoensis from the

Vostok Bay, Sea of Japan. Biologia Morva, Vladivostok 2:

46-50.

Orensanz, J. M, A. M. Parma and O. O. Iribarne. 1991. Pop-

ulation dynamics and management of natural stocks. In:

Shumway, S. E. (ed.). Scallops: Biology, Ecology and Aqua-

culture. Developments in aquaculture and fisheries science

21. Elsevier Science Publishers, New Y'ork, pp. 625-713.

Petuch, E. J. 1987. New Caribbean molluscan faunas. The

Coastal Education and Research Foundation, Charlottes-

ville, N'irginia, 158 pp.

Rohlf, F. J. and F. L. Bookstein. 1987.

as a method for "size correction.

356-367.

Roughgarden, J. and Y". Iwasa. 1986 D\namics of a meta-

population with space-limited subpopulations. Theoretical

Population Biology 29: 235-261.

Roughgarden, J, Y" Iwasa and C. Baxter 1985. Demographic

theor> for an open marine population with space-limited

recruitment. Ecology 66:54-67.

SAS Institute, Inc. 1985. SAS user's guide: statistics. Version

5 edition. SAS Institute, Inc., Gary, North Carolina, 956

pp.

A comment on shearing

S\stematic Zoology 36:

Sa} , T. 1822. .'\n account of some of the marine shells of the

United States. Journal of the Academy of Natural Sciences

of Philadelphia 2(2):257-276.

Selander, R K 1970. Behasiour and genetic variation in

natural populations. American Zoologist 10:53-66.

Simberloff , D. 1988 The contribution of population and com-

munity biology to conservation science. Annual Review of

Ecology and Systematics 19:473-512.

Sinclair, M.,'r. K. Mohn, G. Probert and D. L. Roddick. 1985.

Considerations for the effective management of Atlantic

scallops. Canadian Technical Report of Fisheries and

Aquatic Sciences 1382, 97 pp.

Sneath, P. H. A. and R. R. Sokal. 1973. Numerical taxonomy.

Freeman, San Francisco. 573 pp.

Sokal, R R and F. J Rohlf. 1981. Biometry, 2nd edition. W.

H. Freeman, San Francisco, 859 pp.

Tabb, D. C. and R. B. Manning. 1961. A checklist of the flora

and fauna of northern Florida Bay and adjacent brackish

waters of the Florida mainland collected during the period

July, 1957 through September, 1960. Bulletin of Marine

Science of the Gulf and Caribbean ll(4):552-649.

Volckaert, F. and E. Zouros. 1989. Allozyme and physiolog-

ical \ariation in the scallop Placopecten magellanicus and

a general model for the effects of heteroz\ gosity on fitness

in marine molluscs, Mamie Biology 103:51-61.

Wall, J. R, S. R. Wall and M. Castagna' 1976. Enzymes [sic]

polymorphisms and genetic variation in the bay scallop,

Argopecten irradians. Genetics 83(3, part 1, Suppl.):81.

Waller, T. R. 1969. The evolution of the Argopecten gibbus

stock (Mollusca: Bivalvia), with emphasis on the Tertiary

and Quaternary species of eastern North America. Journal

of Paleontology 43:1-125.

Waller, T. R. 1991. Evolutionary relationships among com-

mercial scallops (Mollusca: Bivalvia: Pectinidae). In:

Shumway, S. E. (ed.). Scallops: Biology, Ecology and Aqua-

culture. Developments in aquaculture and fisheries science

21. Elsevier Science Publishers, New York, p 1-73.

APPENDIX I

Table of morphological characters and abbreviations used

in this study, following Waller (1969). Abbreviations re-

fer to either right or left-valve; apostrophes are added

in te.xt to indicate left valve measurements.

AM

AD

DG

AK

GP

DF

CD

EI

BJ

DE

LO

ad

ce

PW

IW

Valve height

Length of posterior portion of valve

Length of anterior portion of valve

Height of posterodorsal portion of valve

Height of anterodorsal portion of valve

Length of anterior ligament

Length of posterior ligament

Height of anterior auricle

Height of posterior auricle

Length from valve midline to ventral insertion

of

anterior auricle

Valve width

Height of resilial insertion

Length of resilial insertion

Width of medial plicae at ventral margin

Width of medial interplical spaces at ventral

margin

THE NAUTILUS 110(2):42-44, 1997

Page 42

Subspecific Status of Argopecten irradians concentricus

(Say, 1822) and of the Bay Scallops of Florida

Dan C. Marelli

William G. Lyons

William S. Arnold

P'lorida Department of

Environmental Protection

Florida Marine Research Institute

100 8th Avenue SE

St. Petersburg, FL 33701-5095 USA

Maureen K. Krause'

Department of Ecology and

Evolution

State University of New York at

Stonv Brook

Stony Brook, NY 11794-5245 USA

ABSTRACT

Nei genetic distances between Florida and North Carolina pop-

ulations of ba\' scallops, all putatively Argopecten irradians

concentricus, are greater than distances between the North

Carolina population and populations of A. i. irradians from

New York to Massachusetts. The subspecies A. i. concentricus

is probably untenable. Evidence for a subspecies, A. i. taylorae,

in Florida and the eastern Gulf of Mexico is discussed.

Key words: Argopecten. bay scallops, genetics, Pectinidae,

subspecies.

INTRODUCTION

Bay scallops, Argopecten irradians (Lamarck, 1819),

range from New England to Florida, Texas, and north-

eastern Mexico. Northern, southern, and western popu-

lations have been accorded subspecific rank as A. i. ir-

radians, A. i. concentricus (Say, 1822), and A. i. ampli-

costatus (Dall, 1898) (Clarke, 1965; Waller, 1969). We

have found no morphometric or genetic characters to

separate scallops of a recently proposed fourth subspe-

cies, A. i. taylorae Petuch, 1987, from other Florida

populations (Marelli et al., 1996) customarily assigned to

A. i. concentricus. However, our work shows that Nei

(1972) genetic distances between Florida populations and

a North Carolina population of Argopecten irradians,

all putativeK of A. i. concentricus, are greater than dis-

tances between that North Carolina population and pop-

ulations of A. I. irradians from Massachusetts, Connect-

icut, and New York (figure 1), prompting us to review

and reassess the subspecific status of A. i. concentricus.

Say (1822) described Pecten concentricus based upon

' Current address: Southampton College, Long Island Univer-

sity, Southampton, NY 11968 USA.

specimens from the coast of New Jersey and also rec-

ognized as distinct a variety from New England that he

named Pecten borealis-. the latter name is now known

to be a junior synonym of A. i. irradians. Because Say's

specimens of P. concentricus are lost, Clarke (1965) se-

lected a neotype from Great Egg Harbor near Atlantic

City in southeastern New Jersey, thereby fixing the type

locality. Clarke also restricted the type locality of P.

irradians to Waquoit Bay near Falmouth, Massachusetts.

Clarke (1965) evaluated the status of subspecific units

among bay scallops by comparing four "primary char-

acters" [plical count; width/length; height/length; and

color (% white)] of shells from 23 populations throughout

the range of what he called "the Argopecten irradians

superspecies. Clarke's results, derived from relatively

few specimens (.\ = 13) of disparate sizes (height range:

17.7 to 80.6 mm), must be viewed with uncertainty be-

cause of our findings of allometric \ariabilit\ among

individual populations (Marelli et at., 1996). Neverthe-

less, Clarke s four samples from the "New Jersev coast ";

Atlantic City, New Jersey; Sinepuxent Bay, Maryland;

and Hog Island, Virginia are instructi\ e in understanding

his concept of A. i. concentricus. The New Jersey coast

sample (depicted as northern New Jersey but actually

unlocalized) had characters consistent with four other

samples from Massachusetts and Rhode Island (i.e., A. i.

irradians): the Atlantic Cit\ and Sinepuxent Ba\' samples

shared characters of both the more northern and more

southern (North Carolina) samples; and the Hog Island

sample grouped with two samples from North Carolina,

which in turn most resembled five samples from the

eastern Gulf of Mexico between Sanibel Island, Florida,

and Ft. Morgan, Alabama. A specimen from Chandeleur

Island, Louisiana, was depicted as intermediate between

those of eastern and western Guli (Matagorda, Texas, to

Tampico, Mexico; i.e., A. i. aniplicostatus) populations,

but its plical count was clearly assignable to the eastern

group.

Clarke interpreted his results to indicate that northern.

Dan C. Marelli et ai, 1997

Page 43

(0.026)

(0.014)

[(oToos)

— (0.04)

(0.019)

(0.021)

MASS

CONN

LINY

CBNC

SJFL

HOM

RKB

0.15 0.125 0.1 0.075 0.05 0.025 0

GENETIC DISTANCE

Figure 1. UPGMA tree constructed from the Nei (1972) genetic distance matrix for populations of Argopecten irradians. See

Marelli et al. (1996) for methods. Abbreviations are MASS; Martha's Vineyard, Massachusetts; CONN; Niantic River, Connecticut;

LINY; Orient Harbor, Long Island, New York; CBNC; Core Banks, North Carolina; SJFL; St. Joseph Bay, Florida; HOM; Homosassa

Bay, Florida; RKB; Rabbit Key Basin, Florida. Numbers in parentheses are standard errors of branching points estimated using

procedures of Nei et al. (1985).

southern, and western populations of A. irradians are

morphologically dissimilar throughout most of their

ranges but that they intergrade between New Jersey and

Virginia and again in Louisiana. Despite the transitional

features of scallops in the t\ pe population, Clarke tried

to maintain nomenclatural stability by retaining the name

concent riciis for the economically important subspecies

of the southern United States. In retrospect, Clarke s

findings seem to have provided more support for the

existence of clines than for the existence of subspecies.

The subspecies concept is regarded by some as arbi-

trary because it attempts to separate populations that, in

fact, may show gradual transitions in morphology, phys-

iology, and genetics over the range of a species (Wilson

& Brown, 1953; Endler, 1977). The classification of Ar-

gopecten irradians exemplifies difficulties inherent in

attempts to distinguish such populations. The genetic

distances we report for bay scallops from Florida to Mas-

sachusetts could indicate a cline among populations be-

tween which no clear boundaries exist, and that is prob-

abl\' the case among the Atlantic coast populations north

of Florida. The transitional nature of morphological

characters of the type population of A. i. concent ricus

seems to support that explanation, as do genetic similar-

ities we found among populations between North Car-

olina and Massachusetts. Thus, the concept of A. i. con-

centricus as a subspecies seems untenable.

The subspecies concept gains strength, however, w hen

it is applied to populations that are separated by distri-

butional gaps or barriers that constrain contact {sensii

Mayr, 1969). Waller (1969) described a distributional gap

of approximately 764 km (475 miles) between the east-

ernmost population of A. !. aniplicostatus and the west-

ernmost population he considered to represent A. i. con-

centricus. and he proposed that the Mississippi Delta acts

as a barrier to genetic interchange between those stocks.

Thus, subspecific designations for those populations seem

appropriate.

A distributional gap of similar magnitude in the south-

eastern United States has received less attention, perhaps

because it contains no obvious barrier. The northernmost

population of bay scallops along eastern Florida is at

Palm Beach Inlet (Lake Worth Inlet) (Waller, 1969) near

latitude 26°45'\, and the next documented occurrence

of the species seems to be at Lockwood Folly Inlet near

Cape Fear, North Carolina (Porter, 1974), near SS'SO'N;

these populations are separated by a minimum distance

of nearly 800 km. Even if reports of uncommon occur-

rence in South Carolina (Clarke, 1965; Shoemaker et ah,

1978) and an unsubstantiated report from Georgia (Pe-

tuch, 1987) are considered, no population of bay scallops

is known in the region between Palm Beach Inlet and

the Florida-Georgia border at 30°45'N, a gap of nearly

450 km. Thus, despite reported similarities in shell mor-

phology, the genetic gap between the North Carolina

and Florida stocks is not a simple expression of distant

populations separated by other, more contiguous popu-

lations distributed along a cline. Instead, the bay scallop

stocks of Florida (and probably those extending to eastern

Louisiana) seem to comprise an aggregation of popula-

tions isolated b\ large distances from other aggregations

to the north and to the west, i.e., a subspecies, sensu

Mayr (1969).

If A. i. concentricus is not appropriate as a name for

bay scallop stocks of Florida and the eastern Gulf of

Mexico, and if those stocks merit subspecific recognition,

then A. i. taijlorae Petuch, 1987, is the first available

name. Another possibility, Pecten circularis Sowerby,

1835, was shown by Waller (1995) to be a junior synonym

of Argopecten irradians concentricus. Dr. Waller in-

Page 44

THE NAUTILUS, Vol. 110, No. 2

formed us (in litt.. 1/30/96) that the holot\pe of Pecten

circiilaris "compares favorabl\ w ith North Carohna con-

centricus in overall shape, auricular shape, rib shape,

and rib count, and it differs in these features from Gulf

populations."

We advocate caution to those who might immediately

appl\- the name Argopccten irradians taijlorac for the

Florida and eastern Gulf stocks. Our study (Marelli et

al., 1996) was intended only to assess differences between

t\pical A. 1. taylorae and other bay scallops of Florida.

Those differences were refuted using analytical tech-

niques involving both genetics and morphometries. Sim-

ilar work is needed to elucidate the genetic compositions

of populations in the transition zone between New Jersey

and \'irginia, in South Carolina and Georgia (if such

populations exist), in the region between northwestern

Florida and eastern Louisiana, and in the western Gulf

of Me.xico (A. !. amplicostatus) to provide perspective

for evaluating genetic distances among all of the popu-

lations before the question of subspecies is resolved.

ACKNOWLEDGEMENTS

Dr. Arthur Bogan, then at the Academy of Natural Sci-

ences of Philadelphia, provided a copy of Say (1822).

Dr. Thomas R. Waller, National Museum of Natural

History, Washington, D. C, provided additional infor-

mation on the holotype of Pecten circiilaris. This project

was partially supported by funds created by the Florida

Saltwater Products and the Florida Saltwater Fishing

License.

LITERATURE CITED

Clarke, A. H, Jr. 1965. The scallop superspecies Aequtpec/en

irradians (Lamarck). Malacoiogia 2:161-188,

Dall, W. H. 1898. Contributions to the Tertiary fauna of

Florida, with especial reference to the Miocene silex beds

of Tampa and the Pliocene beds of the Caloosahatchie

River; Wagner Free Institute of Science Transactions, Vol.

3, Part 4, 377 p.

Endler, J. A. 1977. Geographic variation, speciation, and clines.

Princeton Universitv Press, Princeton, New Jersey, 246 p.

Lamarck, J. B P. A de VI de. 1819. Histoire natiirelle des

animaux sans vertebres. Paris, Vol. 6, Part 1, 343 p.

Marelli, D. C, M. K. Krause, W. S. Arnold and W. G. Lyons.

1 997. Systematic relationships among Florida populations

of Argopccten irradians scnsu lata (Lamarck. 1819) (Bi-

valvia: Pectinidae), The Nautilus 110(2):31-41.

Mayr, E. 1969. Principles of svstematic zoologv. McGraw-

Hill, New York, 428 p.

Nei, M. 1972. Genetic distance between populations. The

American Naturalist 106: 283-292.

Nei, M., J. C. Stephens and N. Saitou. 1985, Methods for com-

puting the standard errors of branching points in an evo-

lutionary tree and their application to molecular data from

humans and apes. Molecular Biolog\ and E\ olution 2: 66-

85,

Petuch. E, J, 1987 New Caribbean molluscan faunas. The

Coastal Education and Research Foundation, Charlottes-

ville, Virginia, 158 p.

Porter, H, J. 1974, The North Carolina marine and estuarine

Mollusca — an atlas of occurrence. University of North Car-

olina Institute of Marine Sciences, Morehead City. North

Carolina, 351 p.

Say, T, 1822. .\u account of some of the marine shells of the

United States, Journal of the Academs' of Natural Sciences

of Philadelphia 2(2):257-276,

Shoemaker, A, H,, H, J, Porter. B. Boothe, R. E. Petit and L.

S, Eyster, 1978, Marine mollusks. In: R, G, Zingmark (ed.)

An annotated checklist of the biota of the coastal zone of

South Carolina. University of South Carolina Press, Co-

lumbia, SC. p. 123-135.

Waller, T. R. 1969. The evolution of the Argopecten gibhus

stock (Mollusca: Bivalvia), with emphasis on the Tertiary

and Quaternar\ species of eastern North .America, Journal

of Paleontology 43:1-125

Waller. T. R 1995. The misidentified holot\ pe of Argopecten

circularis iBi\aK ia: Pectinidae), The \eliger 38: 298-303,

Wilson, E. O. and W. L. Brown. 1953. The subspecies concept

and its taxonomic applications. Svstematic Zoologv 2:97-

111.

THE NAUTILUS 110(2):45-54, 1997

Page 45

Distribution of Unionoid (Bivalvia) Faunas in Minnesota, USA

Daniel L. Graf

Department of Biology-

Northeastern University

Boston, MA 02115 USA

and

Department of Mollusks

Museum of Comparative Zoology

Cambridge, MA 02138

ABSTRACT

The freshwaters of Minnesota are significant because they form

the conjunction of three major drainage s\ stems: the Mississippi,

the St, Lawrence, and Hudson Bay; 46 species of freshwater

mussels (Bivalvia: Unionoidea) inhabit these rivers and lakes.

Of these, 31 are found only in the Mississippi River and its

tributaries below Minneapolis. Nine species have distributions

that include all of the state's drainage systems; 5 species have

been reliably reported from only the Red River of the North

and Lower Mississippi River drainage systems; and Elliptio

complanata is found only in the Lake Superior Basin. Prob-

lematical records within the state are addressed.

Key words: Biogeography, Unionidae, Margaritiferidae, fresh-

water mussels.

INTRODUCTION

No equal area on earth has such a diversity of Naiad life

or such magnificent shells. Here are found the largest

species in the world; here are forms with knobs, pustules,

angles, lobes, and concentric sculpture. The nacre of

many of them is wonderfully rich in tints ot silver, pink,

purple, salmon or red, and it is equaled in beauty by the

elegant patterns of e.xternal painting, in stripes and mot-

tlings and delicate hair lines.

Charles T. Simpson (1896) on the Lhiionoidea of the

Mississippi Valley

The distributions of the 46 species of freshwater mussels

(Bivalvia; Unionoidea) that inhabit Minnesota were pre-

sented by Charlotte Webster Davvley in her doctoral

thesis (1944) and a widely cited paper (Dawley, 1947).

However, additional data have accumulated in the near-

ly half century since then. This paper seeks to update

the literature record and establish the known distribu-

tions of these mollusks.

The life-cy cle of a freshwater mussel involves the use

of one or more species of fish or, in one case, an am-

phibian as a host for the parasitic larva or glochidium.

In general, the major phase of dispersal occurs via par-

asitism of their host. Thus, barriers that inhibit the va-

gility of fish (drainage divides, waterfalls, etc.) also block

the dispersal of the Unionoidea.

Hydrology of Minnesota. Minnesota's surface area of

218,500 km- is drained by three divergent watersheds

(Figure 1 ). The Lake Superior Sy stem, the smallest, drains

to the St. Lawrence River via Lake Superior. It is defined

as the lake itself and all its Minnesota tributaries. This

system drains about 15,300 km- or 7% of the state.

The next largest watershed, draining 76,500 km- or

35% of the state, runs to Hudson Bay and is subdivided

into the Red River System (the Red River of the North

and all its Minnesota tributaries) and the Lake of the

Woods System (the Rainy Ri\ er and all Minnesota waters

draining to Lake of the Woods). The two systems are

confluent at Lake Winnipeg, Manitoba, but because they

have different mussel faunas, they are considered sepa-

rately.

The remaining 126,700 km- (58''() drains to the Gulf

of Mexico via the Mississippi River. The basin is subdi-

vided into the Upper and Lower Mississippi River Sys-

tems (following L'nderhill, 1989) based on their divergent

histories and \astly disparate mussel faunas. The Upper

Mississippi River System is that portion of the Mississippi

River and all of its tributaries above the Falls of St.

Anthony at Minneapolis. The Lower Mississippi River

System is the river and all of its Minnesota tributaries

below the Falls of St. Anthony. This includes not only

those watersheds draining directly into the Mississippi

River in Minnesota (the St. Croix, Minnesota, Zumbro,

Cannon, and Root River Watersheds), but also those wa-

ters in southwestern Minnesota draining to the Missouri

and Des Moines Rivers.

For the purposes of this study, state boundaries formed

by streams extend to the opposite bank to relax possible

political tensions among mussels on opposing sides of the

same river.

These five drainage systems were in place following

the final northward retreat of Glacial Lake Agassiz. Be-

fore that time, the enormous volumes of meltwater left

Page 46

THE NAUTILUS, Vol. 110, No. 2

Figure 1. The Drainage Systems of Minnesota. The Red River

System (RRS) and Lake of the Woods System (LVVS) are con-

fluent at Lake Winnipeg, eventually emptying to Hudson Bay.

The Lake Superior System (LSS) drains to the Great Lakes.

The remainder of the state drains via the Mississippi and is

comprised of the Upper Mississippi River System (UMRS) and

Lower Mississippi River, Minnesota River, and St. Croi.x River

subsystems (LMRS, MRS, and SCRS, respectively). See te.xt for

e.xplanation of the state s drainage systems.

in the wake of wasting glaciers over-ran present drainage

divides and connected now dissociated basins. At such

times, unionoids were free to migrate up the Mississippi

River from southern refugia (Johnson, 1980) and into

adjacent basins until water levels receded.

MATERIALS AND METHODS

Species have been assigned to watersheds using not only

specimens housed in the University of Minnesota's James

Ford Bell Museum of Natural History (JFB) Invertebrate

Collection but al.so from an extensive literature survey.

In the Appendi.x, for each species, a single lot is listed

for each basin when available, generally those used by

earlier authors.

The following literature references where utilized to

establish the distributions of the Unionoidea in each of

the above defined Minnesota drainage systems. No ar-

cheological or fossil literature has been included.

LakeSuperiorSystem;Dall(1905), Dawley (1944, 1947),

Goodrich and van der Schalie (1932: Lake Superior in

general), Moyle (1947), Walker (1913).

Lake of the Woods System: Baker (1929, 1935), Clarke

(1973), Dawley (1944, 1947).

Red River System: Baker (1929), Clarke (1973), Coker

and Southall (1915), Cvancara (1966, 1967, 1970, 1977,

1979, 1983), Cvancara et al. (1981), Daniels (1909),

Dawley (1944, 1947), Grant (1885), MN DNR (1984,

1986), Radke (1992), Sargent (1895). Wilson and

Danglade (1914).

Upper Mississippi River System: Baker (1929), Bright

(1988), Dawley (1944, 1947), Grant (1885), Movie

(1940), Sargent (1895), Wilson and Danglade (1914).

Lower Mississippi River Subsystem: Baker (1928), Bright

(1988), Bright ct al. (1989),' Coon et al. (1977), Cooper

(1834), Davis (1990), Dawley (1944, 1947), Fuller (1978,

1980), Grant (1885), Grier (1922, 1926), Grier and

Mueller (1922); Havlik (1981), Holzinger (1887),

Hornbachef al. (1992), Johnson (1980), Mathiak (1979),

Nachtrieb (unpublished), Southall (1925), Thiel (1981),

van der Schalie and van der Schalie (1950), Wilson

and Danglade (1914).

Minnesota River Subsystem: Bright et al. (1990), Dawley

(1944, 1947), Grant (1885). MN DNR (1985), Nach-

trieb (unpublished), Wilson and Danglade (1914).

St. Croix River Subsystem: Baker (1928), Bright (1988),

Cooper (1834), Dawley (1944, 1947), Fuller (1978,

1980), Imlay (1972), Mathiak (1979), Stern (1983),

Wilson and Danglade (1914).

Wilson and Danglade (1912) and Ellis (1931), provided

only vernacular names, but these have been reworked in

a scientific context by Wilson and Danglade (1914) and

van der Schalie and van der Schalie (1950), respecti\el\.

Grant (1887) reported collecting unnamed Anodonta and

Lampsilis from unspecified watersheds of the Arrowhead

Region of northeastern Minnesota. Few data are avail-

able on the Unionoidea of the Lake Superior System. Of

the general Great Lakes literature reviewed (e.g., van

der Schalie, 1961; Walker, 1913), most study has centered

on the eastern lakes, emphasizing Lake Erie. The ref-

erences cited aboxe provide almost no information on

the mussels found in the streams of the North Sliore of

Lake Superior (with the exception of the St. Louis River).

Smith and Moyle (1944), however, sur\e\ed the fauna

of these streams, including the macrobenthos, and re-

ported no unionoids. Since Moyle, in his other reports

(1940, 1947), has provided mussel data it can be assumed

that no unionoids are present in these streams.

For mussel distributions adjacent to Minnesota, the

reader is referred to the follow ing references: North Da-

kota: Cvancara (1966, 1967, 1970, 1975, 1976, 1983),

Cvancara et al. (1966, 1972, 1976); South Dakota: Coker

and Southall (1915), Over (1913); Canada: Clarke (1973,

1981); Wisconsin: Baker (1928), Havlik and Stansbery,

(1977), Mathiak (1979), Stern (1983).

RESULTS

Figure 2 presents the distributions of the Unionoidea in

Minnesota. For each species, the systematic assignment

follows Williams et al. (1993), which claims to provide

Daniel L. Graf, 1997

Page 47

Upper Mississippi River Faina —

iMsinifioiui tiimplanaia (B;irnes, IS23) |

Lasmi^ona contpri_'SSti (Lea. IS2'^)) |

Liunpsih:! ciirdium (Rafinesque. IS20) |

Pygdiioiioii grandis (Say. 1824) |

Ugiimia recta (Lamarck. 1X19) |

LumpsiUs siliquoidea (Barnes. 1823) |

AnodonioiJesfenissMioints {Lea. 18.'14) |

Slrophiuis iiiidiilcinis (Say. 1817) Q

UilerlnuUa imbecillis (Say. 1829) Q

D

Red River of the North Fai na

Piikimiliis aliiliis (Say. 1 8 1 7 ) □ □

l.usinigonu coskira (Ra{\ni:f.quc. 1820) Q [31

FiiscoiHiici Jlava (Rafinesque. 1820) Q Q

Ambleniii piuala (Say. 1817)0] CI

yi/<i</n(/(M/»i/(/M(/<; (Rafine.sque. 1820) Q Q

Dl

Elliptu>

Lake Superior Fauna

omphiiuna (Lightfool. 1786) |

D D nnnn

_ -Specimen with LiteraUire Citalicni or

VoLiclier only

H Literature Citation only

\T\ Problematical Record (see te.xt)

□ Not Reported

Lower Mississippi River Faina

Sinipsontilas tunhigiui (Say. 1825)

I'oHiiiidus capax (Green. 1832)

Pli'iiiiibinui ciHiineum (Conrad, 1836)

Anidens ciinpdgiisiis (Say, 1819)

Elliplid cnissidtns (Lamarck. 1819)

PUiliiihasus cyphyiis (Rafinesque. 1820)

Eihptui dilataiiis (Rafinesque. 1820)

Tnincillii do/Hicifoimis (Lea. 1828)

FiiscdiHiui fheiHi (Lea. 1831 )

Vi'iuiskuonclui ellipsifdiniis (Conrad. 1836)

Leplodcii Jrcigilis (Rafinesque. 1820)

QiiadndajragDSci (Conrad. 1836)

Liiinpsilis higgiiisi (Lea. 1857)

Airniiiiiiiuis ligamenlina (Lamarck. 1819)

Ellip.stiriii llneiiliila (Rafinesque. 1820)

Alasiiiulanki iihiri^iiuiki Say 1819

QiuidntUi mekinevw (Rafinesque. 1820)

CiimherUindia nuinodonui (Say. 1829)

Mfgalamuas nenosii (Rafinesque. 1820)

Qiiiidnilu nitdiilaki (Rafinesque. 1820)

Ptiranuliis (ihieiisis (Rafinesque. 1820)

I Ihiivcinci olivtirui (Rafinesque, 1820)

Ttixitltisina purvtis (Banies, 1823)

Qiiadruhi pustuUtsci (Lea. 1831)

Oblicpuiriu njh'xii Rafinesque. 1820

Anddiinia subdibuuUiki Say 1831

Lanipsitis teres (Rafinesque, 1820)

Epiiiblasmu irkptetra (Rafinesque, 1820)

TrunciUii Inincuia (Rafinesque. 1820)

Cyclimaias iiibcriiiUita (Rafinesque. 1820)

Tntiifiiiiiia tcrnuiisti (Rafinesque. 1820)

Figure 2. The Distribution of L'nionoid P^aunas in Minnesota.

the same list as Turgeon et al. (1988) but with spelling

and other such errors corrected. Figures depicting these

mollusks can be found in Cummings and Mayer (1992)

and Fuller (1985). The only exception, Elliptio com-

planala (Lightfoot, 1786), is illustrated in Clarke (1973:

56-57, plate 3). The species list includes only those spe-

cies verified to occur in Minnesota; distributions pre-

sented exclude erroneous records (see Problematical Re-

cords below).

The Lower Mississippi Drainage System is divided into

three subsystems: the Lower Mississippi River Subsystem,

excluding the Minnesota and St, Croix River watersheds;

the Minnesota River Subsystem, w hich includes the main

stem of the river and all its tributaries; and the St. Croix

River Subsystem, also including the main stem of that

river and its tributaries.

Questionable and Problematical Records. Problemati-

cal records, both from literature reports and JFB speci-

mens, may lead to false hy potheses; those identified by

the author are summarized and dealt with below .

Questionable Literature Records. Four species reported

from the Red River System are supported by only a single

literature citation each: Qtiadrula pustulosa by Coker

and Southall (1915), Pleiirobetna coccineum by Wilson

and Danglade (1914), Elliptio dilatata by Daniels (1909),

and Obliquaha reflexa h\ Dawley (1944, 1947). How-

ever, no corroborating voucher specimens exist at the

JFB and none of these species were collected by Cvancara

(1966, 1967, 1970, 1975, 1976, 1983; Cvancara et al,

1966; 1972; 1976) or others (e.g. Clarke, 1973) who have

extensively surveyed basin. Qiiadrula pustulosa, P. coc-

Page 48

THE NALTILLS. Vol. 110. No. 2

cineum, and E. dilatata are likely incorrectly identified

Q. quadrula, Fusconaia flava. and Ligumia recta, re-

spectively. In the case of O. reflexa, both Cvancara (1970)

and Clarke (1973) reported personal communications with

Da\vle\ in which she admitted her misidentification. The

true distributions of these four species are probably lim-

ited to onl\- the Lower Mississippi River System.

Problematical Voucher Specimens. The presence of

Tritogonia verrucosa in the Upper Mississippi River Sys-

tem is indicated by an unpaired valve (JFB 3190) col-

lected b> John Moyle (1940). Mo\le believed the valve

to be evidence of Lower Mississippi species overcoming

the Falls of St. Anthony but not surviving because of

poor habitat. Dawley (1944, 1947) considered the pres-

ence of Tritogonia to be an accident, and her opinion is

likeK correct. Thus, the distribution of T. verrucosa is

limited to only the Lower Mississippi River System. An

unpaired valve of Amblema plicata (JFB 7567) was col-

lected from the Cloquet River in the Lake Superior Sys-

tem. However, since A. plicata has otherwise not been

reported from that system in Minnesota or even from

the Lake Superior Basin in general, this single specimen

is probably not a remnant of a population and is assumed

to be trash or the remains of some introduced individual

or individuals. Amblema plicata has been reliably re-

ported from only the Red and Lower Mississippi River

Systems, and this presumably reflects the mussel's true

distribution within the state.

The problematical distribution of Acttnonaias liga-

mentina in the Minnesota has been addressed by Clarke

(1973). Dawley "s Red River specimen could not be lo-

cated, but records of A. ligamentina reported by her

(1944, 1947: JFB 2863), Moyle (1940: JFB 2872), and

Radke (1992: JFB 7569) have been re-identified as Lamp-

silis siliquoidea. Cvancara (1970) disregarded Dawley "s

Red River voucher, reporting that in a personal com-

munication, she agreed that her record was probably an

error. Thus, Actinonaias is limited to only the Lower

Mississippi River System.

The single lot, JFB 2332 Lampsilis teres, labeled as

coming from Leech Lake in the Upper Mississippi River

System is likely the result of a cataloging error. Since

Dawley (1944, 1947) apparently ignored it, the validity

of this record is questionable.

Other Problematical Data. Utterbackia irnbecillis has

never been reported from any of the Lake Superior or

Hudson Bay drainage systems. This makes L'. irnbecillis

unique among species reliably reported from the Upper

Mississippi Ri\er System. Also making V. irnbecillis dis-

tinctive is the fact that it mav complete its reproductive

cycle with or without a glochidial host (Dawley, 1944;

Fuller, 1985). This would hamper its upstream vagility,

and perhaps 17. imbecillis could not extend beyond the

Upper and Lower Mississippi River Systems before drain-

age connections were severed.

The presence of Anodontoides ferussacianus in the

Lake of the Woods S\stem is not supported by a specimen

in the JFB, though both Baker (1935) and Dawley (1944,

1947) reported the species from that basin.

Strophitus undulatus has never been reported from

the Lake Superior System and its presence in the Lake

of the Woods System is supported by only a single lit-

erature record: Clarke (1973). Dawley (1944, citing Le-

fevre & Curtis, 1911) noted that S. undulatus is another

mussel that can complete its life-cycle independent of a

glochidial host. However, the mussel is not obligately

anomalous in its reproduction; its widespread distribu-

tion indicates that it probably is dispersed by fish at least

some of the time. Three of its published glochidial hosts

(Hoggarth, 1992) are found in the that watershed as well

as Minnesota's four other drainage systems (Underbill,

1989), providing a means of statewide distribution. The

apparent absence of S. undulatus may actually be the

result of the lack of an adequate surve> of the St. Louis

River System.

DISCUSSION

There is a pattern to the distributions of the Unionoidea

in Minnesota. Species with similar distributions can be

grouped into faunas; members of each fauna are h\-

pothesized to have shared modes and tempos of dispersal

into the state s drainage systems. Ninety-eight percent

of the species (45 of 46) are found in the Lower Missis-

sippi River System, supporting the hypothesis that the

freshwater mussel fauna oi Minnesota dispersed north-

ward from southern refugia by way of the Mississippi

River at the close of the Pleistocene (Johnson, 1980). The

remaining species, Elliptio complanata, appears to have

migrated into the state via Lake Superior from the St.

Lawrence watershed.

Of the 45 species of unionoids reported from the Low-

er Mississippi River System, 30 have been reliably re-

corded from onl\ that system; these mussels constitute

the Lower Mississippi River Fauna (Figure 2). Much of

this fauna is made up of mussels ecologically limited to

larger rivers that were unable to extend their ranges

beyond the Lower Mississippi and the lowest reaches of

the Minnesota and St. Croix Ri\ ers (e.g., Fusconaia ebena,

Ellipsaria lineolata). Others of the species characteristic

of the Low er Mississippi River System ma\ be relatively

recent arrivals to the state (e.g., Anodonta suborbiculata,

Quadrula nodulata) since the> were not reported by

earlier authors.

Those freshwater mussels commonly found above the

Falls of St. .^nthonx' are collectively know n as the Upper

Mississippi River Fauna (Figure 2). The Upper Missis-

sippi Ri\ er Fauna must ha\ e been present at a time w hen

all ot the states watersheds were connected. Of the 9

species that belong to this assemblage, only Utterbackia

imbecillis and Strophitus undulatus have not been re-

ported from all five of Minnesota s drainage systems (dis-

cussed above).

Five mussel species are found oni\ in the Red and

Lower Mississippi River Systems; these belong to the Red

River of the North Fauna (Figure 2). .\t the close of the

Daniel L. Graf. 1997

Page 49

Pleistocene, with the formation of Glacial Lake Agassiz,

these t\\o drainage SNStems were connected more than

once, and this allowed the dispersal of the Red Ri\er of

the North Fauna from the Minnesota River basin into

that of the Red River of the North.

Elliptio complanata is the only unionoid of the Lake

Superior Fauna, It is known oiiK' from the Lake Superior

System. It has probabK migrated into Minnesota in the

last 80 years or so from the eastern Great Lakes.

The problematical species discussed above are difficult

for exactly the reason that the\ don not fit neatly into

any of the four described unionoid faunas. However,

except for the case of Actinonaias ligarnentina. these

problems are the result of single specimens.

Further study of the unionoid faunas of the Lake of

the Woods, Lake Superior, and Upper Mississippi River

systems promises to advance our knowledge on the dis-

tributions of freshw ater mussels of the Minnesota region.

ACKNOWLEDGMENTS

This paper is dedicated to my mentor and friend, the

late Dr. Robert C. Bright of the James Ford Bell Museum

of Natural History, University of Minnesota. Dr. Bright

granted me the encouragement, resources, and freedom

to completeK reorganize and computerize the museum's

unionoid collection; this paper is the result. Professor

James C. Underhill, also of the Bell Museum, provided

invaluable intellectual and moral support. N'ersions of

this manuscript were criticalK reviewed by J.C. Under-

hill, E. Ruber of Northeastern University, and K.J. Boss

and R.I. Johnson of the Museum of Comparative Zoology;

all provided worthwhile input. Two anonymous review-

ers suggested numerous improvements. Thanks also to

C. Gatenby of Virginia Polytechnic Institute for iden-

tifying those specimens that I was unable to acquire, T.

Deneka of Macalester College for providing data on spec-

imens he has collected, and all others w ho have assisted

me directK and indirectly in completing this study.

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APPENDIX

Figure Citations and Specimens E.xamined

Provided below are figure references and the James Ford

Bell Museum of Natural History (JFB) specimens used

to construct Figure 2. For each lot, its catalog number

and collection locality are provided. Whenever possible,

specimens cited by previous authors have been selected,

and these references are cited below. Also provided are

corrections and re-identifications with regard to JFB

specimens of historical significance.

The following drainage system abbreviations are used:

LMRS = Lower Mississippi River Subsystem

LSS = Lake Superior System

LWS = Lake of the Woods System

MRS = Minnesota River Subsystem

RRS = Red River System

SCRS = St. Croi.x River Subsystem

UMRS = Upper Mississippi River System

Actinonaias ligamcntina (Lamarck, 1819). Figures:

Cummings & Mayer (1992:105), Fuller (1985:36). LWS:

Dawley's (1944, 1947) Sturgeon River record is JFB 2863

Lampsilis siliquoidea. RRS: Radke's (1992) Otter Tail

River record is JFB 7569 Lampsilis siliquoidea. I'MRS:

Dawley (1944, 1947) and Moyle's (1940) Crow Wing

River record is JFB 2872 Lampsilis siliquoidea. LMRS:

JFB 2847 Mississippi River, Dresbach, Winona Co.: Daw-

ley (1944, 1947), Grant (1885). MRS: JFB 2859 Minnesota

River, New Ulm, Brown Co.: Dawley (1944, 1947). SCRS:

JFB 2878 Kettle River, Rutledge, Pine Co.: Dawlev (1944,

1947).

Alasmidonta marginata Sa\, 1819. Figures: Cummings

& Mayer (1992:85), Fuller '(1985:42). LMRS; JFB 2766

Mississippi River, Winona, Winona Co.: Dawley (1944,

1947), Holzinger (1887). MRS: JFB 2763 Minnesota Riv-

er: Dawley (1944, 1947). SCRS: JFB 2769 St. Croi.x River

at the mouth of Sunrise Ri\er, Chisago Co.; Dawley

(1944, 1947).

Amblema plicata (Saw 1817). Figures: Cummings &

Mayer (1992:41), Fuller (1985:50). LSS; JFB 7567 Clo-

quet River, Brimson, St. Louis Co. RRS: JFB 3105 Red

River, 20 mi. N of Breckenridge, Wilkin Co.; Dawley

(1944, 1947), Grant (1885). LMRS; JFB 3083 Mississippi

River, Dresbach, Winona Co.; Dawlev (1944, 1947). Grant

(1885). MRS; JFB 3093 Minnesota River. Ft. Snelling;

Dawley (1944, 1947), Grant (1885). SCRS; JFB 3112 St.

Croix River at the mouth of Sunrise River, Chisago Co.:

Dawley (1944, 1947).

Anodonta suhorhiculata Sav, 1831. Figures; Cummings

& Mayer (1992:77), Fuller (1985:58).

Anodontoidcs ferussacianus (Lea, 1834). Figures; Cum-

mings k Maver (1992:81), Fuller (1985:57). LSS; JFB

2709 Cloquet River, St. Louis Co. RRS; JFB 6582 Cor-

morant Lake, Becker Co.; Radke (1992). UMRS; JFB

3995 Rum River, Anoka, Anoka Co.; Dawley (1944, 1947),

Grant (1885). LMRS; JFB 2717 Cedar River, Austin,

Mower Co.; Dawley (1944, 1947). MRS: JFB 2742 Cot-

tonwood River, Garvin, Lyon Co.; Dawlev (1944). SCRS:

JFB 2753 Rush Creek; Dawley (1944, 1947).

Arcidcns confragosus (Sav, 1819). Figures; Cummings

& Mayer (1992:89), Fuller (1985:51). LMRS; JFB 2771

Mississippi River, Red Wing, Goodhue Co.; Dawlev (1944,

1947). MRS; JFB 2773 Miimesota River, Ft. Snelling:

Dawley (1944, 1947), Grant (1885).

Cumherlandia monodonta (Say, 1829). Figures; Cum-

mings & Mayer (1992:23), Fuller (1985:16). SCRS; JFB

6690 Rush Creek, Rush City, Chisago Co.

Page 52

THE NAUTILUS, Vol. 110, No. 2

Cyclonaias tuberculata (Rafinesque, 1820). Figures:

Cummings & Mayer (1992:49), Fuller (1985:10). LMRS:

JFB 3198 Mississippi River, Ft. Siielling: Dawley (1944,

1947), Grant (1885). SCRS: JFB 3199 St. Croix River at

the mouth of Sunrise River, Chisago Co.: Dawley (1944,

1947).

Ellipsaria lineolata (Rafinesque, 1820). Figures: Cum-

mings & Mayer (1992:107), Fuller (1985:45). LMRS: JFB

2916 Mississippi River, Nininger, Dakota Co.: Dawley

(1944, 1947). MRS: JFB 2915 Minnesota River, Ft. Snell-

ing: Dawley (1944, 1947), Grant (1885). SCRS: JFB 4200

St. Croix River, Taylors Falls, Chisago Co.

Elliptio complanaia (Lightfoot, 1786). Figures: Clarke

(1973:56-57, plate 3). LSS: JFB 3258 Lake Superior, Du-

luth, St. Louis Co.: Dawley (1944, 1947).

Elliptio crassidens (Lamarck, 1819). Figures: Cummings

& Mayer (1992:67), Fuller (1985:14). LMRS: JFB 3249

Lake IPepin, Goodhue Co.: Dawley (1944, 1947). MRS:

JFB 3245 Minnesota River: Dawley (1944, 1947).

Elliptio dilatata (Rafinesque, 1820): the spike. Figures:

Cummings & Mayer (1992:69), Fuller (1985:15). LMRS:

JFB 3225 Mississippi River, Winona, Winona Co.: Daw-

ley (1944, 1947), Holzinger (1887). MRS: JFB 3242 Min-

nesota River, Ft. Snelling: Dawlev (1944, 1947), Grant

(1885). SCRS: JFB 3244 St. Croix River, Pine Co.: Dawley

(1944, 1947).

Epioblasma triquetra (Rafinesque, 1820). Figures: Cum-

mings & Mayer (1992:163), Fuller (1985:44). SCRS: JFB

5701 St. Croix River, Sand Is., Chisago Co.

Fiisconaia ebena (Lea, 1831). Figures: Cummings &

Mayer (1992:43), Fuller (1985:12). LMRS: JFB 3057 Mis-

sissippi River, Dresbach, Winona Co.: Dawley (1944,

1947), Grant (1885). MRS: JFB 6437 Minnesota River,

Scott Co.: Bright et al. (1990). SCRS: JFB 4044 Lake St.

Croix, Washington Co.

Fitsconaia flava (Rafinesque, 1820). Figures: Cummings

& Mayer (1992:47), Fuller (1985:11). RRS: JFB 3014 Red

River, 20 mi. N of Breckenridge, Wilkin Co.: Dawley

(1944, 1947), Grant (1885). LMRS: JFB 3045 Mississippi

River, Winona, Winona Co.: Dawley (1944. 1947). MRS:

JFB 3044 Minnesota River, Ft. Snelling: Dawley (1944,

1947), Grant (1885). SCRS: JFB .3047 St. Croix River,

Marine on St. Croix, Washington Co.: Dawley (1944,

1947).

Larnpsilis cardiuni (Rafinesque, 1820). Figures: Cum-

mings & Mayer (1992:157), Fuller (1985:28). LSS: JFB

2448 Lake Superior, Minnesota Point, Duluth, St. Louis

Co.: Dawley (1944, 1947). LWS: JFB 2411 Sturgeon Riv-

er, Little Fork River watershed, St. Louis Co.: Dawley

(1944, 19471 RRS: JFB 2396 Red River, 20 mi. N of

Breckenridge, Wilkin Co.: Dawlev (1944, 1947), Grant

(1885). UMHS: JFB 2400 Mississippi River, Brainerd,

Crow Wing Co.: Dawley (1944, 1947), Grant (1885).

LMRS: JF"B 2397 Vli,ssissippi River, Dresbach, Winona

Co.: Dawley (1944, 1947), Grant (1885). MRS: JFB 2404

Minnesota River, Ft. Snelling: Dawlev (1944, 1947), Grant

(1885). SCRS: JFB 2415 St. Croix River, Marine on St.

Croix, Washington Co.: Dawley (1944, 1947).

Larnpsilis higginsi (Lea, 1857). Figures: Cummings &

Mayer (1992:153), Fuller (1985:33-34). LMRS: JFB 2454

Mississippi River, Dresbach, Winona Co.: Dawle\ (1944,

1947). MRS: JFB 6396 Minnesota River, Carver Co.:

Bright pf a/., 1990 (cf. L. /ugginsi); Dawley's (1944, 1947)

record is JFB 2455 Obovaria olivaria. SCRS: JFB 2453

Lake St. Croix, Washington Co.: Dawlev (1944, 1947).

Larnpsilis siliquoidea (Barnes, 1823). Figures: Cum-

mings & Mayer (1992:151), Fuller (1985:29). LSS: JFB

2291 Cloquet River, St. Louis Co.; Dawley (1944, 1947).

LWS: JFB 2983 Sturgeon River, Little Fork River wa-

tershed, St. Louis Co.: Dawley (1944, 1947). RRS: JFB

2305 Red River, 20 mi. N of Breckenridge, Wilkin Co.:

Dawley (1944, 1947), Grant (1885). UMRS: JFB 2242

Mississippi River, Brainerd, Crow Wing Co.: Dawley

(1944, 1947), Grant (1885). LMRS: JFB 2307 Rolling-

stone Creek, Minnesota City, Winona Co.: Dawley (1944,

1947), Grant (1885). MRs! JFB 2253 Minnesota River,

Ft. Snelling: Dawley (1944, 1947), Grant (1885). SCRS:

JFB 2261 St. Croix River at the mouth of Sunrise River,

Chisago Co.: Dawley (1944, 1947).

Larnpsilis teres (Rafinesque, 1820). Figures: Cummings

& Mayer (1992:149), Fuller (1985:31). UMRS: JFB 2232

Leech Lake, Cass Co. LMRS: JFB 2239 Mississippi River,

Red Wing, Goodhue Co.: Dawley (1944, 1947). MRS:

JFB 2236 Minnesota River, Ft. Snelling: Dawley (1944,

1947), Grant (1885).

Lasmigona complanata (Barnes, 1823). Figures: Cum-

mings & Mayer (1992:93), Fuller (1985:39). LSS: JFB

3308 Lake Superior, Minnesota Point, St. Louis Co.: Daw-

ley (1944, 1947); Baker (1928): reporting Lea's type-

locality. LWS: JFB 3305 Sturgeon River, Little Fork

River watershed, St. Louis Co.: Dawley (1944, 1947).

RRS: JFB 3302 Red Lake River, Crookston, Polk Co.:

Dawley (1944, 1947). UMRS: JFB 7507 Mississippi River,

Little Falls public access, Morrison Co. (collected by

Bright et al.); reported by Dawley (1944) & Grier &

Mueller (1922) that Wilson & Danglade (1914) found it

in the I'MRS; however, the data can not be located in

that publication. LMRS: JFB 3301 Mississippi River, Min-

neiska, Wabasha: Dawley (1944, 1947). MRS: JFB 3299

Minnesota River, Ft. Snelling: Dawley (1944, 1947), Grant

(1885). SCRS: JFB 3289 Lake St. Croix: Dawley (1944,

1947).

Lasmigona compressa (Lea, 1829). Figures: Cummings

& Maver (1992:97), Fuller (1985:41). LSS: JFB 3262 Clo-

quet River, St. Louis Co.: Dawley (1944, 1947). LWS:

JFB 3261 Fall Lake, Lake Co.: Dawley (1944). RRS: JFB

7196 Otter Tail River, Otter Tail Co. (collected by Bright

etal.). UMRS: JFB 3265 Sauk River, Stearns Co.: Dawley

(1944, 1947), Movie (1940). LMRS: JFB 5251 North

Branch Middle Fork Zumbro River, Highway 57 Bridge,

Dodge Co.: Bright ct al. (1989). MRS: JFB 6017 Min-

Daniel L. Graf, 1997

Page 53

nesota River, Yellow Medicine Co.: Briglit ct al. (1990).

SCRS. JFB 3266 St. Croix River at the mouth of Sunrise

River, Chisago Co.: Dawley (1944, 1947).

Lasmigona costata (Rafinesque, 1820). Figures: Cum-

mings & Mayer (1992:95), Fuller (1985:4o"). RRS: JFB

3279 Red Lake River, Crookston, Polk Co.: Dawley (1944,

1947). LMRS: JFB 3280 Mississippi River, Winona, Wi-

nona Co.: Dawle\ (1944, 1947). MRS: JFB 3274 Min-

nesota River: Dawley (1944, 1947). SCRS: JFB 4003 Snake

River at the mouth of the Groundhouse Ri\er, Kanabec

Co.

Leptodca fragilis (Rafinesque, 1820). Figures: Cum-

mings & Maver (1992:121). Fuller (1985:20). LMRS: JFB

2919 Lake Pepin: Dawley (1944, 1947), MRS: JFB 2923

Minnesota River, Ft. Snelling: Da\\ lev (1944, 1947), Grant

(1885). SCRS: JFB 4613 St. Croix River, Marine on St.

Croix, Washington Co.

Ligumia recta (Lamarck, 1819). Figures; Cummings &

Maver (1992:137), Fuller (1985:18). LSS: JFB 2205 Lake

Superior: Dawley (1944, 1947). LWS: JFB 2214 Lake of

the Woods, Pine Is., Lake of the Woods Co.: Dawley

(1944). RRS: JFB 2201 Red River, 20 mi, N of Breck-

enridge, Wilkin Co.: Dawley (1944, 1947), Grant (1885).

UMRS: JFB 2994 Mississippi' River, Brainerd, Crow Wing

Co.: Dawley (1944, 1947), Grant (1885). LMRS: JFB 2989

Vlississippi River, Winona, Winona Co.: Dawlev (1944,

1947), Holzinger (1887). MRS: JFB 2998 Blue Earth Riv-

er: Dawley (1944, 1947). SCRS: JFB 2213 Kettle River,

Rutledge, Pine Co.: Dawley (1944, 1947).

Mcgalonaias nervosa (Rafinesque, 1820). Figures: Cum-

mings & Mayer (1992:25), Fuller (1985:52). LMRS: JFB

3071 Mississippi Ri\er, Red ^^ ing, Goodhue Co.: Dawley

(1944, 1947). MRS: JFB 3065 Minnesota River, Cedar

Ave., Hennepin Co.: Dawley (1944, 1947). SCRS; JFB

3063 St. Croix River, Hudson, St. Croix Co., Wisconsin:

Dawley (1944, 1947). Lower Mississippi River below

Minnesota: Baker (1903).

Obliqiiaria reflexa Rafinesque, 1820. Figures: Cummings

& Mayer (1992:101), Fuller (1985:48). LMRS: JFB 2808

Lake Pepin, Lake City, Wabasha Co.: Dawley (1944,

1947), Grant (1885). MRS: JFB 2489 Minnesota River,

Ft. Snelling: Dawley (1944, 1947), Grant (1885). SCRS:

JFB 2804 Lake St. Croix, Washington Co.: Dawlev (1944,

1947).

Obovaria olivaria (Rafinesque, 1820). Figures: Cum-

mings & Mayer (1992:109), Fuller (1985:35). LMRS: JFB

2829 Mississippi River, Dresbach, Winona Co.: Dawle\

(1944, 1947), Grant (1885). MRS: JFB 6520 Minnesota

River, Pike Is., Hennepin Co : Bright et ai. 1990; Daw-

ley's (1944, 1947) record is JFB 2832 Fusconaia ebena.

SCRS: JFB 2826 St. Croix River at the mouth of Sunrise

River, Chisago Co.; Dawle> (1944, 1947).

Plethobasns cyphyus (Rafinesque, 1820). Figures: Cum-

mings & Mayer (1992:51), Fuller (1985:49). LMRS; JFB

3203 Mississippi River, Dresbach, Winona Co.; Dawley

(1944, 1947), Grant (1885).

Pleurobema coccineum (Conrad, 1836). Figures: Cum-

mings & Mayer (1992:59), Fuller (1985:13). LMRS; JFB

3212 Lake Pepin, Goodhue Co.; Dawley (1944, 1947).

MRS; JFB 6068 Minnesota River, Chippewa Co.; Bright

etal. (1990). SCRS; JFB 3219 Snake River: Dawley (1944,

1947).

Potainilus alatiis (Say, 1817). Figures: Cummings &

Mayer (1992:125), Fuller (1985:25). RRS; JFB 2946 Red

River, 20 mi. N of Breckenridge, Wilkin Co.; Dawley

(1944, 1947), Grant (1885). LMRS; JFB 2945 Mississippi

River, Dresbach, Winona Co.; Dawley (1944, 1947), Grant

(1885). MRS: JFB 2941 Minnesota River, Ft. Snelling;

Dawley (1944, 1947), Grant (1885). SCRS; JFB 2957

Snake Ri\er, Kanabec Co.: Dawley (1944, 1947).

Potamilus capax (Green, 1832). Figures; Cummings &

Mayer (1992:129), Fuller (1985:27). LMRS; Johnson

(1980) reporting Green's type locality at St. Anthony

Falls, Minneapolis. Lov\er Mississippi Ri\er below Min-

nesota; Grier & Mueller (1922:9); ". . .the consensus of

opinion is that it ordinariK does not go much north of

Davenport, Iowa.

Potamilus ohiensis (Rafinesque, 1820). Figures: Cum-

mings & Mayer (1992:123), Fuller (1985:22-23). LMRS

JFB 2963 Mississippi River, Red Wing, Goodhue Co.

Dawley (1944, 1947). MRS; JFB 2964 Minnesota River

Ft. Snelling; Dawlev (1944, 1947); Grant's (1885) record

is JFB 2920 Leptodea fragilis. SCRS; JFB 5543 St. Croix

River, Interstate Park, Chisago Co., Minnesota; Dawley s

(1944, 1947) record is JFB 2962 Leptodea fragilis.

Pyganodon grandis (Sav, 1829). Figures: Cummings &

Mayer (1992:79), Fuller'(1985;60). LSS; JFB 2471 Com-

stock Lake, St. Louis Co.; Dawley (1944, 1947). LWS:

JFB 2625 Sturgeon Lake, St. Louis Co.; Dawlev (1944,

1947). RRS; JFB 2568 Red Lake River. Crookston, Polk

Co.; Dawley (1944, 1947). UMRS; JFB 2560 Rum River,

Milaca, Mi'Ue Lacs Co.; Dawlev (1944, 1947), Movie

(1940). LMRS; JFB 3327 Zumbro River, Wabasha, Wa-

basha Co.; Dawley (1944, 1947), Grant (1885). MRS; JFB

2664 Minnesota River, Ft. Snelling; Dawley (1944, 1947),

Grant (1885). SCRS; JFB 2660 O.xbow Lake, Marine on

St. Croix, Washington Co.; Dawley (1944, 1947).

Qnadrula fragosa (Conrad, 1836). Figures: Cummings

& Mayer (1992:29). LMRS; JFB 3127 Mississippi River,

Nininger, Dakota Co. [collected November, 1886 by

Winchell; reported as Q. quadrula by Dawley (1944,

1947)]. MRS; JFB 6356 Minnesota River,'Sibley Co.; Bright

et al. (1990).

Quadrula metaixevra (Rafinesque, 1820). Figures; Cum-

mings & Mayer (1992:35), Fuller (1985:6). LMRS; JFB

3152 Mississippi River, Dresbach, Winona Co.; Dawley

(1944, 1947), Grant (1885). MRS; JFB 6357 Minnesota

River, Sibley Co.; Bright et al. (1990). SCRS; JFB 4607

St Croix River, Marine on St. Croix, Washington Co.

Quadrula nodulata (Rafinesque, 1820). Figures: Cum-

mings & Mayer (1992:37), Fuller (1985:8). MRS; JFB

6424 Minnesota River, Scott Co.; Bright et al. (1990).

Page 54

THE NAUTILUS, Vol. 110, No. 2

SCRS: JFB 4578 St. Croi.x River, Coppermine Dam,

Douglas Co., Wisconsin.

Quadriila pustiilosa (Lea, ISol). Figures: Cummings &

Mayer (1992:39), Fuller (1985:9). LMRS: JFB 3136 Mis-

sissippi Riser, Winona, Winona Co.: Dawley (1944, 1947),

Holzinger (1887). MRS: JFB 3144 Minnesota River, Ft.

Snelling: Dawley (1944, 1947). SCRS: JFB 3147 St. Croix

River, Marine on St. Croix, Washington Co.: Dawley

(1944, 1947).

Quadrula quadrula (Rafinesque, 1820). Figures: Cum-

mings & Mayer(1992:31), Fuller (1985:7). RRS:JFB3125

Red River, 20 mi. N of Breckenridge, Wilkin Co.: Daw-

ley (1944, 1947), Grant (1885). LMRS: JFB 3121 Missis-

sippi River, Red Wing, Goodhue Co.: Dawley (1944,

1947). MRS: JFB 3122 Minnesota River, Ft. Snelling:

Dawley U944, 1947), Grant (1885). SCRS: JFB 4911 St.

Croix River, Stillwater, Washington Co.

Simpsonaias amhigua (Say, 1825). Figures: Cummings

& Mayer (1992:91), Fuller (1985:56). MRS: JFB 6074

Minnesota River, Chippewa Co.: Bright et al., 1990. SCRS:

JFB 5063 St. Croix River, Marine on St. Croix, Wash-

ington Co.

Strophitus iindulatiis (Say, 1817). Figures: Cummings

& Mayer (1992:83), Fuller'(1985:61). RRS: JFB 2784 Red

River, 20 mi. N of Breckenridge, Wilkin Co.: Dawley

(1944, 1947), Grant (1885). UMRS: JFB 7415 Mississippi

River, Crow Wing State Park, Crow Wing Co. (collected

by Bright et al). LMRS: JFB 2793 Mississippi River, Red

Wing, Goodhue Co.: Dawley (1944, 1947). MRS; JFB

6075 Minnesota River, Chippewa Co.: Bright et al. (1990).

SCRS: JFB 2795 Grindstone River, Kettle River water-

shed. Pine Co.: Dawley (1944, 1947).

Toxolasma parvus (Barnes, 1823). Figures: Cummings

& Mayer (1992:131), Fuller (1985:54-55). LMRS: JFB

2974 Vlississippi River, Wacouta, Goodhue Co.: Dawley

(1944, 1947). MRS: JFB 2976 Minnesota River, Ft. Snell-

ing: Dawley (1944, 1947), Grant (1885). SCRS: JFB 2973

St. Croix River, Marine on St. Croix, Washington Co.:

Dawley (1944, 1947).

Tritogonia verrucosa (Rafinesque, 1820). Figures: Cum-

mings & Mayer (1992:27), Fuller (1985:53). UMRS: JFB

3190 Sauk River, St. Cloud, Stearns Co.: Bright (1988),

Dawley (1944), Moyle (1940). LMRS: JFB 3189 Missis-

sippi River, Dresbach, Winona Co.: Dav\ley (1944, 1947),

Grant (1885). MRS: JFB 3181 Minnesota River, Ft. Snell-

ing: Dawley (1944, 1947), Grant (1885). SCRS: JFB 3191

St. Croix River, Tavlors Falls, Chisago Co.: Dawley (1944,

1947).

Truncilla donaciformis (Lea, 1828). Figures: Cummings

& Mayer (1992:117), Fuller (1985:47). LMRS: JFB 4307

Mississippi River, Northern States Power Prairie Is. Nu-

clear Generating Plant, 1.5 miles upstream of Lock &

Dam #3, Goodhue Co. MRS: JFB 2933 Minnesota River,

Ft. Snelling: Dawley (1944, 1947), Grant (1885). SCRS;

JFB 4610 St. Croix River, Marine on St. Croix, Wash-

ington Co.

Trur}cilla truncate (Rafinesque, 1820). Figures: Cum-

mings & Mayer (1992:115), Fuller (1985:46). LMRS; JFB

2896 Mississippi River, Winona, Winona Co.; Dawley

(1944, 1947), Holzinger (1887), MRS: JFB 2897 Min-

nesota River, Ft. Snelling: Dawley (1944, 1947), Grant

(1885). SCRS: JFB 2900 St. Croix River, St. Croix Falls,

Polk Co., Wisconsin: Dawley (1944, 1947).

Utterbackia imbecillis (Sav, 1829). Figures: Cummings

& Mayer (1992:75), Fuller (1985:59). UMRS: JFB 2683

Mississippi River, Fridlev, .\noka Co.: Dawley (1944,

1947), Moyle (1940). LMRS; JFB 2682 Lake Pepin, Wa-

couta, Goodhue Co.; Dawley (1944, 1947). MRS; JFB

2679 Minnesota River, Ft. Snelling: Dawley (1944, 1947),

Grant (1885). SCRS: JFB 2768 Oxbow Lake, Marine on

St. Croix, Washington Co.: Dawley (1944, 1947).

Vcnustaconcha ellipsiformis (Conrad, 1836). Figures:

Cummings&Mayer(1992:141), Fuller (1985:38). LMRS;

JFB 5280 Cascade Creek, Zumbro River watershed,

Rochester, Olmsted Co.; Bright et al. (1989). SCRS; JFB

5736 St. Croix River, Polk Co., Wisconsin.

THE NAUTILUS 110(2):55-75, 1997

Page 55

The Life and Malacological Contributions of

R. Tucker Abbott (1919-1995)

M. G. Harasewych

Department of Invertebrate Zoology

National Museum of Natural History

Smithsonian Institution

Washington, DC 20560 U.S.A.

Dr. Robert Tucker Abbott, one of the best known and

most venerated figures in American Malacology, died

November 3, 1995 after a protracted battle with pul-

monary fibrosis. During his long and multi-faceted ca-

reer, he was an eminent researcher, curator, editor, mu-

seum director, educator, enterpreneur, and an ambas-

sador of his science to a broad and appreciative audience

of enthusiastic shell collectors and weekend naturalists.

R. Tucker Abbott was undoubtedly the most widely

known malacologist in the world. His unique career has

been chronicled like that of no other malacologist. Dur-

ing his life, articles and biographical sketches appeared

in Reader's Digest [August 1961], Hawaiian Shell News

\1(d)~ [1963] and again 24(3):3 [1976], Ttic Wall Street

Journal [March IS, 1994] reprinted in Ihiwaiian Shell

News 42(8):7-8 [1994] and Sniithsunian magazine [Au-

gust, 1995] as well as in numerous newpaper and shell

club articles. Following his death, countless tributes and

obituaries have been published, among them those ap-

pearing in American Conchologist . Journal of Shell-

fisheries Research and The Sew York Times. Most have

emphasized the latter portions of his career.

Born on September 28, 1919 in Waterlown, Massa-

chusetts, to Charles Matthew (a paint manufacturer) and

Frances (a homemaker, nee Tucker) Abbott, Tucker was

the third of four brothers. As a bo\. Tucker spent his

summers in Bermuda, where an uncle. Col. Richard

Tucker was Director of the Bermuda Biological Station.

It was there that he met and was befriended by Dr.

William Beebe, the ichthyologist and pioneer of deep-

sea exploration. Dr. Beebe was to have a profound influ-

ence on Tucker. Many aspects of Tucker's career as a

scientist, popularist, author, and celebrit\ find parallels

in the life of William Beebe.

When Tucker was a young boy, his family moved to

Canada. It was there that he and a friend built their first

natural history museum in the basement of the Abbott

home. Upon graduating from the Humberside Collegiate

in Toronto in 1938, Tucker entered Harvard College.

While at Harvard, Tucker met Dr. William J. Clench,

who was to become his lifelong friend and mentor.

Tucker's career as a malacologist began \\ hile he was

still an undergraduate at Harvard. He was a Research

Assistant in the Department of MoUusks, Museum of

Comparative Zoology, Harvard University from 1938-

1942. During this time he helped establish the mono-

graphic series Johnson ia. serving as its first Business Man-

ager, and published his first mongraphs and taxa (co-

authored with Clench) prior to graduation. He was given

leave from his studies to serve as the malacologist on the

Harvard-Archbold Expedition to Melanesia in 1940-41,

collecting mollusks in Fiji for three months aboard the

junk-yacht Cheng-Ho.

After graduating from Harvard College with a B. S.

degree in 1942, Tucker became a United States Naval

Aviator, and served as a di\e-bomber pilot stationed in

Guantanamo \a\al Station in Cuba until 1944, when he

was transferred to the United States Naval Medical Re-

search Unit 2 (NAMRU-2) based in Guam. As the unit's

first medical malacologist. Lieutenant Abbott traveled

throughout the Marianas, Philippines and the Yangtze

Valley of China, examining liuiidreds of species of fresh-

water mollusks to identify those capable of acting as

intermediate hosts of treniatodes parasitic in man, to

describe and study the life c\ cles of these parasites, and

to develop eftective counternieasures to control them.

In 1946, Tucker left the Navy, married Mary M. Sisler

and was appointed Assistant Curator in the Division of

Mollusks of the National Museum of Natural History,

Smithsonian Institution. In the \ears that followed, the

Abbott family grew with the births of Robert Tucker,

Jr., Carolyn Tucker, and Cvnthia Douglas. Tucker en-

rolled in the graduate program of George Washington

University, and earned a M. S. degree in 1949, com-

pleting his thesis "The Anatonn and Life History of the

fresh-water snail, Thiara granifera (Gastropoda: Proso-

branchiata)" under the direction of Ira B. Hansen. The

same year, he was promoted to Associate Curator. De-

spite maintaining an extensive publication record. Tuck-

er found time in tlie succeeding years to publish the first

edition of American Seashells (1954), one of his land-

mark works, and to complete his Ph. D. at George Wash-

Page 56

THE NAUTILUS, Vol. 110, No. 2

ington University. His dissertation, The Gastropod Ge-

nus Assiminea in the Philippines ' was completed with

Paul Simon Galtsoff serving as his faculty advisor.

In late 1954, Tucker left the National Museum of

Natural History to fill the new i\ established Pilsbr\' Chair

of Malacology at the prestigious Acadenn of Natural

Sciences of Philadelphia. Shortly after arriving in Phil-

adelphia, Tucker founded and served as the first presi-

dent of the Philadelphia Shell C^lub, as well as Editor of

its Proceedings. Following Henr\ Pilsbry's death in 1957,

he became co-editor of The Nautilus together with H.

B. Baker. In 1959, Tucker established the monographic

series Indo-Pacific Mollusca as a complement to John-

sonia, w hich dealt with western Atlantic molluscan fau-

nas, and wrote the first three monographs in rapid suc-

cession. Under Tucker's leadership, and with support

from the Natural Science Foundation, the Department

of Malacology enjoyed unprecedented grow th, launching

collecting expeditions to all areas of the Indo-Pacific

ranging from Madagascar to the Cocos-Keeling Islands,

the Bay of Bengal, tlie Philippines and the Cook Islands.

It was in the middle of his stay at the Academy of

Natural Sciences that I first met Tucker. He was 40,

already at the top oi his field, and just returned from the

du Pont — Academy of Natural Sciences expedition to

the Philippines. My classmate and I were ten years old

and very interested in man\ aspects of natural history,

.^fter spending an afternoon with Tucker, we both be-

Figure 2. Mr, R. Tucker Abbott in the moliusk collectiuiis of

the National Museum of Natural History, Smith,sonian Insti-

tution, 1950.

came avid malacologists and volunteers, a conversion he

effected regularly on enthusiasts of all ages.

During the Philadelphia years. Tucker continued to

publish prodigiously, yet found time to pursue his hobby,

the collection and classification of paper clips. When he

appeared as a guest on the television quiz shows "What's

My Line'r*' and "To Tell the Truth the panel deduced

that he w as a malacologist, but Tucker w as able to elude

them when appearing in his capacity as a "clipologist!"

Tucker was widowed in 1964 and married Sue Sweeney

Darwin in 1966.

Because of his prominence in the field and his large

follow ing among collectors. Tucker was recruited by John

du Pont to help build a new natural history museum in

the suburbs oi Wilmington, Delaware. He could not resist

the challenge and opportunit) to create a new Depart-

ment of Malacology or a new natural history museum

devoted in large part to moUusks. Tucker left the Acad-

emy of Natural Sciences of Philadelphia in 1969 to take

up the ilu Pont Chair ot Malacolog> and serve as Assistant

Director ot the Delaware Museum of Natural History.

Calling upon his professional colleagues and friends in

the world of amateur concholog\ . Tucker assembled a

collection and librar\ that toda\ is one of the ten largest

in the western hemisphere. \\ bile at the Delaware Mu-

M G. Harasewych, 1997

Page 57

Figure 3. Dr. R. Tucker Abbott marking proof pages of Sea-

shells oj the World in his office at the Academy of Natural

Sciences of Philadelphia, circa 1962.

seum of Natural History, Tucker continued as editor of

Indo-Pacific Mollusca, and assumed primary editorship

of The Nautilus upon the death of H. B. Baker in 1970.

He pubhshed numerous papers and books, among them

Kingdom of the Seashel! (1972) and tiie second edition

ot American Scashells (1974), and was appointed Ad-

junct Professor at the College of Marine Studies, Uni-

versity of Delaware in 1973. Despite a distance of some

100 miles between the Campus of the College of Marine

Studies and the Delaw are Nhiseum, Tucker was an active

faculty member, teaching graduate courses in Malacol-

ogy (together with M. R. Carriker) and Evolutionary

Biology, and serving on five graduate committees. His

students, myself among them, remember his enthusiasm

and skill in collecting moilusks in tlie field, as well as in

transforming the "surplus' specimens into a variety of

culinary delicacies, including his "'Seven Species Soup.

Tucker was divorced in 1974. He married Cecelia White

in the spring of 1977.

With his monumental accomplishment of museum

building behind him, Tucker "retired to Melbourne,

Florida in 1978, to concentrate on writing and to build

a successful publishing company, American Malacolo-

gists. Inc. Since Indo-Pacific Mollusca remained at the

Delaware Museum of Natural History, he started another

monographic series, Monographs of Marine Mollusca,

one not encumbered by zoogeographic boundaries. I re-

member Tucker confiding that, as an undergraduate, he

had been torn between pursuing a career in malacology

Figure 4. .A \ iew of Tucker in the specimen preparation room

of the Delaware \hiseum of .Natural History, circa 1972. (Photo

courtesy of Delaware Museum of .Natural History).

and one in writing. .At this point in Tucker's career, his

passion for writing clearly blossomed with a seemingly

endless profusion of books and articles for the amateur

naturalist and collector. His primary mission now more

than ever was to reach and educate an ever-expanding

audience of avid readers.

By the mid 1980 s. Tucker would occasionally talk of

retirement, when he would devote his time to gardening

and writing about gardening. But that was always "in

another five years. Even at this phase of his career, he

remained interested and active in various aspects of mu-

seum work, consulting with the Florida Museum of Nat-

ural History on several exhibits, and completely design-

ing the Hall of Shells at the newly built Fernbank Mu-

seum in .Atlanta.

When he was in his seventies, which he used to say

was middle-aged for a malacologist. Tucker accepted

another challenge, to build another museum, this time

one devoted entirely to moilusks and emphasizing edu-

cation and outreach rather than research. Those who

were aquainted with Tucker knew that he was never

happier than when he had some new project on the

drawing boards. He was grateful for having been given

the opportunity to call upon the experiences of his long

Page 58

THE NAUTILUS, Vol, 110, No. 2

Figure 5. R, Tucker Abbott ut the groundbreaking for the

Bailev-Matthews Shell Museum, JuK 6, 1992.

and varied career to design a museum that incorporated

tlie best elements of all tlie museums that he had worked

in or visited. First as Consultant to the Board, then as

Founding Director, Tucker devoted himself to this new

task with extraordinary energy, enthusiasm and inten-

sity. The Bailey-Matthews Shell Museum, which had its

formal opening No\ember 18th, barely two weeks after

his death, embodies Tucker s concept of the quintessence

of what a shell museum should be. He regarded it to be

the "crowning accomplishment of his life.

Many remember Tucker as "Mr, Seashell, ' a flam-

boyant man of boundless wit and good humor, described

in the Wall Street Journal as "partial to short sleeved

Hawaiian shirts decorated with brightK colored shells"

and wearing multiple pairs ol glasses tethered around

his neck. He had an enthusiasm that was contagious and

the uncanny abilit\ to transform even a casual acquain-

tance into a co-conspirator in the most far-fetched of

projects. Tucker was a man l)lessed with a love of his

work and an unfailing corn iction in its importance. For

him, the joy w as in conceiving and doing a project, rather

than its completion. Tucker also had a very strong sense

of duty and service to the malacological community. In

addition to .serving as editor of The Nautilus { 29 \ears),

Indo-Pacijic Mollusca (19 \ears) and Monographs of

Marine Mollusca (7 years), he was President of the Amer-

ican Malacological Union (1959), Founder and President

of The Philadelphia and Wilmington Shell C^lubs, Hon-

orarv Member of numerous shell clul)s in man\ countries.

Figure 6. R, Tucker .Abbott autographic a cop\ of one of his

books (May. 1992),

Secretary of the Society of Svstematic Zoology (1956-

59), Publications Chairman and later Awards Chairman

for the Conchologists of America, Councilman of the

Harvard Graduate Society for Advanced Study and Re-

search, Member of the Board of Trustees of the Bermuda

Biological Society and the Paleontological Research In-

stitution, and served numerous other organizations in

many and varied capacities.

Among his accomplishments. Tucker leaxes a legacy

of many significant contributions to the infrastructure of

malacology in .America. He enlarged collections, pio-

neered curatorial s\stems, built new museums and De-

partments of Mollusks, and is responsible for creating at

least two positions in the field of malacology. He main-

tained and created new venues for publication of schol-

arly works. He trained malacologists b\ ser\ing on their

graduate committees, and by organizing and adminis-

tering award programs that continue to support students

in malacology. Because of his interest in the historv of

.American malacology, he was instrumental in establish-

ing archives now administered by the .American Mala-

cological Union and housed at the .\cadem> of Natural

Sciences in Philadelphia. Perhaps his most obvious con-

tribution was his strong commitment to popularize the

science of malacolog\' and the hobb> oi shell collecting.

He w as instrumental in building a large, supportive, w ell-

educated, and well-organized constituency for mollusks,

a constituency that few other invertebrate groups enjoy.

In a "Letter to a Beginner." published in .American

Conchologist [15(2): 17], Tucker wrote "You ma\ wonder

why I've taken so much time to answer \ our letter, even

though I am 300 years behind in my w ork and writings.

It s because I wrote a letter similar to yours about fifty

\ears ago to a Harvard shell professor. Dr. William J.

M. G. Harasewych, 1997

Page 59

Clench. He patiently answered me in detail and started

me on the right track. It's m\ turn to pass on his kindness.

I hope \ou 11 have the chance to take your turn. " Tucker

Abbott has done a great deal for malacology during his

life. We would do \\ ell to remember his simple request.

R. Tucker Abbott is survived by his wife, Cecilia White

Abbott, his son, Robert Tucker Abbott. Jr., his daughters

CaroKn Tucker Palmer, C>ntliia Douglas Sullivan, Cher\l

Robin Nelson, Jonna Leigh liobson, and Erika von der

Heyden, and b) his brothers John, Matthew and Peter.

ZOOLOGICAL TAXA OF R. TUCKER ABBOTT

Despite a prolific publication record, R. Tucker Abbott

proposed relatively few new nomina, these being limited

to one family-level taxon, ten genus-level ta.xa (seven as

subgenera) and 71 species-le\el taxa (19 as subspecies).

Of these, 60'"f of the genus-level taxa and 21 ^c of the

species level-taxa were co-authored. Two taxa, the genus

Laevistrombiis and the species Acanthochitona halesae.

have been attributed to Abbott by subsequent authors

because he associated a description and or figure v\ith a

nude name of an earlier author.

Family-Level Taxa

PROPEAMUSSIIDAE Abbott, 1954, American Seashells,

p. 369. For Propcainussiuin deGregorio, 1884.

Genus-Level Taxa

Clenchiella Abbott, 1948. The Nautilus 61(3):76. Type

species: Clenchiella victoriac Abbott, 1948, by original

designation. Amnicolidae.

Echiniiui.'i Clench and .Abbott, 1942, Johnsonia 1(4):3.

New name for \ina Gra\, 1850, not Sina Horsfield,

1829, nor Nina Gray, 1855. Littorinidae.

Fukuia Abbott and Hunter, 1949. Proceedings of the

Helminthological Society of Washington 16(2):79-80.

Type species: Fukuia kurodai Abbott and Hunter, 1949,

by original designation. Amnicolidae.

Glohivasuiu .\bbott, 1950. As a subgenus of Vasum Rod-

ing, 1798. Johnsonia 2(28):215-216. Type species: Tur-

binelta nuttingi Henderson, 1919, by original designa-

tion. Turbinellidae.

Laevistrombus Abbott, 1960. As a subgenus of Strombus

Linne, 1758. Indo-Pacific Mollusca l(7):47-48. Type spe-

cies: Strombus canarium Linne, 1758, by original des-

ignation. Abbott attributed this taxon to Kira, 1955, whose

use was a nomen nudum. See Bieler & Petit, (1996) for

a discussion of the authorship of this taxon. Strombidae.

Namrutua Abbott, 1948. As a subgenus of Hua Chen,

1943. Bulletin of the Museum of Comparative Zoology

Harvard 100(3):296-297. T\pe species: Melania ning-

poensis Lea, 1856, by original designation. Thiaridae.

Pitarenus Rehder and Abbott. 1951. As a subgenus of

Pi tar Romer, 1857. Revista de la Sociedad Malacologica

"Carlos de La Torre " 8(2):57. Type species: Pitaria cor-

data Schwengel, 1951, by original designation. N'eneri-

dae.

Siphovasum Rehder and .Abbott 1951. As a subgenus of

Vasum Rriding, 1798. Re\ ista de la Sociedad Malacolo-

gica "Carlos de La Torre" 8(2):61. Type species: Vasum

(Siphovasum) latiriforme Rehder and Abbott, 1951, by

original designation. Turbinellidae.

Tcctininus Clench and Abbott, 1942. As subgenus of

Echininus Clench and Abbott, 1942. Johnsonia 1(4):4.

Type species: "non umbilicated Echininus nodulosus

(Pfeifter, 1839)", b\ original designation. Littorinidae.

Texadina .\bbott and Ladd, 1951. As a subgenus of Lit-

toridina E\doux and Soule\et, 1852. Journal of the

Washington Academy of Science 41(10):335. Type spe-

cies: T. sphinctostoma Abbott and Ladd, 1951, by orig-

inal designation. Amnicolidae.

Species-Level Taxa

Abbreviations for repositories of type material:

AM = Australian Museum, Sydney.

AMNH = .'\merican Museum of Natural History, New

York.

ANSP = The .-Academy of Natural Sciences of Philade-

phia.

BM(NH) = British Museum (Natural History), London,

[now The Natural History Museum, London]

BMH = Bishop Museum, Honolulu.

CM = Carnegie Museum, Pittsburg.

DMNH = Delaware Museum of Natural History, Wil-

mington.

FSM = Florida State Museum, Gaines\ille. [now Florida

Museum of Natural Histors ]

MCZ = Museum of Comparative Zoology, Harvard Uni-

versity, Cambridge.

MZUM = Museum of Zoolog\, Universit)' of Michigan,

Ann Arbor.

NMV = National Museum of X'ictoria, Brisbane.

PBS = Philippine Bureau of Science, Manila.

PML'H = Poey Museum, University of Havana.

RM = Raffles Museum, Singapore.

UMML = Rosenstiel School of Marine and Atmospheric

Science, L'niversitN of Miami.

I'SNM = National Museum of Natural History, Smith-

sonian Institution, Washington, DC.

IT = Institute of Marine Science, L'niversity of Texas.

WAM = Western .Australian Museum, Perth.

afrobellatus. Strombus terebcllaius Abbott, 1960. Indo-

Pacific Mollusca 1(2):88, plate 61, figure 2. Holotype

ANSP 214295, paratypes ANSP 211423, USNM, ANSP,

Coryndon Museum 1611. Pange Island, west side of Zan-

zibar, Tanzania. Strombidae.

alcaldei. Naniviirea Jaume and Abbott, 1948. Revista de

la Sociedad Malacologica "Carlos de la Torre" Habana

6(l):5-7, figures la-c, 4. Holotype PMUH 12178, para-

Page 60

THE NAUTILUS, Vol. 110, No. 2

types, Jaume collection 25202, USNM 488585. Cuba.

Amnicolidae.

alfredi, Turbonilla (Pyrgisciis) .\bbott, 1958. Mono-

graphs of the Academy of Natural Sciences of Philadel-

phia 11:104-105, text-figure 5, la-c. Holotype ANSP

198692, paratypes ANSP 198693, Ostheimer Collection.

Grand Cayman Island. Pyramidellidae.

andersoni. Phaliuin .Abbott, 1968. New name for Cassis

(Phalium)dalli Anderson, 1929, not Phalium dalli (Dick-

erson, 1917). Miocene, Caribbean coast of Colombia.

Cassidae.

areas, Terebra (Strioterebruni) Abbott, 1954. The Nau-

tilus 68(2):40-41, pi. 2, figure 4. Holotvpe USNM 613882,

paratvpes, USNM 613883, ANSP 191712, Steger collec-

tion (now DMNH 104984, DMNH 118028). Areas Cays,

Campeche Bank, Mexico. Terebridae.

atlanticus, Pleuwbranchus (Susania) Abbott, 1949. The

Nautilus 62(3):73-78, plate 5, figures 1-10. Holotype

USNM 574352, paratvpes USNM 574342, ANSP 184350,

MCZ 165951. Soldier Key, Bisca> ne Bay, Florida. Pleu-

robranchidae.

aiireofasciatus. Conns spuiius Rehder and Abbott. 1951.

Revista de la Sociedad Malacologica "Carlos de La Tor-

re" Habana 8(2):64-65, plate 9, figs. 3, 4. Holotype USNM

597521, USNM 597522, Sennott collection, Woolsey col-

lection. Off Dry Tortugas, Florida, in 20 fms. Conidae.

auricincta, Strombifortnis Abbott, 1958. Monographs of

the Academy of Natural Sciences of Philadelphia 11:

106-107, text-figure 5, 4. Holotype ANSP 198840, para-

types ANSP 199841, USNM 619550, ANSP 75855. Grand

Cayman Island. Melanellidae.

anstini. Conns Rehder and Abbott. 1951. Journal of the

Washington Academv of Sciences 41(l):22-24, figure 7.

Holotype USNM 603017, paratypes USNM 421721,

USNM 603018. Dry Tortugas, Florida. Conidae.

unstini, Stenoihijra Abbott, 1951. Journal ot the Wash-

ington Academy of Sciences 41(1):14-16, figures 1, 3-7

Holotype USNM 603669, paratypes USNM 603670,

USNM 603671, USNM 603672, MCZ. Leyte, Philippines.

Stenothyridae.

balcsae. Acanlliocliituna Abbott, 1954. American Sea-

shells, p. 318. Holotype ANSP 349331, Bonefish Key,

Florida. Abbott attributed this taxon to Pilsbry, 1940,

whose use was a nomen nudum. See Lyons (1988):102

for a discussion of the authorship of this taxon. Acan-

thochitonidae.

bayeri, Olivella Abbott, 1951. The Nautilus 64(4):111-

1 12, plate 7, figures 2a-b Holotype USNM 595132, para-

types USNM 412871, MCZ, ANSP, UMML. Off Ajax

Reef, off Elliott's Key, Lower Florida Keys. Olividae.

blandfurdi, ,\ssiniinea Abbott, 1958. I'roceedings of the

Academv of Natural Sciences of Philadelphia 110:247-

249, plate 21. Holotype USNM 615212. paratypes USNM

513820, ANSP 193360. Samar Island. Philippines. Assi-

mineidae.

boettgeri, Assiminea Abbott, 1958. Proceedings of the

Academy of Natural Sciences of Philadelphia 110:249-

251, plate 15, figure 3, plate 23. Holotype USNM 615214,

paratypes USNM 613828, ANSP 193370. Leyte Island,

Philippines. Assimineidae.

bonnci, Indopijrgns Abbott, 1945. Occasional Papers on

MoUusks l(l):3-4, plate 1, figures 1-6, plate 2, figures

1,2. Holotype, MCZ 147474, paratypes MCZ 147475,

MCZ 147476. Lake Pose, Celebes. Hydrobiidae.

boschi, Cijmatium (Ranularia) Abbott and Lewis, 1970.

The Nautilus 83(3):86-88, figures 1-6. Holotype DMNH

22862, paratypes AMNH 139206. ANSP 188844, Bosch

collection. Al Masirah Island, Oman. Ranellidae.

branluunac. Fasciolaria distans Rehder and Abbott. 1951.

Revista de la Sociedad Malacologica "Carlos de La Tor-

re" 8(2):59-60, plate 8. figs. 4,5. Holotype USNM 597513,

Branham collection, Raines collection. Off Puerto Alvaro

Obregon, Tabasco, Mexico. Fasciolariidae.

cardmca. Coralliophda Abbott, 1958. Monographs of the

Academy of Natural Sciences of Philadelphia 11:166-

67, text-figure 3, plate 1, figures g-h. Holot>pe ANSP

197054, paratvpes ANSP 61054, (DMNH 24282 ex ANSP

61054), ANSP 194091, ANSP 29958, ANSP 36673, ANSP

196397, ANSP 29962, ANSP 29959, ANSP 190548, ANSP

197056, ANSP 197055, ANSP 195857, MCZ 133415, MCZ

38548. Vera Cruz, Mexico. Coralliophilidae.

caribaea, Cosa .\bbott, 1958. Monographs of the Acad-

emy of Natural Sciences of Philadelphia 11:112-113,

text-figure 6. Holotype ANSP 200064, paratype ANSP

205419. Grand Cayman Island. Philobryidae.

clarki. Stcnothyia ansiini Abbott, 1951. Journal of the

Washington Academ> of Sciences 41(1):14-16, figure 2.

Holot) pe USNM 603673, parat> pes USNM 303387. Min-

doro Island, Philippines. Stenoth\ridae.

clarki. Conns Rehder and Abbott. 1951. Journal of the

Washington Academy of Sciences 41(1):22, figures 1-6.

Holotype USNM 485740, parat\ pes USNM 488 465, MCZ.

SSW of Marsh Island, Louisiana. Conidae.

cookei, Omphalotropis Abbott. 1949. Occasional Papers

of the Bishop Museum 19(I5):263-266. figure la-e. Ho-

lotype USNM 591306, paratypes MCZ 182894, BMH

212195, numerous additional paratypes at USNM, MCZ,

BMH. Saipan Island, Marianas. Synceridae.

cosmani, Chicoreus .\bbott and Finlay, 1979. The Nau-

tilus 94(4):159-162, figures 1-9. Holotype USNM 783323,

paratypes AMNH, ANSP, DMNH. Cosman, Finla\. and

Bijur C:ollections. Ocho Rios. Jamaica. Muricidae,

dinagatcnsis. Synccra pscndoquadrasi .\bbott. 1948.

Bulletin of the Museum of Ciomparative ZoologN Har-

vard. 100i3):284-285. Holotspe MCZ 175323, paratypes

MCZ 175324, USNM 485565. Panamoan, Dinagat Island,

NE of Mindanao Island. Philippines. Synceridae.

M. G. Harasewych, 1997

Page 61

cloiseae. Acteon Abbott. 197;3. The Nautilus 87(4):91-

92, figures 1-5. Holotype DMXH 27213, paratypes

DMXH 67746, Bosch collection. Al Masirah Island, Oman.

Acteonidae.

emipowlusi. Ocenebra (Ocenebra) Abbott, 1954. The

Nautilus 68(2):41-42, pi. 2, figure 3. Holotype USNM

613881, paratypes, ANSP 191711, Steger collection (now

DMNH). Off Fort Myers, Florida. Muricidae.

gcrrardi. Transeiiuella .Abbott, 1958. Monographs of the

Academy of Natural Sciences of Philadelphia 11:130-

131, text-figure 7, plate 4, figures a-c. Holotvpe ANSP

199507, paratvpes ANSP 199508, ANSP 199509, (DMNH

21194 ex ANSP 199509), ANSP 199506, USNM 619551,

MCZ, USNM, ANSP 199510. Grand Ca\ man Island. Ve-

neridae.

grandanus, Cantharus midtangulus Abbott, 1986. The

Nautilus 100(4);120-121, figure 1. Holotype USNM

859096, paratypes ANSP, AMNH, MCZ, FSM. Black Isle,

St. Joseph Ba\, Gulf Count\, Florida. Buccinidae.

guamcnsis, Sijncera nitida Abbott, 1949. Occasional Pa-

pers of the Bishop Museum 19(15):273-274, figures 8a-

c, 9a-c. Holotype USNM 589997, paratypes USNM

589992, MCZ 183904, BMH 212193. Banks of the Pago

River, Guam Island, Marianas. Synceridae.

habci. Assiminea habei Abbott, 1958. Proceedings of the

Academy of Natural Sciences of Philadelphia 110:251-

254, plates 16-18, 25. Holotype USNM 615213, para-

types USNM 613801, ANSP 193367, USNM 613831,

ANSP 193366, (DNMH 25311 ex ANSP 193366), ANSP

193368, ANSP 193369. Leyte Island. Philippines. Assi-

mineidae.

hectori, Phalium .\bbott, 1968. New name for Cassis

muricata Hector, 1886, not Cassis muricata Menke, 1828.

Miocene, New Zealand. Cassidae.

hedgpethi, Pleurobranchaca Abbott, 1952. Florida State

University Studies 7:1-2, plate 1, figures 1-8. Holotype

IISNM 574607, paratype, Hedgpeth collection. Port

Aransas, Texas. Pleurobranchidae.

hoffmeyeri, Terebra (Strioicrcbntin) Abbott, 1952. The

Nautilus 65(3):78-80, plate 5, figures 5-9. Holotype

USNM 598077, paratvpes USNM 596972, MCZ, ANSP,

(DMNH 24990 ex ANSP 234286), BMH, PBS, Hadley

Collection. Basay Beach, Manila Ba\ , Luzon Island, Phil-

ippines. Terebridae

bummi, Pohjccra .\bbott, 1952. Florida State University

Studies 7:3-4, plate 2, figures 1-8. Holotype, USNM

574647, paratypes USNM 574648, Humm collection. Al-

ligator Harbor, Franklin Count\ , Florida. PoKceridae.

indomaris, Lambis scorpius Abbott, 1961. Indo-Pacific

Mollusca 1(3):165-166, plate, figure. Holotype ANSP

242210, paratypes ANSP 247564, ANSP 212384, ANSP

214263, ANSP 212968. Nossi-be, Madagascar. Strombi-

dae.

iredalci. Strombus vomer Abbott, 1960. New name for

Strombiis australis Gra\, 1826, not Strombus austraUs

Schroter, 1805. Strombidae.

keelingensis, Siiccinea Abbott, 1950. Bulletin of the Raf-

fles Museum 22:87-90, figures 1-7. Holotype USNM

488579, paratypes, USNM 488580, RM, BM(NH). North

Keeling Island, Cocos-Keeling Islands, Succineidae.

klineorum, Strombus Abbott, 1960. Indo-Pacific Mol-

lusca 1(2):70, plate 20, figures 20-21. Holotype ANSP

247621, paratypes ANSP 211121, ANSP 211173, MCZ

224889. Powder Bay, Sri Lanka. Strombidae.

kurodai, Fuhuia .Abbott and Hunter, 1949. Proceedings

of the Helminthological Societ\ of Washington 16(2):80-

82, plate 1, fig. 6, plate 2, fig. 4.' Holotype USNM 593365,

parat\pes USNM 488538, USNM 488517. Takeda River,

near Maruoka City, Fukui Prefecture, Honshu, Japan.

Amnicolidae.

kurodai. PhaUum .Abbott, 1968. Indo-Pacific Mollusca

2(9):105-106, plate 87. Holot\pe NSMT, paratypes ANSP

241694. Tosa Bay, Shikoku, Japan. Cassidae.

latiriforme, Vasum (Siphovasum) Rehder and Abbott.

1951. Revista de la Sociedad Malacologica "Carlos de La

Torre" 8(2):61-62, plate 9, figs. 5,6. Holotype USNM

597517, paratypes, USNM 597518,

Sennott collection, Raines collection. NW of Cape Ca-

toche, Yucatan, Mexico, in 18-20 fms. Turbinellidae.

luzonica, Assiminea habei Abbott, 1958. Proceedings of

the Academy of Natural Sciences of Philadelphia 110;

254, plates 15, figure 9. Holotype ANSP 96035, paratypes

ANSP 193356, ANSP 193355. USNM 201151. Luzon Is-

land, Philippines. Assimineidae.

margaritensis. Murex Abbott, 1958. Monographs of the

Academy of Natural Sciences of Philadelphia 11:61, plate

1, figures n-o. New name for Murex imperialis Swainson,

1831, not Murex imperiahs G. Fischer, 1807. Muricidae.

marshallensis. Assiminea nitida Abbott, 1958. Proceed-

ings of the Academy of Natural Sciences of Philadelphia

110:256, plate 15, 'figure 8. Holotype USNM 615210,

paratypes USNM 580909, ANSP 193364. Japtan Island,

Enivvetok Atoll, Marshall Islands. Assimineidae.

minirosea, Ocenebra (Ocenebra) Abbott, 1954. The Nau-

tilus 68(2):43-44, pi. 2, figure 1. Holotvpe USNM 103380,

paratvpes, USNM 161271, USNM 103413, USNM 416653,

USNM 416655, USNM 416654. Jamaica. Muricidae.

nworei. Olivella Abbott, 1951. The Nautilus 64(4):112-

114, plate 7, figure la-b. Holotvpe USNM 595133, para-

types USNM 595134, MCZ, ANSP, UMML. Carysfort

Reef, off Key Largo, Florida. Olividae.

morrisoni, Assiminea Abbott, 1958. Proceedings of the

Academy of Natural Sciences of Philadelphia 110:259-

260. Holotype USNM 615211, paratypes USNM 614191,

ANSP 193357. Teteu Island, Raroia Atoll. Tuamotu Is-

lands. Assimineidae.

Page 62

THE NAUTILUS, Vol. 110, No. 2

multistriata, Fukuia Abbott aiul Hunter, 1949. Proceed-

ings of the Helminthological Societ\ ot VV'ashiiigtoii 16(2):

82-83, Plate 1, fig. 5, plate 2, figs. 5 & 11. Holotype

USNM 594185, paratypes USNM 488540, USNM 487383.

Takefu in Kamiyama-mura, Nannjo-gun. Fiikui Prefec-

ture, Honshu, Japan. Amnicolidae.

nipponensis, Casmaria ponderosa .\bbott, 1968. Indo-

Pacific Mollusca 2(9);200, plate 14, figures 13-14. Ho-

lotype ANSP 313258, paratypes ANSP 241636, ANSP

227579, MCZ 233233. Tanabe, Wakayama Prefecture,

Japan. Cassidae.

ochroglottis, Stronibus inutabilis Abbott, 1960. Indo-Pa-

cific Mollusca l(2):74-75, plate 20. figures 9-10. Holo-

type ANSP 250187, paratypes ANSP 185466, USNM

2'6652a. USNM 465685, MCZ. Mauritius. Strombidae.

orrae. Strombus urceits Abbott, 1960. Indo-Pacific Mol-

lusca l(2):66-67, plate 20, figure 28; plate 41, figure 5.

Holotvpe ANSP 247756, para^types MCZ, ANSP 232673,

ANSP 233409, ANSP 240131, ANSP 240137. Augustus

Island, Western Australia. Strombidae.

ostheimerae. Emarginula .Abbott, 1958. Monographs of

the Academy of Natural Sciences of Philadelphia 11:18,

plate 1, figure c. Holotype ANSP 195983, paratypes ANSP

195982, Ostheimer collection. Grand Cayman Island.

Fissurellidae.

parkeri, Ithycythara Abbott, 1958. Monographs of the

.\cademv of Natural Sciences of Philadelphia 11:96, text-

figure 5. Holotype ANSP 198919, paratypes ANSP 209867

Grand Cayman Island. Turridae.

patac. Conns Abbott, 1971. The Nautilus 85(2):49-51,

figures 1-6. Holotvpe DMNH 44097, paratypes DMNH

44096, DMNH 40595, DMNH 44095, Finlay collection.

Pompano Beach, Broward Count\ , Florida. Conidae.

pihbryi, Lambis crocata Abbott, 1961, Indo-Pacific Mol-

lusca 1(3);158-159, plate 128, figures 1-2. Holotype ANSP

156123, paratypes MCZ 49694, BMH 68268. Nuku Hiva,

Marquesas. Strombidae,

pseudoquadrasi, Syncera .Abbott, 1948. Bulletin ot the

Museum of Comparative Zoology Harvard. 100(3):283-

284, plate 1, figs. 11-13, text iig. 12. Holotype MCZ

175325, paratypes MCZ 175326, USNM 485564. Bugo,

Macajalar Bay, northern Mindanao Island, Philippines.

Synceridae.

rasilistoma, Tudicula (Tiuliculu) .Abbott, 1959. Indo-Pa-

cific Mollusca 1(1):29, plate 4, figure 2, plate 10, figures

A-C. Holotype NMV F18189, paratypes NMV, ANSP

227669. Tweed Heads, New South Wales, Australia. Tur-

binellidae.

saipanensis, Paludinella conica Abbott, 1949. Occasional

Papers of the Bishop Museiun 19(15):271. Holotype

ISNM 590891, paratypes MC;Z 182889, USNM, MCZ,

BMH. North end of Saipan Island, Marianas. Synceridae.

satlasi,Murex(Murexjrecurvirostris Hehderand Abbott.

1951. Revista de la Sociedad Malacologica "Carlos de La

Torre" 8(2):58-59, plate 9, figs. 7, 8. Holotype USNM

597515, USNM 597516, Sennott collection. 60 miles NW

of Cape Catoche, Quintana Roo, Mexico, in 18-20 fms.

Muricidae.

sennottoriim. Conns Rehder and .\bbott, 1951. Revista

de la Sociedad Malacologica Carlos de la Torre 8(2):63,

pi. 9, figures 1-2. Holotype USNM 597519, paratypes,

USNM 597520, Sennott collection. SW of Campeche,

Mexico. Conidae.

sphinctostoma. Littoridina (Texadina) Abbott and Ladd,

1951, Journal of the Washington Academy of Science

41(10):335-338. Holotype USNM 596722, paratypes,

USNM 596723, USNM 596724, MCZ, ANSP 187519,

MZUM, UT. N of Webb Point, NW side of San Antonio

Bay, 27 miles NE of Rockport, Texas. Amnicolidae.

springeri, Anadara Rehder and .\bbott, 1951. Revista de

la Sociedad Malacologica "Carlos de La Torre" 8(2):54.

plate 8, figs. 8, 9. Holotype USNM 597375, paratypes

USNM 538689, USNM 603848, USNM 603843. 12 miles

off Dry Tortugas, Florida, in 66-60 fms. Arcidae.

stegeri, Terebra (Myurella) floridana Abbott, 1954. The

Nautilus 68(2):39-40, pi. 2, figures 5,6. Holotype USNM

613884, paratypes, USNM 613885, USNM 605184, ANSP

191709, Steger collection (now DMNH 105856). Off

Campeche, Mexico. Terebridae.

thielei, AssUninea Abbott, 1958. Proceedings of the

Academy of Natural Sciences of Philadelphia 110:242-

244, plate 15, figure 5, plates 9, 22. Holotype USNM

609577, paratvpes USNM 613802, USNM 613813, USNM

613848, USNM 1018, ANSP 193358, ANSP 193359, MCZ.

Leyte Island, Philippines. .Assimineidae.

transversnnt, Microcardium Rehder and Abbott, 1951.

Revista de la Sociedad Malacologica "Carlos de La Tor-

re" 8(2):56, plate 9 figs. 9, 10. Holotype USNM 485734.

50 miles SSW of Marsh Island, Louisiana, in 29 fms

(Pelican station 94-1). Cardiidae.

vector. Phalinm (Seniicassis) Abbott, 1994, The Nautilus

107(3):94-96, figures 1,2. Holotype. USNM 860326,

paratypes, Dan collection. Sa\ a de Malha Bank. Cassidae.

victoriac. Clcnchiclla Abbott, 1948. The Nautilus 61(3);

76-80, plate 5, figures 1-7. Holotype USNM 488534,

paratypes USNM 488535, MCZ. ANSP 18.3548. MZIM,

CM. San Joaquin River, Le> te Island, Philippines. Am-

nicolidae.

virginensis. Latirns (Polygona) Abbott, 1958. Mono-

graphs of the .\cadem\' of Natural Sciences of Philadel-

phia 1 1 :76, text-figure 4, plate 2, figure r. Holot\ pe ANSP

196459, paratypes ANSP 34975, ANSP 34968, ANSP

34969. St. Thomas Island, Virgin Islands, Fasciolariidae.

iLilsoni, SIrondins (Canarium) .\bbott, 1967. Indo-Pa-

cific Mollusca l(7):455-456, plate 328, figures 1-3. Ho-

lotvpe ANSP 253088. paratvpes ANSP 212695. WAM

847-66. WAM 840-66, WAM 838-66. WAM 851-66,

ANSP, BMH. Zanzibar, Tanzania. Strombidae.

M. G. Harasewych, 1997

Page 63

liinkuorthi. Fhalinrn Abbott, 196>S. Intlo-PaciBc Mol-

lusca 2(9):179-181, plates 165-16b. Holot\pe AM, para-

types AiNSP 272396, WAM 159-62. Abrolhos Islands,

Western Australia. Cassidae.

zanzibarica. Titdicula Abbott, 1958. \otulae Naturae

305:1-4, figures 1-7. Holotype AXSP 225261, paratypes

ANSP 225262, 2 additional paratypes. Ras Nungwi, Zan-

zibar, Tanzania. Turbinellidae.

PUBLICATIONS OF R TUCKER ABBOTT

R. Tucker Abbott was a prolific author, including among

his works numerous newspaper articles, book reviews and

sound recordings. Man\' of his popular works were pub-

lished in multiple printings and editions, some were

translated and published in several languages. The var-

ious editions of his works are listed here, but individual

printings are not. Man\' of Tucker's book reviews were

published in se\eral different journals or shell club pub-

lications. The following list of publications is arranged

in chronological order. Dates of publication are provided

when they could be determined. Dates of items published

in publications of the United States National Museum

are from Anonsmous (1971), those published in The

Nautilus are from Coan and Harasewych (1993).

Clench, \V. J. and R. T. Abbott. 19-11. The genus Strombus

in the Western Atlantic. Johnsonia 1(1):1-15, [October 25]

Clench, W. J. and R. T. Abbott. 1942. The genera Tectarius

and Echininiis in the Western Atlantic, Johnsonia 1(4):1-

4. [May 1]

Clench, W. J. and R, T. .Abbott 1943 Tiie genera Cypraea-

cassis. Morum, Sconsia and Dalium in the Western \\.-

lantic, Johnsonia 1(9):1-S. [July 7]

Clench, W. J. and R. T. Abbott. 1943 The Cenera Gaza and

Livona in the Western Atlantic. Johnsonia l(12l;l-9. [De-

cember 31]

Abbott, R. T. 1943. Guantanamo Bay. Cuba Johnsonia 1(12);

10-11. [December 31]

.Abbott. R. Tucker, 1944. The genus Modulus in the Western

Atlantic. Johnsonia l(14):l-6. [October 16]

Abbott, R, T, 1945, A new Celebes Fresliwater Snail (Hy-

drobiidae). Occasional Papers on Mollusks, Harvard, 1(1):

1-4, [February 27]

Abbott, R, T. 1945, The Philippuie Intermediate Snail Host

(Schistomophora quadrasi) of Schistosomiasis. Occasional

Papers on Mollusks, Harvard 1(2):5-16. [April 30]

Clench, William J, and R. Tucker Abbott. 1945. The Genus

Strombus in the Western .Atlantic, Johnsonia 1(18):1, [No-

vember 23]

.Abbott, Lieutenant Tucker, 1946 The Egg and Breeding

Habits of Oncomelania quadrasi Mlldff,. the Schistoso-

miasis Snail of the Philippines Occasional Papers on Mol-

lusks, Harvard 1(61:41-48, [February 14]

Abbott, Tucker, 1947, Review ol "Morris, P A, 1947, A

Field Guide to the Shells of our Atlantic and Gulf Coasts.

Houghton Mifflin Co., Boston. 190 pp The Wood Thrush

3:21, [December]

Abbott, R, Tucker, 1948. A New Genus and Species of Phil-

ippine Amnicolidae. The Nautilus 61(3):75-80, plate 5,

[March 2]

Abbott, Lieutenant R, Tucker, 1948, Handbook of Medicallv

Important Mollusks of the Orient and Western Pacific.

Bulletin of the Museum of Comparati\e Zoology Harvard.

100(3):243-328, pis. 1-5. [April]

Abbott, R, Tucker, 1948, Notes on the .Anatoni) of a giant

Subspecies Mclongena corona from Florida Shell Notes

by Frank Lyman 2(2&3):21-27, plates 1-5, [May]

Jaume, M, L, and R, Tucker .Abbot [sic], 1948, A new Cuban

species of the .Amnicolid genus i\^anivitrea. Revista de la

Sociedad Malacologica "Carlos de la Torre" Habana 6(1):

5-8. [June]

Abbott, R. Tucker. 1948. A potential snail host of oriental

Schistosomiasis in North .America (Pomatiopsis lapidaria).

Proceedings of the United States National Museum Smith-

sonian Institution 98(:3222):57-68, pis. 3,4. [July 2]

Abbott, R. Tucker. 1948. The Spread and Destructiveness of

the Giant .African Land Snail Achatina julica. The Nau-

tilus 62(1 ):31-34. [July 22]

Abbott, R. Tucker. 1949. March of the Giant African Land

Snail. Natural History 58(2):68-71. [February]

.Abbott, R. Tucker. 1949. A .New Florida Species of the Tec-

tibranch Genus Pleurobranchus. The Nautilus 62(3):73-

78, plate 5, figures 1-10. [March 18]

Abbott, R. Tucker. 1949. Mollusks and Medicine in World

War II. Report, Smithsonian Institution for 1947:325-;338,

pis. 1-3. [presented as a paper at the 14th .American Mal-

acological Union meeting, Pittsburgh in 1948. Presentation

reviewed in American Malacological L'nion News Bulletin

and Annual Report 1948:16-17 [March] [reprinted in Ster-

kiana 9:28 (1963)]

Abbott, R, Tucker. 1949. New S\ncerid Mollusks from the

Marianas Islands (Gastropoda, Prosobranchiata, Synceri-

dae). Occasional Papers ot the Bishop Museum 19(15):261-

274. [June 25]

Abbott, R. Tueker and G. W, Hunter 111 1949, Studies on

Potential Snail Hosts of Schistosonm japonicum I. Notes

on the .Amnicolid snails of the genera Blanfordia. Tricula,

and a new genus Fukuia from Japan. Proceedings of the

Helminthological Societv of Washington 16(2):7;3-86, plates

1.2 [July]

Hunter, III, G. W., R. Tucker Abbott, C Pan and E. Stray.

1949. Studies on Potential Snail Hosts of Schistosoma

japonicum II. Infection Experiments on .Amnicolid snails

of the genera Blanfordia. Tricula. & Fukuia. Proceedings

of the Helminthological Societv of Washington 16(2):86-

89. [July]

.Abbott, R. Tucker. 1949 Sexual dimorphism in Indo-Pacific

Strombus. The Nautilus 63(2):58-61, [November 1]

.Abbott, R. Tucker, 1949. An Indian Species of Clenchiella.

The Nautilus 63(2):62. [November 1]

Abbott, R. Tucker. 1949. Samoa, Shell-collector's paradise.

The Scientific MonthU 69(5)319-327, [November 3]

.Abbott, Robert Tucker. 1949 The .Anatonn and Life History

of the fresh-water snail, "Thiara granifera" (Gastropoda:

Prosobranchiata). Master of Science Thesis. George Wash-

ington University, Washington, DC \i + 68, ill. Directed

b>': Ira B. Hansen. [November]

.Abbott, R. Tucker. 1950. The Genus Cyclostrcma in the West-

ern Atlantic. Johnsonia 2(27):193-200. [January 6]

Abbott, R. Tucker 1950 The Genera Xancus and Vasum in

the Western Atlantic Johnsonia 2(28):201-219. [January

6].

Abbott, R, T. 1950, Voyages of the "Eolis", Johnsonia 2(28):

219-220, [January 6],

Abbott, R, Tucker, 1950, Snail Invaders, Natural History .59(2):

80-85, [February]

Page 64

THE NAUTILUS, Vol. 110, No. 2

Abbott, R. Tucker. 1950 Review of •C M Voiige. 1949.

The Sea Shore. \\\ + 311 pp. Collins, LoikIoii" The Sci-

entific Monthly. 70(2):135-136. [February]

.Xbbott, R. Tucker. 1950. The Molluscan Fauna of the Cocos-

Keeling Islands, Indian Ocean. Bulletin of the Raffles Mu-

seum, Singapore 22:68-98. [.-Vpril].

Abbott, R. Tucker. 1950. The \'enomous Cone Shells. The

Science Counselor 13(4)T25-126, 153. [December]

Abbott, R. Tucker. 1951. New stenothyrid gastropods from

the Philippines (Rissoidae). Journal of the Washington

.i^cademy of Sciences 41(1):14-16. [January 15]

Rehder, Harald A. and R. Tucker Abbott. 1951. Two new-

Recent cone shells from the Western Atlantic (Conidae).

Journal of the \\ ashington Academy of Sciences 41(1 ):22-

24. [January 15]

.'\bbott, R. Tucker. 1951. The use of infra-subspecific names

in Doctor de la Torre's essay on Polymita. The Nautilus

64(3):103-104. [February 15]

.■\bbott, R. Tucker. 1951. Review of "de la Torre y Huerta,

C. 1950. El Cencro Polymita. Memorias Soc. Cubana

Hist. Nat. 20(11:1-20, pis. "l-ll." The Nautilus 64(3):107.

[February 15]

Abbott, R. Tucker. 1951. Review of "Moreno, A. 1950. Es-

tudio Anatomico del genero Polymita Beck. Memorias

Soc. Cubana Hist. Nat. 20(l):21-35, pis. 12-22." The Nau-

tilus 64(31:108. [February 15]

Abbott, R. Tucker. 1951. Review of "Colnian, J. S. 1950.

The Sea and Its Mysteries. Norton, New York, xvi + 261

pp." The Scientific Monthly 72(2): 132. [February]

Abbott, R. Tucker. 1951. In Search of the Golden Cowrie.

Natural History 60(3):104-110, 144. [March]

Abbott, R. Tucker. 1951. New Deep-Water Olivellas from

Florida, with Notes on the O. jaspidea-nivea Complex.

The Nautilus 64(4):110-1 16. plate 7, figures 1-5. [May 7]

Abbott, R. Tucker. 1951. Operation Snail-Folk. Natural His-

tory 60(6):280-285. [June]

Abbott, R. Tucker. 1951. Review of "Pearse, A. S. 1950.

The Emigration of Animals from the Sea. Sherwood Press,

Drvden, NY xii + 210 pp." The Scientific Monthly 73(1):

68.' [July]

Abbott, R. Tucker. 1951. Eastern Pacific Poromya and Ce-

toconcha Names. The Nautilus 65(1 ):33. [August 27]

Abbott, R. Tucker. 1951. Review of "Carson, R. L. 1950.

The Sea Around Us. Oxford University Press, NY, vii -I-

230 pp." The Scientific Monthly 73(2):'l37-138. [August]

Abbott, R. T. and H. S. Ladd. 1951. A New Brackish-water

gastropod from Texas (Anmicolidae: Littoridina). Journal

of the Washington Academs of Science 41(101:335-338.

[October]

Rehder, Harald A. and R. Tucker Abbott. 1951. Some New

and Interesting Molhisks Irom the Deeper Waters of the

Gulf of Mexico. Revista de la Sociedad Malacologica "Car-

los de La Torre" 8(2):53-66, 2 plates. [November 12]

Abbott, R. Tucker. 1951. Battle of the Snails. Science Digest,

for November 1951: 15-19. [November]

Abbott, R. Tucker. 1951. Genetic Analysis of Wild Popula-

tions of the Genus Cerion. Colonial Inheritance of the

multiple factor for shell length. [Abstract and Discussion

of paper presented at 17tli Annual Meeting of the Amer-

ican Malacological Union, August 22-24, 1951 ] American

Malacological Union News Bulletin and .-Viinual Report

1951:10-11. [December) [Reprinted in Sterkiana 9:38

(1963)]

Abbott, R Tucker. 1952. Re\ie« of "Klingel, G. 1951. The

Bay. Dodd, Mead, New York \ -I- 287 pp " The Scientific

Monthly 74(11:58. [January]

Abbott, R. Tucker 1952. \ new Terebra (hoffmeyeri) from

the Philippines The Nautilus 65(31:77-80, plate 5, figures

5-9. [Februarx 25]

.■\bbott. R Tucker 1952 .\ Stud\ of an Intermediate Snail

Host (Thiara granifera) of the Oriental Lung Fluke (Par-

agonimus). Proceedings of the United States National Mu-

seum 102(32921:71-116, pis. 8,9. [February 26]

Rehder, Harald .A , R Tucker .Abbott, Joseph P. S. Morrison,

David Nicol, Julia Gardner, W. P. Woodring, John B.

Reeside, Jr., Ralph W. Imla\ , L. W. Stephenson, and W.

A. Cobban. 1952 Objection to M. Gilbert Ranson's .Ap-

plication in Regard to the Generic Name "Cryphaea"

Lamarck, 1801 (Class Pelecypodal. Z. N. (S.)365. Bulletin

of Zoological Nomenclature 6(61:188-189. [Ma> 22]

Abbott, R. Tucker. 1952. Review of "Berrill, N. J. 1951.

The Living Tide. Dodd, Mead, New York 256 pp " The

Scientific Monthly 74(51:313. [May]

.Abbott, R. Tucker 1952. Two New Opisthobranch MoUusks

from the Gulf of Mexico belonging to the Genera Pleu-

robranchaea and Polycera. Florida State I niversitv Stud-

ies 7:1-7. [October 15]

Abbott, R. Tucker. 1952. Review of "Morris, P. A. 1952. A

Field Guide to Shells of the Pacific Coast and Hawaii.

Houghton Mifflin, Boston." Atlantic Naturalist 8(2):I00.

[November]

Abbott, R, Tucker. 1952. Review of "Morris, P. A. 1952. A

Field Guide to Shells of the Pacific Coast and Hawaii.

Houghton Mifflin, Boston." The Scientific MontliK 75(6):

384. [December]

Abbott, R. Tucker. 1952. Neu Federal Regulations on Im-

porting Mollusks. [Abstract and Discussion of paper pre-

sented at 18th .Annual Meeting of the .American Malaco-

logical Union, August 20-23, 1952 ] American Malacolog-

ical Union Annual Report 1952:9. [December]

Abbott, R. Tucker. 1952. Opportunities in Medical Malacol-

ogy. [.Abstract and Discussion of paper presented at 18th

Annual Meeting of the .American Malacological L nion,

August 20-23, 1952 ] .American Malacological I'nion .An-

nual Report 1952:14-15. [December]

Abbott, R. Tucker. 1953. Federal Regulations on Importing

Living Mollusks. The Nautilus 66(31:104. [Februar\ 2 ]

Abbott, R. Tucker. 1953. Review of "Hedgpeth, J. 1952.

Between Pacific Tides. Ricketts and Calvin. 3rd Edition"

Scientific Monthly 76(11:49. [February]

Warinke, G. L. and R. Tucker .Abbott 1953. The Gross Anat-

omv and Occurrence in Puerto Rico ol the Pelecypod

Yoldia perprotructa. Journal of the \\ ashinglon .Academy

of Science 43(8):260-261. [August]

Abbott, R. Tucker. 1954. How to Collect Shells. Natural Hi.s-

tory 63(I):32-37. [January]

Abbott. R. Tucker. 1954. Section 10. MedicalK Important

Mollusks. /)!. Hunter, G. W., W. W . Fr>e and J C Swa-

trzw elder (eds.) .A Manual of Tropical Medicine. W B.

Saunders C;o, Philadelphia, pp. 625-640. [Jaiiuar\ ] [Second

Edition 1954, Third Edition 1960]

Abbott, R. Tucker. 1954. The Habits and Occurrence of the

Nudibranch, Armina tigrina. in southeast I nited States.

The Nautilus 67(3):83-86. [Februar> 18]

Abbott, R. Tucker. 1954. Review of the Atlantic Periwinkles,

Sodilittorina. Echininus, and Tectarius. Proceedings of

the Ignited States National Museum l()3(3328):449-464

[March 23]

.Abbott, R Tucker [Illustrations bv Frederick M Ba\er] 1954

M. G. Harasewych, 1997

Page 65

American Seashells. D \'aii Ndstraiul t^o.. Int. New York

xiv + 541 pp., 40 plates. [April 1.5]

Abbott, R. Tucker. 1954. New Gulf of Mexico Gastropods

(Tercbra anil Ocenebra). The Nautilus 68(2);37-44, pi. 2.

[November 15]

Abbott, R. Tucker [illustrations b\ Frederick M. Ba\er]. 1955.

Introducing Seashells. D. \'an Nostrand Compan\, Inc.

New York, vi + 64, 8 plates. [Februar\ 17]

Abbott, R. Tucker. 1955. Secrets of the Pearl. Natural Histor\

64(3):130-133. [March 7]

Abbott. R, Tucker. 1955. The Titian R. Peale Shell Collection.

The Nautilus 68(4): 123-127, plate 4, figures 1-8. [April

28]

.\bbott, Robert Tucker. 1955. The Gastropod Genus As.si-

minea in the Philippines Doctor of Philosophy Disserta-

tion. George Washington L niversit\, Washington DC. i\

+ 136 pp. Directed by Paul Simon Galtsoft. [June]

.•\bbott, R. Tucker, George M. Moore, Jeanne S, Schwengel,

and Margaret C, Tesks . 1955. Forward, p. i. In: Abbott,

R. Tucker, M. K. Jacobson, and M. C. Teskey (Editors).

How to Collect Shells. (A Symposium). American Mala-

cological Union, Buffalo, NY [i\ ] + 75 + [vii] pp. [ p. i in

second edition, 1961; p. i in third edition, 1966, p. ii,]

.\bbott, R. Tucker 1955 Introduction p ii In: Abbott, R

Tucker, M. K. Jacobson, and M. C. Teske\ (Editors). How

to Collect Shells. (A S\ mposium). .\merican Malacological

Union, Buffalo, NY [iv] + 75 + [vii] pp. [ p. ii in second

edition, 1961; p. ii in third edition, 1966, p. ii, modified

in fourth edition, 1974]

Abbott, R. Tucker. 1955. The Shell collection, pp. 40-43.

(reprinted in part from "American Seashells ), In: .Abbott,

R. Tucker, M. K. Jacobson, and M. C. Teskey (Editors)

How to Collect Shells. [\ Sv mposium). .\merican Mala-

cological Union, Buffalo, NY [iv] + ~b + [vii] pp. [pp. 79-

83 in second edition, 1961; pp. 80-84 in third edition, 1966

and fourth edition, 1974]

Abbott, R. Tucker. 1955. Oiitstandnig Shell collections, pp.

68-70. (reprinted in part from ".\merican Seashells"). In:

.\bbott, R. Tucker, M. K Jacobson, and M. C. Teskey

(Editors). How to Collect Shells. (.\ Symposium). American

Malacological Union, Buffalo, NY [iv] -1-75-1- [vii] pp.

[pp 85-87 in second edition, 1961; pp. 93-95 in third

edition, 1966; pp. 94-95 in fourth edition, 1974]

Abbott, R. Tucker. 1955, Review of "Abbott, R. T., M. K.

Jacobson, and M. C Tesk\ . 1955, Hou to Collect Shells.

American Malacological Union, Buffalo Museum of Sci-

ence. New York" The Nautilus 69(1 );36. [August 1]

Abbott, R. Tucker. 1955. Review of "Perry, L. M. and J. S.

Schwengel. 1955. Marine Shells of the West Coast of

Florida Paleontological Research Institution, Ithaca, NY".

198 pp, 55 pis." The Nautilus 69(1):.36. [August 1]

Abbott, R, Tucker. 1955. Review ol "Bartsch, P. 1955. The

Pyramidellid Mollusks of the Pliocene Deposits of Sorth

St. Petersburg, Florida. Smithsonian Miscellaneous Col-

lections 125(2):1-102, pis, 1-18." The Nautilus 69(I):iii.

[.August 1]

Abbott, R. Tucker. 19.55. Shells of Superstition. Frontiers 20(1);

12-13, inside back cover. [September 20]

Abbott, R. Tucker. 1955. How to Catch Mice. Frontiers 20( 1 );

32. [September 20]

Abbott, R. Tucker. 1955. Anatomy of the Venezuelan Gas-

tropod, Doryssa kappleri. The Nautilus 69(2):44-46, plate

4, figures a-j. [November 5]

Abbott, R. Tucker. 1956. Comments on the Nomenclature of

the Cypraeacea in Kaicher's "Indo-Pacific Sea Shells". Ha-

waiian Shell News 4(7):68-69. [Mas 2]

A[bbott], R. T. 1956. The Kimball \alentine Collection. The

Nautilus 69(41:141. [May 10)

.Abbott, R. Tucker. 1957. Conns niusicns nanus in Hawaii.

Hawaiian Shell News 5(I0):101-105. [August 7]

Abbott, R. Tucker. 1956. Notes on Mitrex scorpio and its

allies. Hawaiian Shell News 4(1I):I13-114 [September 5]

Abbott, R. Tucker 1956. Shell Clubs in America. Proceedings

of the Philadelphia Shell Club l(l):l-5. [September]

.Abbott, R. Tucker. 1957 The Tropical Western .Atlantic Prov-

ince. Proceedings of the Philadelphia Shell Club 1(2):7-

11. [September]

Abbott, R. Tucker. 1957. Prunnni ruscidum in New Jersey.

The Nautilus 71(2):52-53, plate 4, Bgures 4, 4a. [Novem-

ber 4]

Abbott, R. Tucker. 1958. Western Atlantic Marine Mollusks

[Title only]. Bulletin of the American Malacological Union

24:16. [.American Malacological Union Annual Report for

1957] [January 1]

Abbott, R, Tucker, 1958, From the Pilsbr\ Chair of Mala-

cology. The Nautilus 71(3):100-103. [March 4]

.Abbott, R. Tucker. 1958. Redisco\er\ of a Rare Caribbean

Conus. The Nautilus 71(4):117-118. [April 24]

.Abbott, R. Tucker. 1958. A New Recent Species of Tudicula

from Zanzibar (Gastropoda: X'asidae). Notulae Naturae

305:1-4, figures 1-7. [April 25]

.Abbott, R, Tucker, 1958. Shells. National .Audubon Society

Nature Program. Nelson Doubleday, Inc. Garden City,

NY', 56 pp, 29 adhesive stamp plates [April; Second Edi-

tion, 1966, Revised Edition, 1968]

.Abbott, R. Tucker. 1958. The gastropod genus Assiminea in

the Philippines. Proceedings of the .Academy of Natural

Sciences of Philadelphia 11021.3-278, plates 15-25. [Au-

gust 22]

Abbott, R. Tucker. 1958. A Brief Histor> of Shell Auctions.

Buttonwood Farm Shell Auction Catalogue pp. 11-12.

[September 13]

Abbott, R. Tucker. 1958. The Radula of Vuhitoconus grossi

Iredale. Buttonwood Farm Shell .Auction Catalogue p. .35.

[September 13]

Abbott, R. Tucker. 1958. The Mollusk Diar> of Doc Bales.

Proceedings of the Philadelphia Shell C;lub 1(3):6-12. [Oc-

tober]

.Abbott, R. Tucker. 1958. Notes on the .Anatomy of the .Aus-

tralian Volutes, bednalli and grossi. Journal of the Mala-

cological Society of Australia l(2):2-7. [November]

Abbott, R. Tucker. 1958. Marine Mollusca of Rennell Island,

Solomon Islands. The Natural History of Rennell Island,

British Solomon Islands, Copenhagen, 2(28):203-206, plate

20 [December]

.Abbott, R. Tucker. 19.58. The Marine Molluscs of Grand Cay-

man Island, British West Indies. Monographs of the Acad-

emy of Natural Sciences of Philadelphia ll:i-viii, 1-138,

plates 1-5. [December 31] [.A second printing was pub-

lished in October 1967, identical to the first, but with an

inde.x to Common Names, consisting of 16 unnumbered

pages, appended follow ing the plates].

Abbott, R. Tucker. 19.59. The FamiK \asidae in the Indo-

Pacific. Indo-Pacific Mollusca l(l)T5-.32. [March 31]

Abbott, R. Tucker. 1959. How to Collect Shells. /;!. Weyer,

E. M. (ed.). The Natural Sciences Illustrated. J. J Little

and Ives, Co., New Y'ork. pp. 2578-2589.

Abbott, R. Tucker. 1960. Presidential Remarks. [Abstract of

remarks at opening of presented at 26th .Annual Meeting

Page 66

THE NAUTILUS, Vol. 110, No. 2

of the .\inerican Malacological I iiioii, June .jI-JuK 3, 1959]

Bulletin of the .\merican

Malacological I nion 26:2. [American .Malacological I nion .An-

nual Report for 1959] [Januars 1]

.•\bbott, R, Tucker. 1960 Expedition to the Philippine Islands.

Proceedings of the Philadelphia Shell t:lub 1(4)1-6. [April]

Abbott, R. Tucker. 1960. Basic Research — Is It Dying? .

Proceedings of the Philadelphia Shell C:lub l{4):22-25.

[April]

.Abbott, R. Tucker 1960. Gift Irom the Sea of Japan. Frontiers

24(4):120-122. [March 22]

A[tibott], R. T. 1960. Dr. Paul Bart.sch [Note]. The Nautilus

74(1 );33. [July 1]

A[bbott], R. T. 1960. Norman T. Matto.x [Note], The Nautilus

74(1 ):33. [July 1]

A[bbott], R. T. 1960. Department of Living Invertebrates re-

established at the American Museum [Note]. The Nautilus

74(2):S1. [October 5]

Abbott, R. Tucker. 1960. The Genus Strombus in the Indo-

Pacific. Indo-Pacific Mollusca 1(2);33-144. [November 23]

[Baker, H B., C. B. Wurtz and R. Tucker Abbott] Editors.

1961. Jeanne Sanderson Schuengel [notice ot death]. The

Nautilus 74(4):165-166.[April 6]

.Abbott, R. Tucker. 1961. Hov\ to Know American Marine

Shells. Signet Key Book, The New American Library, Nev\-

York. 222 pp. 12 plates. [May; Revised Edition, 1970]

Warmke, G. L. and R. Tucker Abbott. 1961. Caribbean Sea-

shells, A Guide to the Marine Mollusks of Puerto Rico and

other West Indian Islands, Bermuda and the Lower Flor-

ida Keys. Livingston Puhlisliing C^o,, Narbeth, Pennsyl-

vania. i-.\ -I- 1-346 (44 plates). [May; Reprinted in 1975

by Dover Publications]

[Baker, H. B., C. B Wurtz and R. Tucker Abbott] Editors.

1961. Jeanne Sanderson Schwengel, Sc. D., 1889 to 1961.

The Nautilus 75(l);36-39, plate 5. [July 3]

Abbott, R. Tucker. 1961. The Genus Lambis in the Indo-

Pacific. Indo-Pacific Mollusca 1(3):147-174. [September

29]

Abbott, R. Tucker 1961. Forward. In: Jacobson. M. K. and

W. K. Emerson, Shells of the New York City Area. Ar-

gonaut Books, Inc. New York. p. \ii.

Abbott, R. Tucker. 1961. Comments on the proposal to place

the generic name Gari Schumacher, 1817, on the official

list unemended, Z, N. (S.) 1461. Bulletin of Zoological

Nomenclature 1S(5):301. [November 10]

Abbott, R. Tucker. 1961. Oldest Mollusk Research Center in

America. Shells and Their Neighbors 8:1,3. [December]

Abbott, R. Tucker. 1961. Review of ■Cameron, R. 1961.

Shells. G. P. Putnam's Sons, New York. 128 pp. " Frontiers,

December 1961:59. [December]

.Abbott, R. Tucker. 1962. Historical Notes on the American

Journal of Conchology. Sterkiana 6:1-4. [February]

Abbott, R. Tucker. 1962. Review of "Jacobson, M. K. and W

K. Emerson. 1961. Shells of the ^'eu: York City Area.

.Argonaut Books, Larchmont. NY. 142 pp. Frontiers 26(4):

123. [March 22]

Abbott. R. Tucker 1962 Recent I sesoi Non-binoniial Works.

The Veliger 4(4);21.3. [April 1]

Abbott, R. Tucker. 1962. U.se ol the Term "Hvpotypc". The

Veliger 5(2);93-94. [October 1]

Abbott, R. Tucker. 1962. America's oldest mollusk research

center. P'rontiers 26(3):84-85.

Abbott, R. Tucker 1962. Sea Shells of the World. A Guide

to the Better-Known Species. .A Golden Nature Guide.

Golden Press, New York, 160 pp, [Dutch Kdilion, 1963;

French Edition, 1964; Norwegian Edition 1965; Italian

Edition 1966; British Edition, 1967, Danish Edition 1968,

Spanish Edition, 1973, German Edition, 1975, Revised

.American Editions 1969. 1985]

Abbott, Robert Tucker, 1962, Harvard Class ol 1942. Records

of the Class, 1962, Harvard .Alumni .Association, Cam-

bridge, MA. p. 1.

Abbott, R. Tucker. 1963. Further comments on the name of

the type species of Xenophora Fischer von Waldheim,

1807. Bulletin of Zoological Nomenclature 20(1):15. [March

18]

Abbott, R. Tucker. 1963. .A Visit to Some European Shell

Collections, Part I - Great Britain. Proceedings of the

Philadelphia Shell Club 1( 7);25-31, [March]

Abbott, R. Tucker, 1963, The Janthinid Genus Recluzia in

the Western Atlantic, The Nautilus 76(4):151, [April 19]

Abbott, R. Tucker. 1963. Expedition to the Bay of Bengal

[Discussion of paper presented at 29th .Annual Meeting of

the .American Malacological L nion, June 18-21, 1963] Bul-

letin of the .American Malacological L'nioii 30:5, [Decem-

ber 1]

.Abbott, R, Tucker, 1963, Schelpen van der W ereldzeen, J,

M. Meulenhoff, .Amsterdam. 160 pp [Dutch Edition of

"Sea Shells of the World "]

Abbott, R. Tucker 1963. Report on Collections made at Phu-

ket Island, Thailand: International Indian Ocean Expe-

dition, 1963. .Academy of Natural Sciences, Philadelphia,

33 pp, 1 map, [Mimeographed report]

Abbott, R. Tucker, 1964, America's Shelling Mecca, Infor-

mation for Visitors to Sanibel and Captiva, Special Shell

Fair Edition:9, [March]

Abbott, R, Tucker, 1964, Anolher Cypiaea gut tain. Hawaiian

Shell News, 12(6):2. [April]

.Abbott, R Tucker, 1964. Three Conus glorianiaris in Phil-

adelphia. Hawaiian Shell News 12(8):6. [June]

.Abbott, R. Tucker 1964. Littorina ziczac (Gmelin) and L.

lineolata Orbignv. The Nautilus 7S(2):65-66. [October 15]

Wagner, Robert J.\.'and R. Tucker Abbott. [Editors] 1964

Van Nostrand's Standard Catalogue of Shells. [First Edi-

tion]. D. \an Nostrand Co. Inc., Princeton, N], 190 pp.

[November 16, 1964; Errata, dated March 1965; Second

printing with Errata (pages 192-195), May 1965]

Abbott. R. Tucker. 1964. Indo-Pacific Mollusca. [Abstract and

Discussion of paper presented at .30th .Annual Meeting of

the .American Malacological L'nion. July 21-24, 1964] Bul-

letin of the American Malacological l'nion 31:11. [.Amer-

ican malacological L nion .\nnual Report for 1964] [De-

cember 1]

Abbott, R. Tucker. 1964. Report on the Marine Mollusks

Collected in the Seychelles for the International Iiulian

Ocean Expedition. .Academy of Natural Sciences, Phila-

delphia. 6 pp. [Mimeographed report]

Abbott, R. Tucker. 1964. Report on the Marine Mollusks

Collected in the Bav of Bengal on Oiiise One ot the ".Anton

Bruun". Department of Mollusks. The .Academv of Nat-

ural Sciences of Philadelphia 34 pp. [Mimeographed re-

port]

.Abbott, R. Tucker. 1964. Cxjquillages. Especes du Monde

Entier. Le Petit Guide, Hachette. 160 pp. [French Edition

of "Sea Shells of the World"]

Abbott, R. Tucker 1965. Review of "Kennelly, D. H. 1964.

Marine Shells oj Southern Africa. Frontiers 29(3):94.

[February]

.Abbott. R Tucker 19()5, Miocene .S/ro)ii//!/.v {0(;/o;(i(7i(i 1 Iroiii

India, Indo-Pacific Mollusca U6):401-402, [April .30]

M. G. Harasewych, 1997

Page 67

Abbott, R- Tucker. 1965. Seashells from Cat Food. Hawaiian

Shell News 13(12):6. [October]

.\bbolt, R. Tucker. 1965. Cypiuca arenona C.ra\. 1824. Ha-

waiian Shell News 14(2):8. [December]

Abbott, R. Tucker. 1965. The wonderful world of shells Sci-

ence Digest 5?>i6);66-73. [December]

Abbott, R. Tucker. 1965. Giant Clams. Frontiers 30(2):46-

49. [December]

Abbott, R. Tucker. 1965. Snegler og Skjell: Fra Hele Verden.

Fredhois Forlag A/S, Olso. 160 pp. [Norwegian Edition

of "Sea Shells of the World"]

Abbott, R. Tucker. 1965. The Shell Desk t:alendar lor 1966.

Shell Oil Co., New York. 120 pp.

.-Vbbott, R. Tucker. 1966. Concholog): Queen ol the Natural

Sciences. Frontiers 30(3):68-73. [Februar\ ]

Abbott, R. Tucker. 1966. Curator's Choice: Glor\-of-India

Cone. Frontiers 30(3):90. [Februar>]

Abbott, R Tucker. 1966. Conchiglie, Specie di Tutto il Mon-

do. Mondadori, Milan. 160 pp. [Italian Edition of "Sea

Shells of the \\'orld"] [Februarv ]

Abbott. R. Tucker. 1966. Quiz-Me: Seashells. .\ Junior Golden

Guide. Golden Press, New \ork 4S pp [.\pril. Braille

Edition, 1967]

Abbott, R. Tucker. 1966. Comments on the proposed vali-

dation of Valuta episcopalis Linnaeus, 1758. Z.N.{S.) 1728.

Bulletin of Zoological Nomenclature 23(2/3 ):80. [July 29]

Abbott, R. Tucker. 1966. Comment on A/i(ra per/a/a Roding,

1798, as a nomen oblitum. Z.N.{S.) 1726. Bulletin of Zoo-

logical Nomenclature 23(2/3);90. [July 29]

Abbott, R. Tucker. 1966. New England's Rarities, 1672. New

York Shell Club Notes 1245. [September]

Abbot [sic], R. Tucker. 1966. Shells. Nelson Doubledav , Inc.

and Odham Books, Ltd. 63 pp.. 29 adhesive stamp plates.

[Revised Edition, 1968]

Abbott, R. Tucker. 1966. The Shell Desk Calendar for 1967,

With Illustrations by Several Hands Showing Sea Shells in

Aspects Practical, Scientific, and Ornamental Shell Oil

Co., New York. 120 pp.

Abbott, R. Tucker. 1967. Re\iew of "Dance, S. P. 1966.

Shell Collecting: An Illustrated History." Frontiers 1(3):

66-67. [February 1]

Abbott, R. Tucker, and Russell H Jensen. 1967. Molluscan

Faunal Changes around Bermuda. Science 155(3763):687-

688. [Februar^ 10]

Abbott, R. Tucker. 1967. Review of "Dance, S P. 1966.

Shell Collecting: An Illustrated History." Natural Historv

76(3):66-67. [March]

Abbott, R. Tucker. 1967. N'enom apparatus and Geographical

Distribution of Conus glorumwn.'i. Notulae Naturae 400:

1-8. [May 3]

Jung, Peter and R. Tucker .Abbott. 1967. The Genus Tere-

bellum (Gastropoda: Strombidae). Indo-Pacific Mollusca

l(7):445-454. [May 15]

Abbott, R. Tucker. 1967. Strombus (Canarium) wilsoni. new-

species from the Indo-Pacific Indo-Pacific Mollusca 1(7):

455-456. [May 15]

Wagner, R. J. L. and R. Tucker Abbott. [Editors] 1967. \'an

Nostrand's Standard Catalogue of Shells. [Second Edition].

Van Nostrand Co. Inc., Princeton, NJ, i-xi + 1-303. [Sep-

tember 19]

Abbott, R. Tucker. 1967. Review of 'Tebble, N. 1966. Brit-

ish Bivalve Seashells. A Handbook for Identification.

Trustees of the British Museum (Natural History) London.

212 pp + 12 pis." Quarterly Review of Biology 43(3):343-

344. [November 25]

.Abbott, R. Tucker ami H, S. Zim. [co-authorship misattributed]

1967. Sea Shells of the World. A Guide to the Better-

Known Species .\ Little Guide in Colour. Paul Hamlyn

Ltd, London. 160 pp. [British Edition of "Seashells of the

world "]

Abbott, Robert Tucker. 1967. Harvard Class of 1942, Records

of the Class, 1967. Harvard Alumni .Association, Cam-

bridge, MA. p. 1.

Abbott, R Tucker. 1967. Re\iew of "Dance, S. P. 1966.

Shell Collecting: An Illustrated History." Quarterly Re-

view of Biolog) 42:549-550.

Abbott, R. Tucker. 'l967. Review of "Tebble, N. 1966. Brit-

ish Bivalve Seashells. A Handbook for Identification.

Trustees of the British Museum (Natural History) London.

212 pp -I- 12 pis." Frontiers 32(2):30.

.Abbott, R. Tucker. 1967. Terebellum's Blue-E\ ed Periscope

Frontiers December :8-9. [December]

Abbott, R. Tucker. 1967. Quiz-Me; Seashells, .A Junior Golden

Guide. Twin Vision, California. 37 pp. [Braille Edition]

Abbott, R. Tucker. 1967. The Shell Desk Calendar for 1968,

.A Journal Book Well .Adapted for the Purpose of .Noting

Events and Expectations w itli the Greatest Readiness, Be-

ing Calculated to Sa\e Much tune and Trouble. Shell Oil

Co., New York. 120 pp.

.Abbott, R. Tucker, and Russell H. Jensen. 1968. Portuguese

Marine MoUusks in Bermuda. The Nautilus 8 1(3): 86-89.

[Januar\' 25].

Abbott. R Tucker 19fj8. Giant Cowries. The Nautilus 82(1):

32. [July 24]

Abbott, R.Tucker. 1968. The Helmet Shells of the World

(Cassidae). Part 1. Indo-Pacific Mollusca 2(9):15-202. [Au-

gust 30]

.Abbott, R Tucker. 1968. [Editor] William Swainson's "Exotic

ConchologN," .A Facsimile Edition of the 1841 Edition,

with Essential Excerpts from the First and Second Issues

U821-1835I. Delaware Museum of .Natural History, Re-

production Series, Number 1. D. \'an Nostrand & Com-

pan>, Princeton, NJ. i-xxi\ -(- 4-47 4- 48 plates. [.Abbott

contributed the Preface (v) and the Modern Explanation

of Plates (41-47)] [September 11]

Abbott, R Tucker. 1968. Shells. Nelson Doubleday, Inc. and

Odham Books, Ltd. 63 pp. 29 adhesive stamp plates [Re-

vised Edition] [September]

Abbott, R. Tucker 1968. Review ot "Knudseii, J 1967. The

Deep-Sea Bivalvia. The John Murray Expedition, 1933-

34, Scientific Reports ll(3);235-343 -t- 3 pis." Quarterly

Review of Biology 43(3):344. [September]

Stix, Hugh, Marguerite Stix and R. Tucker .Abbott [photographs

b> H. Landshoft ]. 1968. The Shell, Five Hundred Million

\ ears of Inspired Design. Harr\ N. .Abrams, Inc. New York.

188 plates, plus unpaginated text. [October; French Edi-

tion. 1969; Italian Edition 1969; .Abridged English Editions

1972. 1973, 1978, 1988]

.Abbott, R. Tucker and Herbert S. Zim. [co-authorship misat-

tributed] 1968. Fakta om Ivonkylier. Lademann, Copen-

hagen 160 pp. [Danish Edition of "Sea Shells of the World ']

[November 1]

.Abbott, R. Tucker [recorded b\]. 1968. Pronouncing the Sci-

entific .Names of Seashells of North .America. R. Tucker

.Abbott and the Delaware Museum of Natural History,

Greenville, Delaware. One, High Fidelit\, 33 13 RPM

recording (36 minutes).

.Abbott, R Tucker [Illustrations b\ George F. Sandstrom]. 1969.

Seashells of North .America; .A Guide to Field Identifica-

Page 68

THE NAUTILUS, Vol. 110, No. 2

tion. Golden Press, New York. 2.S0 pp [Revised edition,

1986] [January]

Richards. Horace G , R. Tucker .\bl)ott. and Thomas Skymer.

1969. Marine Pleistocene Mollusks from Bermuda. The

Geological Societ\' of .America, Abstracts with Programs

for 1969 (4):69. [.\bstract] [April 9]

Richards, Horace G., R. Tucker .Abbott, and Thomas Skymer.

1969. The Marine Pleistocene Mollusks of Bermuda. No-

tulae Naturae 425:1-10. [October 10]

Sti.\, Hugh, Marguerite Stix and R. Tucker Abbott [photographs

by H. Landshoff]. 1969. Le Conchiglie. Garzanti, Italy.

188 plates, plus unpaginated text. [October 15]

.Abbott, R. Tucker. 1969. Achatina fulica invades Florida.

The Nautilus 83(2):75. [October 31]

Stix, Hugh, Marguerite Stix and R. Tucker Abbot [sic] [pho-

tographs by H. Landshoff]. 1969. Les Coquillages, les

chefs-d oeuvre de la vie sous-marine. Seghers, Paris. 188

plates, plus unpaginated text. [November 25]

Abbott. R Tucker. 1969. The Shell Desk Calendar for 1970

Shell Oil Co., New York. 116 pp.

Abbott, R. Tucker and Hal Lewis. 1970. Cymatiitm boschi.

New Species from the Arabian Sea. The Nautilus 83(3):

86-88. [January 23]

Abbott, R. Tucker. 1970. .-Kmerican Malacological Union

Symposium on the rare and endangered mollusks of North

America, 7. Eastern marine mollusks. Malacologia 10:47-

49. [May]

Abbott, R. Tucker. 1970. How to Know American Marine

Shells [Revised Edition]. Signet Key Book, The New Amer-

ican Library, New York. 222 pp. 12 plates. [May]

Abbott, R. Tucker. 1970. Conchology: Queen of the Natural

Sciences. La Conchiglia 11(1 1-12) [21-22]:14-15. [Decem-

ber]

Wagner, R. J. L. and R. Tucker Abbott. 1970. Van Nostrand's

MonthK Supplement. Number 1. New subgenera of Con us

proposed since 1966. Value changes and additions La

Conchiglia II (ll-12)[21-22]:3, 22-23. [December] *

Abbott, R. Tucker. 1971 The Shell Desk Calendar for 1971,

with photographs suggesting the color, range and balance

of life in the sea. The Shell companies. New York. 116 pp.

Wagner, R J. L. and R. Tucker Abbott. 1971. Van Nostrand's

Monthly Supplement. (Number 2, January, 1971). The Vo-

lutes (Volutidae). La Conchiglia III (1)[23]:15. [January] *

Wagner, R. J. L. and R. Tucker Abbott. 1971. Van Nostrand's

Monthly Supplement. Continuation of Number 2. The

Volutes (Volutidae), La Conchiglia III (2)[24];15. [Feb-

ruary] *

Wagner, R. J. L. and R. Tucker Abbott. 1971. Supplemento

mensile al catalogo del Van Xostrand. La Conchiglia III

(3)[25]: 15. [March]*

Abbott, R. Tucker. 1971. .-^re marine mollusks endangeredr'

La Conchiglia III (4)[26]:1 1-12. [April]

Wagner, R J L. and R Tucker Abbott 1971 \an Nostrand's

*.\ccording to an announceinent in La Coiwhiglia II( 1 1-12)[21-

22];3 (1970), supplements containing value changes in Van

Soxtrand's Standard Catalog of Hhell.s (Second Edition) were

originally published in issues of La Conchiglia. These were

subse(|ueiitly reprinted (unpaginated) in a format suitable for

insertion into the second edition ot Van Nostrand s Standard

Catalog of Shells and sold separately.

Monthly Supplement. (Number 5, .\pril, 1971 ) The Cow-

ries (Cypraeidae). La Conchiglia III (4,l[26]:15. [.April] *

Wagner, R. J. L. and R. Tucker Abbott 1971. Supplemento

mensile al catalogo del \ an Nostrand, \ an Nostrand s

Catalog Monthlv Supplement Familia Cvpraeidae. La

Conchiglia III (5)[27]:15. [May]*

Wagner, R. J. L. and R. Tucker Abbott. 1971. Supplemento

mensile al catalogo del \'an Nostrand, Van Nostrand's

Catalog MonthK Supplement. Familia Cvpraeidae La

Conchiglia III (6)[28]:15. [June] *

Abbott, R. Tucker and Charles B. Wurtz. 1971. Horace Bur-

rington Baker, 1889-1971. The Nautilus 85(l):l-4. [July

20]

Wagner, R. J. L. and R. Tucker Abbott. 1971 Family Cy-

praeidae. Van Nostrand's Monthly Supplement. Number

3. La Conchiglia 30: insert of 8 unnumbered pages. [Sep-

tember] *

Abbott, R. Tucker. 1971. Conus patae, \ .New Caribbean

Gastropod. The Nautilus 85(2):49-51. [October 4].

Wagner, R. J. L. and R Tucker Abbott 1971 Van Nostrand's

Monthly Supplement. (Number 10, October 1971). Value

Changes and Additions to Strombidae, .Additions to the

Helmet Shells (Cassidae). La Conchiglia III (10)[32]:15.

[October] *

Abbott, R. Tucker. 1971. Seashell Safari to the Solomons. New

York Shell Club Notes 176:3-5. [November]

Abbott, R. Tucker. 1971. Bivalves, Magnificent Jewelry of

the Sea. International Wildlife l(6):58-63. [November]

Wagner, R. J. L. and R. Tucker Abbott. 1971. Van Nostrand's

Monthly Supplement. (Number 11-12, Nov. -Dec. 1971).

Additions to the Helmet Shells (Cassidae). La Conchiglia

III (11-12)[33-34]:15. [November-December]*

Abbott, R. Tucker. 1971. The Shell Desk Calendar for 1972,

with ilustrations showing how the art of many ages has

been enriched b\ the form and s> mmetrv of the shell. The

Shell Companies, New York. 116 pp.

Wagner, R. J. L. and R. Tucker Abbott. 1972. \aii Nostrand's

Monthly Supplement. (Number 1, January 1972). .Addi-

tions to the Helmet Shells (Cassidae). La Conchiglia I\'

(1)[35]:15. [January]*

Wagner, R. J L. and R Tucker Abbott. 1972. Van Nostrand's

Monthly Supplement. (Number 2, Februarv 1972). .Ad-

ditions to the Helmet Shells (C:assidae). La Conchiglia IV

2(36):15. [February] *

Wagner, R. J. L. and R. Tucker Abbott. 1972. Van Nostrand's

Monthly Supplement. (Number 2, February 1972). Ad-

ditions to the Helmet Shells (Cassidae). La Conchiglia IV

3(37):15. [March]*

Wagner, R. J. L. and R Tucker Abbott. 1972 \aii Nostrand's

Monthly Supplement. (.Number 2, Februarv 1972). .Ad-

ditions to the Helmet Shells (Cassidae). La Conchiglia I\'

4(38): 15. [April]*

Abbott, R. Tucker. 1972. Review of "Keen, A. M. 1971. Sea

Shells of Tropical West America." Quarterlv Reviev\ of

Biology 47:240. [June]

Abbott, R Tucker. 1972. C:onchologv— Hobbv ami Science.

Jacksonville Shell Show Program:7, [JuK 29]

Abbott, R. Tucker. 1972. New MoUusk Exhibits in America.

La Conchiglia IV 7-8(41-42):220-21. [July-August]

Wagner, R. J L and R. Tucker Abbott. 1972. Van Nostrand's

Monthly Supplement. (Number 2, February 1972). .Ad-

ditions to the Helmet Shells (Classidae). La Conchiglia I\'

7-8(41-42):23, [July-August] *

Abbott, R, Tucker. 1972. Kingdom of the Seashell. Crouii

M G. Harasewych, 1997

Page 69

Publishers, Inc., New York. 256 pp. [August] [French Edi-

tion, 1974; Itahan Edition, 1974; Revised Edition, 1993]

Keen, A, M. and R. Tucker Abbott. 1972. Problem of the

type species of Lucina (Mollusca: Pelecypoda) Z.N.(S. )

2001. Bulletin of Zoological Nomenclature 29{3):15S-161.

[No\ ember 30]

Abbott, R. Tucker. 1972. The Bermuda "VVilke." Newsletter,

Bermuda Biological Station lor Research. 2(1)2. [Decem-

ber]

Salvini-Plawen, L. v. and R. Tucker Abbott. 1972. The Mol-

lusks. pp. 19-28. In: Grzimek, B., O. Kraus, and R. Reidel

(eds.) Grzimek's Animal Life Encyclopedia. Volume 3.

Mollusks and Echinoderms. Van Nostrand Reinhold Com-

pany, New York. 541 pp [paperback edition, 1984]

Salvini-Plawen, L. v. and R. Tucker .\bbott. 1972. The So-

lenogasters and Chitons, pp. 29-42. In: Grzimek, B., O.

Kraus, and R. Reidel (eds.) Grzimek's .Animal Life En-

cyclopedia. Volume 3. Mollusks and Echinoderms. Van

Nostrand Reinhold Company, New York. 541 pp [paper-

back edition, 1984]

Salvini-Plawen, L. v. and R. Tucker Abbott. 1972. The Shelled

Mollusks pp. 43-46. /().■ Grzimek, B , O. Kraus, and R

Reidel (eds.) Grzimek's Animal Life Encyclopedia. \'ol-

ume 3. Mollusks and Echinoderms. Van Nostrand Rein-

hold Companv, New York. 541 pp. [paperback edition,

1984]

Salvini-Plawen, L. \'. and R. Tucker .Abbott. 1972. The Gas-

troverms. pp. 47-49. In: Grzimek, B., O. Kraus, and R

Reidel (eds.) Grzimek's Animal Life Encyclopedia. Vol-

ume 3. Mollusks and Echinoderms. \'an Nostrand Rein-

hold Companv, New York, 541 pp. [paperback edition,

1984]

Salvini-Plav\en, L. v. and R. Tucker ,\bbott 1972. The Gas-

tropods, pp. 50-135. In: Grzimek, B , O Kraus, and R.

Reidel (eds.) Grzimek's Animal Life Enc\clopedia. Vol-

ume 3. Mollusks and Echinoderms. \an .Nostrand Rein-

hold Companv, New York. 541 pp. [paperback edition,

1984]

Salvini-Plawen, L. \ . and R. Tucker Abbott. 1972. Scaphopods

and Bivalves, pp. 136-189. In: Grzimek, B., O. Kraus, and

R. Reidel (eds.) Grzimek's .\nimal Life Encyclopedia. Vol-

ume 3. Mollusks and Echmoderms. Van Nostrand Rein-

hold Companv, New York. 541 pp. [paperback edition,

1984]

Stix, Hugh, Marguerite Sti.v and R, Tucker .Alibott [photographs

by H. Landshoff ], 1972. The Shell, Five Hundred Million

years of Inspired Design. Ballaiitine Books, Inc. New York.

[Abridged version of 1968 eilition] 135 plates, plus unpa-

ginated text.

Abbott, Robert Tucker, 1972. Har\ ard Class of 1942, Records

of the Class, 1972. Har\ard .-Xlumni Association, Cam-

bridge, \\.\ p 1.

Abbott, R. Tucker. 1973. Review of "Halliday, P. 1972

Snailij Verse. \'olturna Press, Scotland. 48 pp. " The Nau-

tilus 87(1 ):4. [January 24]

Abbott, R Tucker. 1973. Review of 'Goodman, S. and L.

Goodman. .1972. Art From Shells. Crown Publishing

Co., New York. 210 pp." The Nautilus 87(1 ):23. [Januarv

24]

Abbott, R. Tucker. 1973. Spread ot Melanuidcs tiibcrculata

The Nautilus 87(1 ):29. [January 24]

Abbott, R. Tucker. 1973. Review of "Boss, K. J. 1972. The

Genus Semele in the ivestern Atlantic. Johnsonia 5(49):

1-32." The Nautilus 87(1 ):29. [Januar\ 24]

Abbott, R. Tucker. 1973. Mollusks Dangerous to SCLBA di-

vers. Of Sea and Shore 3(4):161-162. [January]

Sandved, Kjell B. and R. Tucker Abbott. 1973 Shells in Color

\'iking Press, New York. 112 pp. [June]

Abbott, R. Tucker. 1973. Review of "Jenkins, M. M. 1972.

The Curious Mollusks. Holiday House, New York. 224

pp." Quarterly Review of Biology 48:376. [June]

Abbott, R. Tucker. 1973. Review of "Andrews, J. 1971. Sea

Shells of the Texas Coast. Lhiiversity of Texas Press, Aus-

tin, xvii + 298 pp." Quarterly Review of Biology 48:376.

[June]

Abbott, R. Tucker, 1973. In search of seashell treasures. Sci-

ence Digest 74(1):16-21. [July]

Abbott, R. Tucker. 1973. Mollusks Dangerous to SCUBA Di-

vers. Delaware Medical Journal 45(6):161-164. [Jul\]

Abbott, R. Tucker. 1973. Acleon etoiseae, a New Opistho-

branch from Arabia. The Nautilus 87(4):91-92. [October 4]

Abbott, R. Tucker. 1973 Review of "Furlong, M. and \' Pill

1972. Edible}' Incredible:' Ellis Robinson Publishing Co."

The Nautilus 86(2-4):66. [December 1]

Abbott, R. Tucker. 1973. Mollusks Dangerous to SCUBA Di-

vers. The Shell Case [Naples Shell Club] 3(l);5-6.

Abbott, R. Tucker. 1973. Review of "Bequart, J. C. and W.

B. Miller. 1973. The Mollusks oj the Arid Southivest.

The University of Arizona Press, Tucson, xvi -(-271 pp."

QuarterK Review of Biolog\' 48:644-645. [December]

Abbott, R. Tucker. 1973. What do Shells Tell':' Young Nat-

uralist 13(12):25-31. [December] [In Russian — translated

by V. Lesiyovsky]

Abbott, R. Tucker [Editor]. 1973. American Malacologists, A

National Register of Professional and Amateur Malacol-

ogists and Pri\'ate Shell Collectors and Biographies of Early

American Mollusk Workers Born Between 1618 and 1900.

First Edition. American Malacologists, Falls Church, \'ir-

ginia. i-iv + 1-494.

Stix, Hugh, Marguerite Stix and R. Tucker Abbott [photographs

by H. Landshoff]. 1973. The Shell. Harry N. Abrams,

NY [Abridged version ot 1968 edition] 135 plates, plus

unpaginated text.

.•\bbott, R. Tucker. 1973. Conchas marinas, Ediciones Dia-

mon, Barcelona, 1(50 pp, [Spanish Edition of "Sea Shells

of the World"]

.Abbott, R, Tucker. 1974. A tribute to a conchological bicen-

tenary. New York Shell Club Notes 200:2. [March 18]

.•\bbott, R. Tucker. 1974. American Malacologists — A Bio-

graphical Survey (1618-1973). Bulletin of the American

Malacological Union, Inc. for 1973:8-11. [May]

Abbott, R. Tucker, 1974, .American Seashells, The Marine

Mollusca of the Atlantic and Pacific Coasts of North .Amer-

ica, Second Edition, \an Nostrand-Reinhold Publishing

Company, New York. 663 pp. [October]

.Abbott, R. Tucker. 1974. Au Royaume des Coquillages. Edi-

tions des Deux Coqs d'Or, Paris. 255 pp [Kingdom of the

Seashell, French Edition]

Abbott, R. Tucker. 1974. II Meraviglioso Monde della Con-

chiglie. .Arnoldo Mondadori Editore, Milan. 256 pp. [King-

dom of the Seashell, Italian Edition]

Abbott, R Tucker. 1975. Obituary, John Quincy Burch (1894-

1974). The Nautilus 89(1):12-13. [January 31]

Abbott, R. Tucker. 1975. Beware the Asiatic Freshwater Clam.

Tropical Fish Hobbyist 23(6): 15. [February]

Abbott, R. Tucker. 1975. The Favored Shell. The Original

Tide-ings 5(4/5):55. [April]

Abbott, R. Tucker. 1975 Thomas Sav — America's First Ma-

Page 70

THE NAUTILUS, Vol. 110, No. 2

laeologist. [.Abstract] Bullt-tiii o\ tht- .\merican Malacolog-

ical Union, Inc. for 1974:66, [May)

Abbott, R Tucker [Editor], 197.5, American Malacologists, A

National Register of Professional and .\mateur Malacol-

ogists and Pri\ ate Shell Collectors. Supplement. American

Malacologists, Greenville, Delaware, pp. 495-609. [June

or later]

Abbott, R. Tucker. 1975. Shells and Our \ anishing World.

Jacksonville Shell Show Program: 9. [JuK 26]

.Abbott, R. Tucker, 1975, Note on Thais haemastoma. Te.xas

Conchologist 12i2):30-31, [December]

Abbott. R, Tucker, 1975. Muscheln und Schnecken des Meeres.

Delphin Verlag, Stuttgart and Zurich. 160 pp. [German

Edition of "Sea Shells of the World"]

.Abbott, R. Tucker, 1976. Cittarium pica (Trochidae) in Flor-

ida. The Nautilus 90(I):24. [January 30].

Abbott, R. Tucker. 1976. Review of "Kay, E. A. and W. J.

Clench, 1975, A Bibliography of William Harper Pease.

Malacologist oj Polynesia. Nemouria 16:1-50, Hawaiian

Shell News 24( 1 ): 10 [January ]

Abbott. R Tucker 1976 Seashells, Ridge Press, Bantam Books,

New York. 160 pp. [subsequent printings, 1978, 1983]

[February]

Abbott, R. Tucker, 1976, Review of "Murray, S. B, 1975.

Seashell Collector's Handbook 6 Identifier. Covering the

Shells of the Coasts of the .Anxericas. Sterling Publishing

Co., New York. 240 pp. (Quarterly Review of Biology

51(1):149, [March]

Abbott, R, Tucker, 1976, Early Conchological Immigrants.

Jacksonville Shell Show Program for 1976:16-17. [July]

Abbott, R. Tucker [Editor]. 1976. The Best of T/ie iVauJi/us,

\ Bicentennial Anthology of American Conchology.

American Malacologists, Inc., Greenville, Delaware, i-viii,

1-280.

Abbott, R. Tucker. 1977. In Memonum [W illard Lee Mo-

horter and John Dyas Parker]. New sletter. .American Mal-

acological Union 8(2):2. [Spring]

.Abbott, R. Tucker. 1977, Shells — .An Emperor's Treasure.

Palm Beach Life 70(7):17-19, 56-57. [July]

Abbott, R. Tucker, 1977. Review of "Yonge, C. M. and T.

E. Thompson. 1976, Living Marine Molluscs. William

Collins & Son, London. 288 pp." Quarterly Review of

Biolog) 52(4):438. [Decemljer]

Abbott, R. Tucker and P. Alcolado M. 1977. Bivalves. FAO

Species Identification Sheets. Fishing .Area 31. (W. Central

Atlantic) 37 pp.

Abbott, R. Tucker and P. Alcolado M. 1977. Gastropods. FAO

Species Identification Sheets. P'ishing Area 31. (W. Central

Atlantic) 18 pp.

Abbott, R. Tucker. 1977. Chitons. F"AO Species Identification

Sheets. Fishing .Area 31. (W, Central .Atlantic) 2 pp

Abbott. Robert Tucker. 1977 Harvard C:lass of 1942, Records

of the Class, 1977. Harvaril .\luinni Association. Cam-

bridge, M.A, p 1,

Abbott, R Tucker, 1977. EarK Shell ( iollet tors in Florida,

The Shell Case [Naples, FLJ 5(1): 1-2

V\agner, Robert J. L. and R. Tucker Abbott [Editors], 1978.

Standard t^atalogue of Shells. [Third Edition]. American

Malacologists, liic , Greenville, Delaware i-v -I- 1-17 +

misc. cosmically numliered pages. [February]

Abbott, R, Tucker, 1978 Review of "Andrews, J. 1977. Shells

and Shores of Texas " Science Books and Films 14(1):25.

[May]

Wagner, Robert J L and R Tucker Al)l)otl[EdU()rs]. 1978

Standard Catalogue of Shells, [Supplement 1] .\merican

Malacologists, Inc, Greenville, Delaware, 28 pp, [Septem-

ber]

Sti,\, Hugh, Marguerite Stix and R Tucker .Abbott [photographs

by H. Landshoff]. 1978, The Shell. Harry N, Abrams,

Inc. New York. 163 pp. [a concise version of the 1968

edition; Japanese edition of this version, 1980]

.Abbott, R. Tucker. 1979, The shell game. Sojourn, Summer

1979:31-33. [July]

Richardson, L,, R, Tucker Abbott, and G. M. Davis, 1979.

Early references to the figures in the Conchylien Cabinet

of Martini and

Chemnitz: Volumes 1-12. Tryonia 2(l):l-225; 2(2):226-427,

[.August 1]

Abbott, R, Tucker, 1979, The Shell Game. Diversion 7(8):

118-120. [August]

.Abbott, R, Tucker, and C. John Finlay , 1979, Chicoreus cos-

mani. .A new Muricid Gastropod from the West Indies.

The Nautilus 94(4): 1.59-162. [October 30J

.Abbott, R, Tucker, 1980. Further notes on Lambis. .New York

Shell Club Notes 258:1-2. [January]

Abbott, R Tucker, 1980. Are shells disappearing':' Shell Col-

lector 2:47-49. [July]

Abbott, R, Tucker, 1980, A misfit, tortuous tellin. Shell Col-

lector 2:60. [July]

Abbott, R. Tucker. 1980 Morris Karl Jacobson (1906-1980).

An Obituary. The Nautilus 94(4):129. [October ,30].

Abbott, R, Tucker, 1980 The shell trade in Florida, status,

trade, and legislation. Trade Records .Analysis of Flora and

Fauna in Commerce (TR.AFFIC) USA Special Report 3:1-

85. [October]

.Abbott, R. Tucker. 1980, Feed-back [letter], Conchologists of

America Bulletin 22:2. [December]

Stix, Hugh. Marguerite Stix and R. Tucker .Abbott [photographs

by H, Landshoff ], 1980, Kai, sonobunkatobi, [Shells,Their

culture and beauty] Harry N. .Abrams, Inc, New York,

163 pp, [Japanese edition of 1978 version]

■Abbott, R, Tucker 1981 Review of "Gordon, .A, and Lorna

Strong Melvin, 1981, l.UUO World Sea Shells: Rare to

Common, icith Values. Charles E, Tuttle Company , Rut-

land VT " The Quarterly Review of Biology 56(2):182.

[June]

.Abbott, R. Tucker, 1981, Some comments on that mystery

Hydatinid in color, Hawaiian Shell .News 29(1 1):6. [No-

vember]

.Abbott, R. Tucker and S Peter Dance. 1981. Compendium

of Seashells, E, P. Dutton, Inc., New York, NY, x -I- 411

pp. [December; Copyrights. 1981, 1983. 1986, 1990; Sec-

ond Edition 1983, Third Edition 1986, reprinted 1990;

Japanese Edition, 1985; .Australian Edition, 1990]

Abbott, R. Tucker 1982, Catalogue of the Recent Shells of

Georgia— 1849, The Shell-O-Gram 23(3):3. [March]

Abbott, R. Tucker, 1982. The C. O, A, Sanibel Sojourn of

1982, Conchologists of America Bulletin 10(3): 1,8-9, [Sep-

tember]

Abbott, R, Tucker 1982 Review of "Rehder. H A 1981

The .Kudubon Field Guide to Sorth American Seashells.

Alfred .A. Knopf, New York. 894 pp " New York Shell

club Notes 285:6. [December]

Wagner, Robert J. L, and R. Tucker Abbott [Editors]. 1982.

Standaril Catalogue of Shells. [Supplement 2], .American

.Malacologists. Inc., Melbourne, Florida. 28 pp.

Abbott, R, Tucker, 1983, Harrv S(tephen) Ladd, The Nautilus

97(1 );43, [January 31].

Abbott, R Tucker. 1983. C;harles B. Wurtz. The Nautilus

97(1):43, [January 31],

M G. Harasewych, 1997

Page 71

Ahhdtt. R, Tucker. 1983. William Euwood) Old. Jr The

Nautilus 97(l);4o-44. [Jaiuiarv 31]

Abbott, R. Tucker. 1983. Reviews ol "Relider. H. A. 1981.

The Audubori Society Field guide to Sorth American

S/ie//s, Chanticleer Press.; Clarke, A. H. 1982. The Fresh-

water Mollusks of Canada. The University of Chicago

Press; and Smythe, K. 1982. Seashells of the Arabian

Gulf" The Quarterly Review of Biolog> 58(1 ):70 [March]

Abbott, R. Tucker and S. Peter Dance. 1983. Compendium

of 8easliells. E. P. Dutton, Inc., New York, NY, x + 411

pp. [Second Edition]

Deisler, Jane E. and R. Tucker .\bbott. 1984. Range Exten-

sions of some Introduced Land Mollusks in the Bahama

Islands, with First Reports for Four Species. The Nautilus

98a):12-17. [January 30J

Abbott, R. Tucker and S Peter Dance. 1984. Compendium

comment. Hawaiian Shell News 32(I):6. [January]

.•Xbbott, R- Tucker. 1984. Giant Octopus comes Ashore. New

York Shell Club Notes 290: 13. [March]

Abbott, R. Tucker. 1984. A Farewell to Bill C:lench. The

Nautilus 98(2):55-58. [April 27J

Abbott, R. Tucker. 1984. Review of "Jokinen, E. H. 1983.

The Freshuater Snads of Connecticut. State Geological

and Natural History Survey of Connecticut, vii + 83 pp."

The Nautilus 98{2):79. [reprinted New York Shell Club

Notes 292:3-4]

Abbott, R. Tucker. 1984. Collectible Florida Shells, American

Malacologists, Inc. Melbourne, Forida, 1-64, [JuK] [Pub-

lished in Regular and Waterproot Editions] [Re-issued the

same year as "Collectible Shells ot Southeastern L .S., Ba-

hamas & Caribbean"]

Abbott, R. Tucker. 1984. "Collectible Shells ol Southeastern

U.S., Bahamas & Caribbean [Regular and Waterproof

Edition], .\merican Malacologists, Inc., Melbourne, Flor-

ida. [Identical to above, except for title]

Abbott, R T. 1984. Obituary [Zinaida A. Filatova], The Nau-

tilus 98(4):166. [October 31]

Abbott, R. Tucker. 1984. Seashells. pp. 103-109. In: Griffis,

Nixon (ed. ) The Mariner's Guide to Oceanograpin . Hearst

Marine Books, New York. 272 pp.. Ifci plates. [October]

Abbott, R. Tucker. 1985. IRS Not Charitable to Some Con-

chologists. Conchologists of .America Bulletin 13(2):22

[June]

Abbott, R. Tucker. 1985. Review of "Anderson, R. 1985.

Guide to Florida Seashells. Winner Enterprises. Florida.

56 pp." Shell-0-Gram [Jacksonville Shell Club] 26(4):6.

[July-August][reprinted The Busycon [BrowartI Shell C'lub]

20(8);2-3; New York Shell Club Notes 29610-11, Ha-

waiian Shell News 33(9):9.]

.•\bbott, R. Tucker. 1985. Kirk .\nders — Shell Guide Supreme

(1945-1985). Thatcheria 20(9):[2] [November]

Abbott, R. Tucker. 1985. Kirk Anders, 1945-1985. Hawaiian

Shell news 33(12):12. [December]

Abbott, R. Tucker. 1985. Kirk Anders. New York Shell Club

Notes No. 297:10-11. [December]

Abbott, R. Tucker. 1985. Seashells ot the W orld, A Guide to

the Better-Known Species. [Revised Etiition]. \ Golden

Nature Guide. Golden Press, New York. 160 pp.

Wagner. Robert J. L. and R. Tucker Abbott [Editors]. 1985.

Standard Catalogue of Shells. [Supplement 3 World Size

Records]. .American Malacologists, Inc., Melbourne, Flor-

ida. 28 pp.

Abbott, R. Tucker and S. Peter Dance. 1985. Compendium

of Seashells. Japanese Edition [translated by T. Habe and

T. Okutani], Heibonsha Ltd, Tokyo, pp. 1-445.

[Abbott, R. Tucker]. 1986. The Nautilus, Its 100th Anniver-

sarv and \oluine 100. The Nautilus 100(11:1-8. [January

31]'

RTA. 1986. Tom Pullev and the Train. The Nautilus 100(1):

8. [January 31]

Abbott, R. T 1986. Rev lew ol "Wells, F. R. 1986. Seashells

of Western Australia. Western .Australian Museum, Perth.

207 pp. "The Nautilus 100(3): 113. [July 30, 1986] [reprint-

ed New York Shell Club Notes 300:18]

Abbott, R. T. 1986. Review of "Reid, D. G. 1986. The

Littorinid Molluscs of mangrove Forests in the Indo-

Pacific Region. British Museum (Naturl History), London.

228 pp." The Nautilus 100(31:113. [July 30, 1986]

Abbott, R. T. 1986. Review of "Gate, J. M. and S. Raskin.

1986. It's Easy to Say Crepidula! Prettv Pretty Press,

Santa Monica, 155 pp." The Nautilus 100(3); 113. [July 30,

1986] [reprinted New York Shell Club Notes 300:19]

Abbott, R. T. 1986. Review of "Just, H. and M. Edmunds

1985. iVor(/i Atlantic Sudibranchs (Mollusca) Seen by

Hcnning Lemche" The Nautilus 100(3);1 14. [July 30, 1986]

.Abbott, R. Tucker 1986. Review of "Rios, E. C. 1986. Sea-

shells of Brazil. Museu Oceanographico. Rio Grande. Bra-

zil, 328 pp., 102 pis." Hawaiian Shell News 34(9):8. [Sep-

tember] [reprinted New York Shell Club Notes 300:16]

.Abbott, R. Tucker. 1986. Cantharus multangulus new sub-

species grandanus from Northwest Florida (Buccinidae).

The Nautilus 100(4);120-121. [October 31J

Abbott, R Tucker. 1986. To Joe Rosewater— From R. Tucker

Abbott. The Nautilus 100(4):152. [October 31]

Abbott, R. Tucker[lllustrationsb> George F, Sandstrom]. 1986.A

Guide to Field Identification, Seashells of North .America.

Golden Press, New York. 280 pp. [Revised Edition]

.Abbott, R. Tucker and S. Peter Dance. 1986. Compendium

of Seashells. American Malacologists, Inc., Melbourne,

Florida, x -I- 411 pp. [Third Edition]

Abbott, R. Tucker. 1987. The Spell ot the Shell. Golden Years,

Melbourne, FL 9(3):18-19. [January]

Abbott. R Tucker. 1987. New hsln home for mollusks. The

Capsule 19(4)1, 4. [Januarv] [Reprinted in COA Bulletin,

March 1987]

Abbott, R. Tucker. 1987. Concerning Olala lactea (Miiller,

1774). Festivus 19(2);12. [February 12]

Abbott, R. Tucker. 1987. That Old Food Chain at Work. The

Capsule 19(5):5. [March]

Abbott, R. Tucker. 1987. Win moon-struck snails face east.

Conchologists of America Bulletin 15(1 ):4. [March]

■Abbott, R. Tucker. 1987. New fish> home for mollusks. Con-

chologists of .America Bulletin 15(1 ):8. [March]

Abbott. R. Tucker. 1987. Letter to a Beginner— With Apol-

ogies, [letter]. American Conchologist 15(2):17. [June]

Houbrick, Richard S., Robert Robertson, and R. Tucker .Abbott

1987. .Anatomv and Systematic Position of Fastigiella

carinata Reeve (Cerithiidae; Prosobranchia). The Nautilus

101(3);101-I10. [July 31]

Abbott, R. Tucker. 1987. Marine Genus New to Atlantic Dis-

covered. The Capsule 20(1):1. [Julv]

.Abbott. R- Tucker. 1987. Letter [concerning editors of mala-

cological journals]. Shells and Sea Life 19(7 ):4. [September]

Abbott, R. Tucker. 1987. Review of "Petuch, E. J. 1987.

Neic Caribbean Molluscan Faunas. CERF, Charlottes-

ville, Virginia. 154 -I- 4 pages, 28 -I- 1 plates." The Festivus

19(10):98-99. [October 8] [Reprinted in New York Shell

Club Notes 305:6-7]

A[bbott], R. T. 1987. New LL S. Shell Stamp Laicovered.

.American Conchologist 15(4 ):2. [December]

Page 72

THE NAUTILUS, Vol. 110, No. 2

Abbott, R. T. 1987. Review of "Petucli, E. J. 1987. New

Caribbean Molhiscan Faunas. CERF, C^liarlottesviile, VA.

154 + 4 pages, 28 + 1 plates." Conchologists of ,\merica

Bulletin 15(4):19. [December]

Abbott, R.T. 19S7 Review of 'Winner, B. E. 1987. A Field

Guide to Molluscan Spawn, volume 1. E.B.M. North Palm

Beach, PL. 139 pages. Conchologists of .\merica Bulletin

15i4);19, [December]

Abbott, R. Tucker [Editor], 1987. Register of .American Ma-

lacologists : .A National Register ot Professional and Am-

ateur Malacologists and Private Shell Collectors. [Second

Edition], American Malacologists, Inc., Melbourne, Flor-

ida. i-.\iii, 1-168.

Abbott, Robert Tucker. 1987. Harvard Class of 1942, Records

of the Class, 1987. Harvard .\lunini Association, Cam-

bridge, MX. p. 1,

Abbott. R, Tucker. 1987, Pronouncing the Scientific Names

of Scashelh of S'orlh America. (.Cassette), .American Ma-

lacologists, Inc. Melbourne. FL

Abbott, R. Tucker. 1987. E.xploring Collectible Shells. 90

minutes of commentary by the author. (Cassette). Amer-

ican Malacologists, Inc. Melbourne, FL

Abbott, R. Tucker. 1988. Dear COA Reader: [Editorial],

American Conchologist 16(1 ):2. [March]

Abbott, R. Tucker. 1988. Emerald Green Snails. American

Conchologist 16(1 ):8. [March]

Abbott, R. Tucker. 1988. Green Snails. American Conchol-

ogist 16(1 ):9. [March]

Abbott, R. Tucker. 1988. Teskey Collection Enhances Florida

Research. New York Shell Club Notes 306:6. [March] [re-

prmted, see below]

Abbott, R. Tucker, 1988, Teske\ Collection Enhances Florida

Research. Hawaiian Shell News 36(4 ):.5. [.April]

Abbott, R. Tucker, 1988, 100Ct)nchological \ears Ago, Amer-

ican Conchologist 16(2):10, [June]

Wagner, R J L, and R, Tucker Abbott. 1988. World Size

Records. American Conchologist 16(2): 17. [June]

Court, I. and R Tucker Abbott. 1988. William Daniel Bledsoe

(1920-1987). American Conchologist 16(3):16. [Septem-

ber]

Abbott, R, Tucker, 1988, What's New in Molluscan Research':'

American Conchologist 16(3):17. [September]

Abbott, R. Tucker, 1988. Ecphora Fever. American Con-

chologist 16(3):20. [September]

Abbott, R. Tucker. 1988. What's New in Molluscan Research?

.American Conchologist 16(4):19. [December]

Abbott, R. Tucker. 1988. Limerick Bargain. American Con-

chologist 16(4):19. [December]

Sti\, Hugh, Marguerite Sti\ and R. Tucker Abbott [photographs

by H. Landshoff]. 1988. The shell: h\e hundred million

years of inspired design. Harry .\ Abrams, Inc New York

188 plates -I- unpaginated text

Abbott, R. Tucker. 1989. Review of "Long islanil Shell Club

1988. Seashclls of Long Island, Long Island Shell Club,

Manhasset, NY. 209 pp." The Capsule 21(4):.\. [January]

Abbott, R. Tucker, 1989, The Spectacular Scallop. .American

Conchologist I7(l);4-5. [March]

Abbott, R. Tucker. 1989. Renate Wittig Skinner, New York

Shell Club Notes 31 1:4-.5 [June]

Abbott, R Tucker. 1989. What's Neu in Molluscan Research'?

.American Conchologist 17(2):9 [September]

.Abbott, R Tucker. 1989. Museum with a Molluscan Mission.

.American Conchologist 17(2): 17. [September]

Abbott, R. Tucker. 1989. Renate Wittig Skiimer 1922-1989.

American Conchologist 17(2):24. [September]

Abbott, R. Tucker 1989, Shells Portland House, New York,

160 pp [December; German Edition, 1990; French Edi-

tion, 1990]

Abbott, R, Tucker. 1989. Compendium of Landshells, .A Color

Guide to More than 2,000 of the World's Terrestrial Shells.

American Malacologists, Inc , Melbourne, Florida, i-viii +

1-240. [December]

.Abbott, R. Tucker. 1989. Snorkeling Sea Shells. .American

Conchologist 17(3):17. [December]

Abbott, R, Tucker, 1989. Corinne Elizabeth Edwards (1905-

1989). American Conchologist 17(3):19, [December]

Wagner, Robert J. L, and R. Tucker Abbott [Editors], 1990.

Standard Catalogue of Shells. [Supplement 4, World Size

Records]. American Malacologists, Inc., Melbourne, Flor-

ida. 80 pp. [April 27]

Abbott, R. Tucker. 1990. Seashells Photo Postcards: 24 full-

color ready-to-mail cards. Dover Publications, Inc, Min-

eola, NY. 16 pp. [Ma\]

Abbott, R. Tucker. 1990. Correcting the Book. "Shells".

American Conchologist 18(2 ):25, [June]

Abbott, R. Tucker. 1990. Yo\o Clams discovered m East

Florida. New York Shell Club Notes No. 315: 10 [June]

[Reprinted from Capsule 22(6) .Astronaut Trail Shell Club].

Abbott, R. Tucker. 1990. The Pocket Guide to Seashells of

the Northern Hemisphere. Dragons' World Ltd., Limps-

field and London, 176 pp. [.August]

Abbott, R, Tucker, 1990. Comments on the proposed con-

servation of Limax fimbnatus Martsn, 1784 and Merita

hebraea Martyn, 1786 (currently Placostylus fimbriatus

and Natica hebraea, MoUusca, Gastropoda). Bulletin of

Zoological Nomenclature 47(3):202. [September 28]

Abbott, R. Tucker. 1990. .American .Nature Guides. Seashells.

Gallerv Books, New York. 1-176, [September] [.American

Edition of Seashells of the Northern Hemisphere]

Abbott, R, Tucker, 1990. Sowerb\ Reborn New York Shell

Club Notes 317:8-9. [December]

Abbott, R. Tucker. 1990. Zebra Mussel Watcli Invoked in

Florida. New York Shell Club Notes 317: 11. [December]

Abbott, R, Tucker and S. Peter Dance. 1990, Compendium

of Seashells. Crawford House Press, Bathurst, Australia,

411 pp, [.Australian Printing]

.Abbott, R, Tucker, 1990. Muscheln unil Meeresschnecken.

Karl Muller, \'erlag. Erlangen, 160 pp, [German Edition

of Shells, 1989]

Abbott, R, Tucker. 1990. Coquillages. Editions Soline, Cour-

bevoie. 160 pp. [French Edition of Shells, 1989],

Abbott, R. Tucker. 1991. The Life Span of Mollusks. Amer-

ican t;onchologist I8(4):25. [Januarv 23]

Abbott, R, Tucker. 1991. Shellmg in Southeast Asia. Of Sea

and Shore 13(4):148-154. [February 4]

Abbott, R. Tucker. 1991. Seashells of the Northern Hemi-

sphere. Gallery Books, New York. 1-191. [March]

Abbott, R. Tucker. 1991 Review of "Sabelli, B.. R Gianuzzi

Savelli and B. Bedulli [eds ]. 1990 .\nnotaled Check-list

of Mediterranean Marine Mollusks, Vol. I. Soc. Italiana

Malacologia, Bologna, ItaK. 348 pp." American Conchol-

ogist 19(1):23. [March]

Abbott, R. Tucker, 1991, Review of "Rios, Cedar Garcia

1990, Los (^)uintones de Puerto Rico. University de Hu-

macao, Humacao, PR, 48 pp," .American Conchologist

19(1):23. [March]

Abbott, R. Tucker, 1991, Seashells ol Southeast Asia, Tvnron

Press, Thornhill, Dummfriesshire 1-145, plates 1-52 [Ma\ ]

Lipe, Robert E, and R Tucker Abbott 1991 Living Shells

M. G. Harasewych, 1997

Page

of the Caribbean and the Florida Ke\s, American Mala-

cologists. Inc. Melbourne, Florida. HO pp. [May 27]

Abbott, R. Tucker. 1991. Mollusk Research ni Bermuda

American Conchologist 19(3):7. [September]

Abbott, R. Tucker. 1991. Review of "Dance, S. Peter and

David Heppell. 1991. Shells — Classical Natural History

Prints. Studio Editions, London, 128 pp.' American Con-

chologist 19(3): 11. [September]

Abbott. R. Tucker. 1991. Those Puzzling Olive Shells. Amer-

ican Conchologist 19(3):23. [September]

Abbott, R. Tucker. 1991. Review of 'Dance, S. P. and D.

Heppell. 1991. Shells — Classical Natural History Prints.

Studio Editions, London 128 pp." New York Shell Club

Notes 323:8. [June]

Abbott, R. Tucker. 1991. The Shell Collector Threat. Sea

Frontiers 37(5): 14-19. [October]

Abbott, R. Tucker. 1991. Was Christopher Columbus a shell

collector'-' Shell Museum Musmgs. Island Reporter [Sani-

bel-Captiva] 19(43):5B. [November 1]

Abbott, R. Tucker. 1991. The first lady of Sanibel shelling.

Shell Museum Musings. Island Reporter [Sanibel-Captua]

19(45);5B. [November 8]

Abbott, R. Tucker. 1991. What's in a name'.-' Shell Museum

Musings. Island Reporter [Sanibel-Capti\ a] 19(46):5B. [No-

vember 15]

Abbott, R. Tucker. 1991. Golden shells. Shell Museum Mus-

ings. Island Reporter [Sanibel-Captiva] 19(47):4B, 5B. [No-

vember 22]

Abbott, R. Tucker, I99I. Your shell collection - dump it or

save it':* Shell Museum Musings. Island Reporter [Sanibel-

Captiva] 19(48):4B. 5B. [November 29]

Abbott, R Tucker. 1991. Commensals, parasites, and pearls.

Shell Museum Musings. Island Reporter [Sanibel-Captiva]

19(49):4B. [December 6]

Abbott, R. Tucker. 1991, Edgar Allan Poe and other Pizzle

Editions. Shell Museum Musmgs. Island Reporter (Sanibel-

Captiva] 19(50):5B, 14B. [December 13]

Abbott, R. Tucker. 1991. Honey, I shrunk the shells! Shell

Museum Musmgs Island Reporter [Sanibel-Captiva] 19(51):

4B, 5B. [December 20]

Abbott, R. Tucker. 1991. With lose Irom Russia. Shell Mu-

seum Musings. Island Reporter [Sanibel-Captiva] 19(52):

4B, 5B. [December 27]

Abbott. R. Tucker. 1991. New C;0.\ Grants. American Con-

chologist 19(4):17. [December]

Abbott, R. Tucker. 1991. Let's go shellingl Or shall we'r" New

York Shell Club Notes 321.14-17. [December] [Reprinted

from "Sea F'rontiers " October 1991 under the title "The

Shell Collector Threat"]

Abbott, R. Tucker. 1992. Showers of Shells. Shell Museum

Musings. Island Reporter [Sanibel-Captiva] 20(1):4B. [Jan-

uary 3]

Abbott, R. Tucker. 1992, Dangerous mollusks and zebras.

Shell Museum Musings. Island Reporter [Sanibel-C^aptisa]

20(2):4B. [January 10]

Abbott, R. Tucker 1992. Museum acquisitions. Shell Museum

Musings. Island Reporter [Sanibel-Captiva] 20(3):8B-9B.

[January 17]

Abbott, R. tucker. 1992. The Sanibel Loosa Beads. Shell

Museum Musings. Island Reporter [Sanibel-Captiva] 20(4):

13B. [January 24]

Abbott, R. Tucker. 1992. Bhie-Ribbon conchologists. Shell

Museum Musings. Island Reporter [Sanibel-Captiva] 20(5):

6B-7B. [Januarx 31]

Abbott, R. Tucker. 1992. Shells kill birds Shell Museum Mus-

ings. Island Reporter [Sanibel-Captiva] 20(6):11B. [Feb-

ruary 7]

Abbott, R. Tucker. 1992. The Spectacular Scallop Shell Mu-

seum Musings. Island Reporter [Sanibel-Captiva] 20(7):

6B-7B. [February 14]

Abbott, R. Tucker. 1992. The day an oil spill hits Sanibel.

Shell Museum Musings. Island Reporter [Sanibel-Captiva]

20(8): 17B. [February 21]

Abbott, R. Tucker. 1992. Dogs and shell collectors. Shell Mu-

seum Musings. Island Reporter [Sanibel-Captiva] 20(9):

17B. [February 28]

Abbott, R. Tucker. 1992. It's show and shell time! Shell Mu-

seum Musings. Island Reporter [Sanibel-Captiva] 20(10)

Shell Fair Guide: 8-10. [March 6]

Abbott, R. Tucker. 1992. Fallen Angels, part one. Shell Mu-

seum Musings. Island Reporter [Sanibel-Captiva] 20(11):

lOB-llB [March 13]

Abbott, R. Tucker. 1992. Fallen Angels, part two. Shell Mu-

seum Musings. Island Reporter [Sanibel-Captiva] 20(12):

9B. [March 20]

Abbott, R. Tucker, 1992. Earl\ Florida shell collectors. Shell

Museum Musings. Island Reporter [Sanibel-Captiva] 20(13):

16B-17B. [March 27]

Abbott, R, Tucker. 1992. Review of "Leal, Jose H 1991.

Marine Prosobranch Gastropods from Oceanic Islands

off Brazil. V. B. S. Publishers, Holland 418 pp." New

Y'ork Shell club Notes 322:5 [March]. The identical review

appeared in Hawaiian Shell News 40(3):8 [March] and

American Conchologist 20(2):20. [June]

Abbott, R. Tucker. 1992. The miracle of Granular Ark Mus-

sels. Shell Museum Musings. Island Reporter [Sanibel-Cap-

tiva] 20(14):5B. [April 3]

Abbott, R. Tucker. 1992. [Untitled]. Shell Museum Musings.

Island Reporter[Sanibel-Captiva]20(15):4B-5B [April 10]

.Abbott, R. Tucker. 1992. Museum mission — inspiring kids.

Shell Museum Musings. Island Reporter [Sanibel-Captiva]

20(16):5B. [April 17]

.Abbott, R. Tucker. 1992. Giant African snails — again! Shell

Museum Musings. Island Reporter [Sanibel-Captiva] 20(17):

5B-6B. [April 24]

.Abbott, R. Tucker. 1992. Thomas Sa\ — America s first con-

chologist. Shell Museum Musings. Island Reporter [Sanibel-

Captiva] 20(18):6B. [May 1]

Abbott, R, Tucker. 1992. The shell people - the Calusa In-

dians. Shell Museum Musings. Island Reporter [Sanibel-

Captiva] 20(19):5B, 9B. [May 8]

Abbott, R. Tucker. 1992. [Untitled]. Shell Museum Musings.

Island Reporter [Sanibel-Captiva] 20(20):4B. [May 15]

.Abbott, R, Tucker 1992. Pearl buttons from river mussels.

Shell Museum Musings. Islaiul Reporter [Sanibel-Captiva]

20(21):10B. [May 22]

Abbott, R. Tucker, 1992. Pioneer in nature's garden. Shell

Museum Musings, Island Reporter [Sanibel-Captiva] 20(22):

2B. [May 29]

Abbott, R. Tucker. 1992. Review of the new journal RL TH-

ENICA. Hawaiian Shell News 40(5):7 [Ma\ ]

Abbott, R. Tucker. 1992. Review of "Rice, T. 1992. Catalog

of Dealers' Prices, including Freshwater and Terrestrial

Shells 1 1th Edition. Of Sea and Shore, Port Gamble, WA.

112 -H 44 pp." Hawaiian Shell News 40(5):7 [May]

Abbott, R. Tucker. 1992. Shelling laws and your license. Shell

Museum Musings. Island Reporter [Sanibel-Captiva] 20(23):

2B, 7B. [June 5]

Abbott, R. Tucker. 1992. Museums aren't attics Shell Mu-

Page 74

THE NAUTILUS, Vol. 110, No. 2

seum Musings, Island UepDrtiT [banibei-tJaptiva] 20(24):

3B, 5B. [June 12]

.■\l)l)ott, R. Tucker. 1992, Biodiversity — buzzword or boon-

doggle? Shell Museum Musings. Island Keporter [Sanibel-

Captiva] 20(25):2B-3B. [June 19]

Abbott, R. Tucker, 1992, VMien the eyes have it. Shell Mu-

seum Musings, Island Reporter [Sanibel-Captiva] 20(26):

2B-.3B, [June 26]

Abbott, R, Tucker, 1992. COA Grant Helps Scallop Study.

American Conchologist 20(2): 19. [June]

Abbott, R. Tucker. 1992, Shell \Uiseum Musings. American

Conchologist 20(2 ):22. [June]

Abbott, R. Tucker. 1992. RLTHENICA — a new journal.

New York Shell Club Notes 323:8, [June]

Abbott, R, Tucker. 1992. Scallop Aquaculture. Shell Mu-

seum Musings, Island Reporter [Sanibel-Claptiva] 20(27):

lOB, [JuK 3]

Abbott, R, Tucker, 1992, Supreme Court and shells. Island

Reporter [Sanibel-Captiva] 20(28):2B, 5B, [July 10]

Abbott, R. Tucker. 1992. Dreams do come true. Shell Museum

Musings. Island Reporter [Sanibel-Captiva] 21(28):2B-3B.

[July 17]

Abbott, R, Tucker. 1992. Endangered Florida species. Shell

Museum Musings, Island Reporter [Sanibel-Captiva] 21(29):

5B. 9B, [July 24]

Abbott, R. Tucker, 1992, Calilornia I, O, Us and Pismo Clams.

Island Reporter [Sanibel-Captiva] 21(30):4B-.5B. [July 31]

Abbott, R. Tucker, 1992. Those sex-crazed squids. Island Re-

porter [Sanibel-Captiva] 21(32):2B, 4B. [August 7]

Abbott, R. Tucker. 1992, Honey, I blew up the shells! Island

Reporter [Sanibel-Captiva] 21(33):3B. [August 14]

Abbott. R. Tucker, 1992, The making ol a Florida shell book.

Island Reporter [Sanibel-Captiva] 21(34):2B, .5B, [August

21]

Abbott, R, Tucker, 1992, Ghosts ot Clornell's conchological

past. Island Reporter [Sanibel-Captiva] 21 (35):.5B, 6B. [.-Vu-

gust 28]

Abbott, R. Tucker. 1992. Subtropical red tides; Sanibel shells.

Island Reporter [Sanibel-Captiva] 21(.36):2B, [September 4]

Abbott, R, Tucker. 1992. Psychiatr> , shells and Sanibel Island

Reporter [Sanibel-Captiva] 21 (.37 ):3B-4B, [.September 11]

Abbott, R, Tucker, 1992. The making ol a museum exhibit.

Island Reporter [Sanibel-Captiva] 2) (38):2B-3B. [Septem-

ber 18]

Abbott, R. Tucker, 1992. The Sacred Chank. Island Reporter

[Sanibel-Captiva] 21(39):2B-3B. [September 25]

Abbott, R. Tucker. 1992 Review ol "Winner, Beatrice B.

1992. A Field Guide to MoUuacun Spawn. Volume II.,

E. B. M. Publishing, North Palm Beach, FL94 pp." Amer-

ican Conchologist 20(3):20, [September]

Abbott, R, Tucker, 1992, Is the Shell Museum relevant '' Island

Reporter [Sanibel-Captiva] 21(40):2B-3B, [October 2]

Abbott, R. Tucker. 1992. Museum traveling exhibits. Island

Reporter [Sanibel-Captiva] 21(41):2B-3B, [October 9]

Abbott, R Tucker 1992. Atlanta's new hall of shells. Island

Reporter [Sanibel-Captiva] 21(43);2B-3B. [October 23]

Abbott, R, Tucker, 1992. Spondvlus Medicine— good and bad

American Conchologist 20(4): 1 8 [December]

Abbott, R. Tucker, 1992, Dreams do come true. The Junonia

December 19927-9

Abbott, R, T, 1993, Robert J, L, Wagner (190.5-1992). Amer-

ican Conchologist 21(I):18, [March]

Abbott, R. Tucker. 1993. The makings ol a shell show. The

56th Annual Shell Fair (iuide [Supplement to the Island

Reporter, Sanibel] pp. 1.3-15, 22, 24. [March]

.■\bbott, K, Tucker, 1993, "\\ orld Size Records Continues.

American Conchologist 21(3):5. [September]

Abbott, R. Tucker 1993, Review of "Winner, Beatrice B.

1993, Life Styles of the Seashells. E, B. M, Publishing,

North Palm Beach, EL 61 pp, " .-\mericaii Conchologist

21(4):18, [December]

Abbott, R. Tucker. 1993. Seashells ol Great Britain and Eu-

rope. Junior Nature Guide. Dragon's World, Ltd., London,

80 pp, [Edited b\' T, Rowland-Entwistle]

Abbott, R. Tucker, 1993. Kingdom of theSeashell, the color! ul

story of shell collecting and living mollusks. [revised edi-

tion]. American Malacologists, Inc. Melbourne, Florida 256

PP

Abbott, R, Tucker, 1993, Robert J, L, Wagner (190.5-1992).

New York Shell Club Notes .326:16, [March]

Abbott, R, Tucker, 1994, Phalium (Semica.ssis) vector, a New

Deep- Water Species from the Central Indian Ocean, The

Nautilus 107(3):94-96. [February 2]

.Abbott, R. Tucker. 1994. You can fool some of the people. . .

Island Reporter [Sanibel-Captiva] 23(16):10A. [April 22]

Abbott, R. Tucker, 1994. Review of ■'Abbot(t), R.Tucker,

1993 Seashells of North America. Dragon's World, Lon-

don , 81 pp. American Conchologist 22(1 ):18, [June]

Abbott, R, Tucker. 1994. Review of "Ingoglia, G, 1994.

Sea.ihells. Science Close-Up Series. Western Publishing

CO., Racine, WI, 24 pp.. .American Conchologist 22(1):

19, [June]

Abbott, R. Tucker. 1994, Fareu ell to a Shelling Friend. [Obit-

uary— Mary Kline]. Hawaiian Shell News 42(9):9. [Sep-

tember]

Abbott, R. Tucker. 1994. The best beaches for shell collectors.

Travel & Leisure 24(12):76, 80, [December]

Hess, D F,, R T, Abbott, J, Hamann. K Meyer, S Millen. T

Gosliner, N. Sefton and Roger T, Hanlon, 1994 9, Ma-

rine Molluscs of the Cayman Islands. In: Brunt, M, .\. and

J. E. Davies (eds. ) The Cayman Islands: Natural History

and Biogeography Kluwer .Academic Publishers, Nether-

lands, pp, 139-189,

Abbott, R, Tucker and Perc> A. Morris. 1995. A Field Guide

to Shells, .Atlantic and Gulf Coasts and the West Indies.

Fourth Edition, Houghton Mifflin Compain. New York

350 pp., 74 pis, [.April]

Abbott, R. Tucker, 1995, My Favorite Seashells, Boys Life,

June 1995:24-27, [June]

Abbott, R, Tucker. 1995. A Guide to Shelling, Siiiuu Day

Guide 3(1 ):28-30,32, 34.

Abbott, R. T, 1995. Bailey-Matthews Shell Museum, Cultural

Jewel of Sanibel, Sunny Day Guide 3(1):.33,

Abbott, R. Tucker, 1996, Some Children's Shell Books— A

Review. Of Sea and Shore 19(3):150, [Fall— published

posthumously]

,At the time of hi.s death, R. Tucker .Abliott was working

on a number of books. Among these was our collaboration

on the third edition of American Seashells, on wiiicli he

w ill be a co-author.

BOOKS EDITED BY R. TUCKER ABBOTT

.Abbott, R, Tucker, M K, Jacobsciri. and M G Teskc\ (Editors).

1955. How to Collect Shells (A Symposium). .American

Malacological Union, Bulfalo, New York, [iv] -t- 75 + [vii]

pp

M. G. Harasewych, 1997

Page 75

Abbott, R. Tucker, M. K^ Jacobson, and M. C. Teske\ (Editors)-

1961. How to Collect Shells. (A Symposium). Second Edi-

tion. American Malacological I'ninn, Marinette, Wisconsin

iv + 92 + [i\ ] pp.

.Abbott, R. Tucker, M. K, Jacobson, and M. C:. Teske\ (Editors).

1966. How to Collect Shells. {.\ S\niposiuni). Third Edi-

tion. American Malacological L nion, Marinette, Wisconsin

iv -I- 101 -1- [v] pp.

Abbott, R. Tucker, M. K. Jacobson, and M. C. Teskey (Editors).

1974. How to Collect Shells. (A Symposium), Fourth Edi-

tion. American Malacological Union, Wrightsville Beach,

North Carolina [vi] + 107 pp.

Hasting, Louise Burrell and Margaret Crockett Teske\ . 1979,

[Edited b\ R. Tucker Abbott], Indexes to The Nautilus:

Geographical {\'ols. 1-90) and Scientific Names (X'ols, 61-

90). .-Kmerican Malacologists, Inc. Melbourne, Florida. i\

-I- 238 pp.

Sutty, Lesley. 1986. [Edited b\ R. Tucker Abbott]. Seashell

Treasures of the Caribbean. Dutton, New York. 128 pp.

Bratcher, T. and W. Cernohorsk). 1987. [Edited b> R. Tucker

Abbott]. Living Terebras ot the World : .\ Monograph of

the Living Terebridae of the \\ orld .American Malacol-

ogists, Melbourne, Florida. 240 pp.

Naught, K, C, 1989, [.\bbott, R. Tucker and Kenneth J. Boss,

Editors], .\ Classification of Living Mollusca, .American

Malacologists, Inc, Melbourne, Florida, i-,\ii + 1-195,

ACKNOWLEDGMENTS

I am most grateful to Cecelia W, .Abbott for allowing

access to Tucker's personal files and scrapbooks and for

supplying many of the photographs as well as biograph-

ical information, Jose H. Leal and Edith Chippeau.x

brought to m\ attention some of Tucker's notes and card

files that were found at the Bailev -Matthews Shell Mu-

seum. Ellen Strong kindly pro\ ided detailed information

on his Master s Thesis and Doctoral Dissertation. I thank

Richard E. Petit for comments on tiie manuscript, ad-

dition of citations, and information on editions and sup-

plements. Alan l\. Kabat brought to my attention several

Abbott ta.\a. Melbourne R. Carriker kindly made avail-

able a draft of his "In Memoriuni manuscript. Paula

M. Mikkelsen and Robert Robertson generously provided

photographs of Tucker. Special thanks are due Cecelia

Abbott, Kenneth J. Boss, Melbourne R. Carriker, Russell

H. Jensen, Jose H. Leal, Paula M. Mikkelsen, and f^obert

Robertson for their comments on a draft of this manu-

script.

LITERATURE CITED

.Anonymous, 1971, Publications of the I'nited States National

Museum (1947-1970). Bulletin 298 (final volume of series).

Smithsonian Institution Press, Washington, DC. 77pp.

Bieler, R. and R. E. Petit. 1996. Additional notes on nomina

first introduced by Tetsuaki Kira in Coloured Illustrations

of the Shells of Japan. Malacologia 38(l-2);33-34.

Coan, E. V. and M. G. Harasewvch, 1993. Publication Dates

of The Nautilus. The Nautilus 106(4):174-180.

Lyons, W. G. 1988. A Review of Caribbean Acanthochiton-

idae (Mollusca: Polyplacophora) with descriptions of si.x

new species of Acanthochitona Gray, 1821. American

Malacological Bulletin 6(1):79-114.

THE NAUTILUS 110(2): 76, 1997

Page 76

110(2)

PetitiUa, New Name for Petitella Wise, 1996, a Preoccupied Name

(Mollusca: Gastropoda: Pyramidellidae)

John B. Wise

Houston Museum of Natural History

Oiif Hermann Circle Drive

Houston TX 77030-1799 USA

In a recent study of pli\logeneti(.' relationships within

the famiK PyramideUidae, I proposed the generic name

Petitella \\'ise, 1996, witiiiii the new subfamily Sayel-

hnae. The editors of Zoological Record have since brought

to my attention that Petitella is preoccupied, having

previously been apphed to a genus of fish (Gery & Bou-

tiere, 1964). A replacement name is therefore proposed.

PetitiUa. new name for Petitella Wise, 1995:495, not

Petitella Gery & Boutiere, 1964:474.

Type species, by original designation of Petitella Wise,

1996, Sayella crosseana Dall, 1885, Recent, Western At-

lantic.

The new name honors Richard E. Petit for his many

invaluable contributions to malacology, as did the name

it replaces.

1 thank the eilitors of Zoological Record lor bringing

this to m\ attention.

LITERATURE CITED

Dall, W. H. 1885. Notes on some Floridian land and fresh-

water shells with a revision of the .'\uriculacea of the east-

ern United States. Proceedings of the L'nited States Na-

tional Museum 8:225-289, pis. 17-18.

Gery, J., and A. Boutiere. 1964. Petitella georgiae gen. et.

sp. nov. (Pisces CypriniformesCharacoidei). X'ieet Milieu,

Supplement 17:47:3-484. 6 figs.

Wise, J. B. 1996. Morpholog) and pin logenetic relationships

of certain pyramidellid ta\a (Heterobranchia). Malaco-

logia 37(2):443-511.

INSTRUCTIONS TO AUTHORS

THE NAUTILUS publishes papers on all aspects of the

biology' and svstematics of mollusks. Manuscripts de-

scribing original, unpublished research as well as review

articles will be considered. Brief articles, not exceeding

1000 words, will be published as notes and do not re-

(juire an abstract. Notices of meetings and other items

of interest to malacologists will appear in a news and

notices section.

Manuscripts: Each original manuscript and accompa-

nying illustrations should be submitted in triplicate. Text

must be t\ped on one side of 8V2 X 11 inch white paper,

double spaced throughout (including literature cited, ta-

bles and figure captions), with at least 1 inch of margin

on all sides. All pages must be numbered consecutivelv.

If printed on a word processor, the right margin should

be ragged rather than justified. Authors should follow

the reconunendations of the Council of Biologi/ Editors

Sti/lc ManiKii which is available from the Council of

BiologN' Editors, Inc., 9650 Rockville Pike, Bethesda,

MD 20814, U.S.A. The first mention of a scientific name

in tlie te.\t should be accompanied by the tiLXonomic au-

thority, including year. Latin names and words to be

printed in italics must be underlined; leave other indi-

cations to the editor. Metric and Celsius units are to lie

used.

The sequence of sections should be: title page, ab-

stract page, introduction, materials and methods, results,

discussion, acknowledgments, literature cited, tables, fig-

ure captions, figures. The title page should include the

title, author's name(s) and address(es). The abstract page

should contain the title and abstract, which should sum-

marize in 250 words or less the scope, main results and

conclusions of the paper. The abstract may be followed

by a maximum of 8 key words. All references cited in

the text must appear in the literature cited section and

vice versa. In the literature cited section, all authors

nmst be fully identified and listed alphabetically. Follow

a recent issue of THE NAUTILUS for bibliographic

style, noting that journal titles must be unabbreviated.

Information on plates and figures should be cited only

if not included in the pagination. Tables nmst be num-

bered and each placed on a separate sheet. A brief leg-

end must accompany each table. Captions for each

group of illustrations should be tvped on a separate sheet

and include a key to all lettered labeling appearing in

that group of illustrations.

All line drawings must be in black, high quality ink,

clearly detailed and completely labeled. Photographs

must be on glossy, high contrast paper. All figures are to

be consecutively numbered (figs. 1, 2, 3, ... , NOT figs,

la, lb, Ic, . . . NOR plate 1, fig. 1 . • .). Illustrations

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THE NAUTILUS

Voluini' no. \'iiiiihcr 3

Julij IS, 1997

ISSN 0028-1344

A quarterly devoted

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TH EfcNAUTI LUS

CONTENTS

\\>litntc HI). Xitmhcr 3

juhj IS, 1997

ISSN '002S-1344

Claudia Julia del Ri'o

(>\'n()/.()if Bioijeotiraphic Histon' ot the Eiintlit'iinal (hmius

Rclroldpcs. New Cieiius (SuhtainiK" Tapctiiiae) Iniiii

Sdutlii'in Soutli America and Antarctica 77

Iphittts roheiisi (Epitoniidae: Nvsticllinae). A New Species

of Deep-Sea Gastropod from the Gnlf oi Mexico 94

N'erification of the Specific Status of tlie Eutlangered

Anthony's River Snail. Atlicdniin (iiilliniii/i. Using Alloz\ine

Electroplioresis 97

Inilnced Metamorphosis of Fresliwater Mussel Cilochidia

on Nonliost Fisli 102

Bruno Sabelli

Marco Ta\ iani

Robert T. Dillon, Jr.

Steven A. Alilstedt

Sheila G. Kirk

James B. Lavzer

THE NAUTILUS 110(3):77-93, 1997

Page 77

Cenozoic Biogeographic History of the Eurythermal Genus

Retrotapes, New Genus (Subfamily Tapetinae) from

Southern South America and Antarctica

Claudia Julia del Rio

Centre de Investigaciones en Recursos

Geologicos

Ramirez de Velazco 847

(1414) Buenos Aires ARGENTINA

and

Facultad de Ciencias Exactas y

Naturales

L'niversidad de Buenos Aires

Ciudad Universitaria

Pabellon 2

Nunez (1428) ARGENTINA

ABSTRACT

Retrotapes. new genus, comprises a group of Neoaustral bi-

valves that appeared in the southern circumpolar regions b\

the Eocene and ha\e since been confined to the high latitudes

of the southern hemisphere. Its presence in the Tertiar\ strata

of Antarctica and southern South America reflects an active

fauna! interchange between both regions during the Eocene.

Retrotapes is here proposed to include to those Recent and

Tertiarv representatives of the Subfamily Tapetinae (Family

Veneridae) from southern South .America and .Antarctica that

had been previous!) placed in Venus Linne. 1758, Marcia H.

and .•\. .Adams, 1857, Eurhomalea Cossmann,1920, Samarangia

Da!!, 1902, and Kalelijsia Romer, 1857. The Argentine Tertiar>

species Retrotapes r^infasiensis, new species, R. fuegoensis.

new species, R. striatolaniellata (Ihering, 1897) and fi. scutata

(Ihering, 1907) are here described and illustrated. The Recent

Argentine Venus exalbida Chemnitz, 1795 and Venus lenlic-

ularis Sowerby, 1835, the Antarctic V. antarctica Sharman and

Newton, 1894 and \', newtoni Wilckens, 1911 (Eocene-early

Oligocene', La Meseta Formation), and the Neogene Chilean

species V. navidadis Philippi, 1887 and \'. colchaguensis Phi-

lippi, 1887 are also included in this new genus.

Key words. Tapetinae, Retrotapes, new genus, biogeograpin,

Neoaustral, Tertiary, .Argentina, Chile, .Antarctica.

INTRODUCTION

The Subfamii\ Tapetinae (FamiK Veneridae) shows a

moderatel) high degree of endeniism in Recent as well

as in Tertiary faunas. The known geographic distribution

of most living tapetines is mainly restricted to the south-

ern hemisphere. This subfamiK is particular!)' abundant

in the southern Indo-Pacific region, where it is repre-

sented by more than sevent) species. Most of the Indo-

Pacific genera, among them Katelysia Romer, 1857,

Marcia H. & A. Adams, 1857, Granicorium Hedley,

1906, Hemitapes Romer, 1864, Notirus Finlay, 1928,

Notopaphia Oli\er, 1923, Eumarcia Iredale, 1924, Pa-

phirus Finia), 1927, and Gomphinella Marwick, 1927,

are restricted to New Zealand and Australia. Others,

including Gonjp/nna Morch, 1853, Venerupsis Lamarck,

1818, Ruditapes Chiamenti, 1900, Tapes Mergele von

Miihlfeld, 1811, and Paphia Roding, 1798, are also found

beyond those regions.

In contrast. Recent tapetines are poorly represented

in the littoral zones along both coasts of the American

continents. The subfamily is known from only twelve

living North American species distributed among the

genera Riulitapca, Liocyma Dall, 1870, Psephidia Dall,

1902 and Inn Schmidt, 1818, as well as five South Amer-

ican taxa assigned to the genera Eurhomalea Cossmann,

1920 and Retrotapes. new genus.

The fossil record in the .Americas reveals that Tape-

tinae were more abundant in the Tertiary than in the

Recent fauna. Two diverse and different assemblages

appeared during the Tertiary, one restricted to North

America, the another to austral latitudes. The northern

assemblage comprises the endemic genera Liocyma, Cy-

clorisma Dall, 1902, Psephidia and Sii}onia Stephenson,

1952, as well as taxa with european affinities such as

Mercimonia Dall, 1902, Flaventia Jukes-Browne, 1908,

Paraesa Case) , 1952 , Legumen Conrad, 1858 and Tex-

tiveniis Cossmann, 1886. The less diverse austral assem-

blage, characterized by Eumarcia, Katelysia, Atamarcia

Marwick, 1927 and Retrotapes. appeared during the ear-

ly Tertiary in the southernmost region of South America

and Antarctica.

Page 78

THE XALTILLS. \ ol 110, No. 3

Figure I. Stratigr.ipliie disliibutiuii ot Rftrutapes, new genus,

in South America and Antarctica.

The species that are here inckidetl in Rctrotapcs occur

in the Tertiary rocks of Chile, Antarctica and Argentina,

and are at present distributed in the httoral zones along

both coasts of South .America (F'igure 1), where thev are

represented by R. lenticularis (Sovverby, 1835) and R.

exalinda (Chemnitz, 1795).

This paper includes the systematic description of the

Tertiary .Argentine representatives of Rctrotapcs: R. niu-

fasiensis new species (Puerto Madr\ n Formation, middle

Miocene), R. jucgocnsis new species (Carmen Silva For-

mation, late Oligocene-early Miocene), R. striatolarnel-

lata (Ihering, 1897) (Monte Leon Formation, late Oh-

gocene-eariy Miocene) and El Chacay F'ormation (late

Eocene) and R. scutala (Iliering, 1907) (San Julian For-

mation, late Eocene). Venus antarctica Sharman and

Newton, 1894 and V^ newtoni Wilckens, 1911 from the

Eocene-early Oligocene? La Meseta Formation (Antarc-

A FOSSIL LOCALITIES

1.

2

3

4.

5.

Punto Norte

Puerto Pirdmide

Cerro Prismdfico

Fondeodero Nlnfos

Canadon El Lobo

Monte Entroda

Yegua Quemada

Cerro Puntudo

9. Cerro Castillo

10. Eo Lq Fedenco

Figure 2. Geographic distribution ot fossil localities referred

to in this paper.

tica), and the Chilean species V. navidadis Phiiippi, 1887

(early Miocene), V. colchaguensis Phiiippi, 1887 (early

Miocene-Pliocene) and V. lenticularis (Pliocene-Holo-

cene), along with the Recent V. exalbida, are also placed

in Retrotapes.

MATERIAL AND METHODS

The fossil material reported here came from the thick,

marine, sedimentar\, Tertiar\ sequence exposed along

the eastern coast of Patagonia from San Antonio Oeste

(Rio Negro Province), southward to Isla Grande of Tierra

del Fuego, and at Cerro Puntudo (Lago Cardiel), a lo-

caiit\ situated in the western region of Patagonia (Santa

Cruz Province) (Figure 2) Lithostratigraphic units \ ield-

ing representatives of Retrotapes are the Puerto Madryn

Formation (Haller, 1978), the Monte Leon Formation

(Bertels, 1970), the Carmen Silva Formation (Codignotto

& Malumian. 1981), the San Julian Formation (Bertels,

Claudia Julia del Rio. 1997

Page 79

1970) and the El Chacav Formation (Chiesa & Camacho,

1984).

The Puerto Madr\n Formation, exposed at Peninsula

Valdes (Chubut Pro\ince). is believed to represent one

of the youngest Tertiary marine units recognized in Pa-

tagonia, having been reported b\ del Rio (1988, 1992)

as being of middle Miocene age. This formation consists

of 150 meter thick, whitish cinerites and Nellowish sand-

stones alternating with highly fossiliferous calcareous

sandstones and muddy or sandy shelly beds. Retrotapes

is abundant at the basal ochreous shelly sandstones that

are e.xposed at Cerro Prismatico (horizon N 2, del Rio.

1992). where it is associated with "Clilainys" actinodes

(Sowerby, 1846), Purpiirocardia Icoucnsis del Rio, 1986,

Glycymerita magna del Rio, 1992, Aequipecten para-

nensis (d'Orbign\ , 1842), Dosinia meridionalis (Ihering,

1897) as well as with the gastropod Valdcsia valdcsiensis

del Rio. 1985, Rclrotapes is less common in the gray,

massive, fine sandstones that comprise the base of the

section at Punta Norte (horizon PN 2, del Rio, 1992),

where articulated specimens have been found in life

positions along with Anadara (Rasia) lirata (Philippi,

1893), Glycymeris longiorijormis del Rio, 1992, Lucinisa

sp., "Cyclocardia nortensis del Rio, 1986, Dosinia mer-

idionalis (Ihering, 1897), Dosinia cuspidata del Rio, 1994,

Ameghinomya meridionalis (Sowerby, 1846) and

A. argentina (Ihering, 1897). Retrotapes also occurs in

the strata placed at the top of the sequence exposed at

Punta Ninfas (horizon F 9, del Rio, 1992) and is associated

v\ith Glycymerita magna del Rio, 1992, Amusium paris

del Rio, 1992, and Aequipecten paranensis (d'Orbignv,

1842).

Retrotapes has also been collected from exposures of

the late Oligocene Monte Leon Formation that crop out

at the mouth of the Santa Cruz Ri\er (Santa Cruz Prov-

ince). Here it is localK abundant in the highK fossilif-

erous, coquinoid sandstones situated 80 meters above sea

level at Monte Entrada and in the sandy lenses present

at the sea cliffs at Las Cue\as. Fossil material is well

preserved at both localities where disarticulated valves

are associated with a rich bivalve fauna characterized

by Cucullaea alta (Sowerby, 1846), Neilo ornata (Sow-

erby, 1846), Pecten proximus Ihering, 1897, Venericar-

dia inaequalis Philippi, 1887, Limopsis insolita Sowerby,

1846. Australocallista iheringii (Cossmann,lS98), Do-

sinia meridionalis Ihering, 1897, Perna quadrisulcata

Ihering, 1897 and Phacoides critcialis Ihering,1907.

Specimens of Retrotapes found at the Carmen Silva

Formation (late Oligocene-earK Miocene) occur in peb-

bl\ shelly sandstones at the top of the sequence that crops

out at Cerro Castillo and Estancia La Federica. Material

is abundant and well preserxed. with most of the spec-

imens consisting of articulated \aKes.

Material coming from the San Julian Formation was

collected by C. Ameghino from sedimentary marine out-

crops at Caiiadon El Lobo (= Canadon Tournouer), close

to Punta Casamayor. The age of the San Julian Formation

is still debated, ranging from late Eocene to early Oli-

gocene (Naiiez, 1988; Camacho, 1984, 1995), as recently

documented b> del Rio (1995). Only one fairK- well-

preserved valve of Retrotapes is known from the up-

permost, highly-fossiliferous, orange coquinoid sand-

stones that alternate with green or yellowish gra\ silstones

and sandstones exposed at Cafiadon El Lobo.

Retrotapes is poorly represented in the El Chacay

Formation, exposed at Cerro Puntudo (Lago Cardiel).

Chiesa and Camacho (1994) placed this unit in the late

Eocene based on biostratigraphic correlations with the

San Julian Formation.

Specimens described in this paper are housed in the

Museo Argentino de Ciencias Naturales "Bernardino Ri-

vadavia (MACK), Buenos Aires, Argentina, in the Cen-

tro de Investigaciones en Recursos Geologicos (CIRGEO-

PI), Buenos Aires, Argentina, Facultad de Ciencias Ex-

actas y Naturales of the Universidad de Buenos Aires

(CPBA), Buenos Aires, .Argentina, Direccion Nacional

Servicio Geologico (DNSG), Buenos Aires, .Argentina and

in the Department of Earth and .\tmospheric Science,

Purdue University (PU), Indiana, USA.

SYSTEMATIC PALEONTOLOGY

Phylum MOLLUSC A Linne, 1758

Class BIVALVIA Linne, 1758

Subclass HETERODONTA Neumavr, 1884

Order VENEROIDA H. & A. Adams, 1856

Superfamilv N'ENEROIDEA Rafinesque, 1815

Familv \'ENERIDAE Rafinesque, 1815

Subfamilv TAPETIXAE H. & A. Adams, 1857

Diagnosis: Members of this subfamily are characterized

b\ having smooth inner ventral margins and three car-

dinal teeth in each valve, with at least two of them

grooved or bifid. Most Tapetinae have exterior surfaces

with commarginal lines and/or grooves, while others

have polished surfaces or very fine radial threads. Though

no attention has been paid to the presence of a lunule

or the arrangement of cardinal teeth, these characters,

along with the general outline of the cardinal platform,

are of taxonomic importance at the generic level. When

present, and viewed dorsally, the lunule may be concave

or convex, and is limited b> a groove and/or ridge that

is distinguished from the remaining surface of the shell

b>' its different ornamentation, or b> the presence of a

groo\e.

Hinge characters ha\e pro\en to be the most useful

morphological feature for distinguishing among genera.

Cardinal teeth ma\' be placed on a thin, slender, cardinal

platform with a straight ventral margin, or situated on

a short, broad, strongly arcuate platform. While some

genera are characterized b\ basing teeth that diverge

from a point situated below the beaks, others ha\e non-

divergent teeth (Figure 3). Divergent teeth are charac-

terized by a strongly forward-inclined anterior tooth, a

vertical median tooth, and a backward-inclined posterior

tooth. Genera with non-divergent teeth may have three

cardinal teeth sloping backwards (posterior tooth nearly

Page 80

THE NAUTILUS, Vol 110, No. 3

pas

divergent teeth

non-divergent teeth

Figure 3. Hinge teeth t\ pes in Tapetinae. aas, anterior atlductor

muscle scar; lu, lunule; pas, posterior adductor muscle scar;

l,3a,3b, cardinal teeth.

horizontal) or have the median and posterior teeth in-

clining backwards, with the anterior tooth being vertical.

Genus Retrotapes, new genus

Patagornalea del i^io, 1991 ;93, nomen nudum.

Diagnosis: Shell large to medium sized, highly variable

in outline. Escutcheon facing opposite valve, usually wid-

er, smoother on left valve than on right in adults. Lunule

large to medium sized, deeply impressed, strongly dif-

ferentiated from remaining shell surface, highK to mod-

erately concave, nearly vertical, with longitudinal me-

dian sulcus, bounded b\ deep lunular groove. Hinge

heavy, short, arcuate behind teeth, with three long car-

dinal teeth. Right and left posterior teeth nearly hori-

zontal in some species, left and right anterior teeth nearly

vertical to strongly sloping backw ards, nearK paralell to

posterior teeth; left median tooth, right median and pos-

terior teeth bifid or grooved.

Type species: Retrotapes ninfasiensis , new species.

Type locality: Valdes Peninsula, Chubut Province, Ar-

gentina, Puerto Madryn Formation (Middle Miocene).

Stratigraphic range: Eocene to Holocene, southern South

America and .Antarctica.

Etymology: retro- L. backwards. Referring to the incli-

nation of non-divergent cardinal teeth towards the pos-

terior region of the shell.

Remarks: Recent as well as fossil species belonging to

Retrotapes ha\e pre\ iousK- been placed in Venus Linne,

1758, (Philippi, 1887; Ihering, 1897), in Marcia (Dall,

1902; Ihering. 1907; Riveros & Gonzalez, 1950), in Sa-

marangia Dall, 1902 (Jukes-Browne, 1908; C^arcelles,

1944; Carcelles & Williamson, 1951; Castellanos, 1970

Rios, 1975), in Eurhonialea (Keen, 1954; Soot-Ryen, 1959

Raniorino, 1968; Fisher-Piette & Vukadinovic, 1977

Malumian el al.. 1978; Zinsmeister, 1984; Stilwell & Zins-

meister, 1992) and in Katelifsia (Dall, 1902; Jukes-Browne,

1908; Riveros & Gonzalez', 1950).

The taxonomic placement ot the Holocene South

American species Venus exalbida (Figures 22-23, 41)

and Venus lenticularis (Figures 19-21), here assigned to

Retrotapes. has long been debated, and the phylogenet-

ical relationships of these species within the family Ve-

neridae were misunderstood. Dall (1902) incorrectly in-

LEFT VALVES

46 2b 2o

RIGHT VALVES

UM

3o 1 3b

/ ropes //ferafus {Lmnoeui]

2- Katelysia scalarina (Lomorck)

/vAA fvAJX

3- Eufhomalea rufo (Lamarck)

tyJXh

aiu\

4 - Retrotapes ninfosiensis n. sp.

/u/nn

KS\J\

5- Retrotopes striotolomelloto (Ihering)

fvnsK /uui

6- Retrotapes fuegoensis n. sp.

7- Retrotapes lenticularis (Sower by)

/UHA

AAn

8- Retrotopes exalbida (Chemnitz)

Figure 4-. Schematic cross-sections of hinge teeth. Tapes lit-

eratiis (Linne). M.-\CN 266-1, Indian Ocean, Recent 1,2.4 x);

Kalclysia scalarina (Lamarck). PU 863-5, N'ictoria, Australia.

Recent (2.4 x); Eurhumalea rufa (Lamarck). PL 165, Chile,

Recent. (1.2 x); Retrotapes ninfasier^sis. new species. CPBA

15.110, Fondeadero Ninfas, middle Miocene, Puerto Madr\n

Formation, Argentina (1.2 x); Retrotapes striatolamellata

(Ihering). MACN 2639, Yegua Quemada, late Oligocene, Mon-

te Leon Formation, Argentina (1.2x); Retrotapes fuegoensis,

new species. PU 355-12, Cerro Castillo, late Oligocene-early

Miocene, Carmen Silva Formation, Argentina (1.2 x); Retro-

tapes lenticularis (Sowerby), PL' 165, Chile. Recent (1.2 x);

Retrotapes exalbida (Chemnitz) MACN 21 172, .Argentina. Re-

cent (1.2 X).

terpreted Fischer's (1887) statement about V. exalbida

being the type species of the genus Marcia, considereil

Sainarangia (t\ pe species Venus quadrangularis Adams

& Reeve. 1850) to be a section of Marcia. antl placed V.

lenticularis in Sauiarangia. He thought V. exalbida, V.

lenticularis and V. quadrangularis to be part of a closeK

related group of species related to Marcia Jukes-Browne

Claudia Julia del Rio. 1997

Page 81

Figures 5-9. Tapes litcratus (Linne). MACX 266-1. Indian Ocean, Recent. 5. Exterior view of left valve (IX). 6. Interior view

of right valve (IX). Enlargements of 7. right and 8. left hinges (2X). 9. Dorsal view of an articulated specimen (IX). Figures 10-

14. Eiirhomalea nifa (Lamarck). PU 165, Chile, Recent. 10. Exterior and 11. interior views of right valve (IX) 12. Dorsal view

of an articulated specimen (IX). Enlargement of 13. left and 14. right hinges (2X).

Page 82

THE NAUTILUS, Vol. 110, No. 3

(1908) designated Venus pinguis Chemnitz, 1782 (= V.

opima Gmelin, 1791 ) as the t\ pe species of Marcia, and

excluded V. exalbida from this genus.

Venus quadrangularis (see Fischer-Piette & Vukadi-

novic, 1977:22, figs.207-211, for illustrations of the spe-

cies) cannot be related to either \'. exalbida or \'. Icn-

ticiilaris because of the presence in V. quadrangularis

of a smooth exterior surface, a pustular left anterior lat-

eral tooth and an entire pallial line, characters that al-

lowed Keen (1969) to segregate Samarangia in the Sub-

family Samarangiinae. Other diagnostic features, in-

cluding the hinge and lunular characteristics of V, ex-

albida, V. lenticularis, as well as those of their Tertiary

ancestors, require the erection of the new genus Retro-

tapes. It is placed in the SubfamiK Tapetinae on the

basis of the presence of shells with a smooth inner ventral

margin, moderate to very deep pallial sinus, three car-

dinal teeth in each valve with two of them deeply grooved

or bifid, and the absense of lateral teeth. Morphological

features that distinguish Retrotapes from the remaining

genera of the subfamiK are the presence of non-diver-

gent cardinal teeth and a well defined, concave lunule,

bounded by a deep lunular groove and ridge.

Eurhomalea, represented by its t\pe species E. rufa

(Lamarck, 1818) (Figures 4, 10-14) and by £. salinensis

Ramorino, 1968 (see Ramorino. 1968:218, pl.3, fig.2, pi. 9,

figs.2-3 for illustrations of the type species), is distributed

along the Peruvian and Chilean coasts and has recentlv

been placed in the Subfamily Chioninae (Fischer-Piette

& Vukadinovic, 1977). However, the presence of a smooth

inner ventral margin in Eurhomalea indicates that it is

more accurately assigned to the Subfamily Tapetinae. It

clearly diff^ers from Retrotapes in lacking both lunule

and escutcheon, in having a straight dorsal margin, a

more antero-posteriorly elongate shell, and a narrower

hinge plate than Retrotapes. Moreover, the cardinal teeth

are markedly smaller than those of Retrotapes and quite

divergent from a point situated below the beaks. The

anterior cardinal teeth of both valves in Eurhomalea

slope forward. The left posterior and right anterior car-

dinal teeth are much lower than in Retrotapes. and the

left middle, and right posterior and middle teeth are

only shallowly grooved.

Katelysia (type species: Venus scalarina Lamarck,

1818) (Figures 4, 25-28), a middle Miocene-Holocene

genus confined to New Zealand and Australia, differs in

having a diff^erent outline, a narrower lunule and es-

cutcheon, a shallower pallial sinus, and a shorter cardinal

teeth than Retrotapes.

Marcia opima. the Indo-Pacific type species of Mar-

cia, was illu.slrated by Abbott and Dance (1986:363). The

genus differs from Retrotapes in having a smooth exterior

shell surface, a high umbonal area, a weakly defined

escutcheon, and a nearly smooth and lightK impressed

lunule that lacks a lunular groove. These genera may

also be separated becau.se Marcia has lower, narrower

and more widely divergent teeth than Retrotapes with

the anterior cardinal teeth inclining forward.

Eurnarcia (t\pe species Venus fumigata Sowerby,

1853, illustrated by Lamprell & Whitehead, 1992, pl.74,

fig.589; Abbott & Dance, 1986:363) a common genus in

New Zealand and Australia, is easih separated from Re-

trotapes on the basis of its oval and smooth shells, a lunule

that is not impressed and that is bounded by a weak line,

divergent teeth with left anterior and middle cardinal

teeth deepK grooved and equal in size, and in having a

left posterior tooth that is fused to nymph.

The Tertiary New Zealand Atamarcia (type species

Eurnarcia sulcifera Marwick, 1927, figs. 200, 203, 205,

earl\ Oligocene-late Pliocene) differs from Retrotapes

in being sculptured with commarginal grooves, in having

a fairly impressed lunule, with a shallo\\- lunular groove,

divergent teeth with curved posterior teeth, and with

the left anterior and middle teeth being of equal width.

The monot\pic, lower Pliocene Opimarcia Marwick,

1948, (type species: O. healyi Marwick, 1948, pi. 5, figs. 1,

2, 4) is distinguished from Retrotapes in having oval,

inflated shells, very prominent beaks, a poorly defined

lunule not bounded by a lunular groove, a fairly well-

developed escutcheon, fine irregular commarginal ridges

on the posterior and anterior ends of valves, a shorter

pallial sinus than Retrotapes, and divergent teeth with

a strongly curved right posterior tooth and a triangular,

deeply grooved left anterior tooth.

Tapes (type species: Venus literatus Linne, 1758)

(Figures 4-9) has shells that are more antero-posteriorK

elongated than those of Retrotapes, and have a straight,

horizontal dorsal margin, acuminate anterior margin, a

rounded pallial sinus, straight or slightK arcuate hinge

margin and a grooved left anterior cardinal tooth.

The oldest presentK known record of Retrotapes dates

to the late Eocene. This genus occurs in the San Julian

Formation and the El Chaca> Formation (.\rgentina),

where is represented b\ R. scutata and R. striatolamel-

lata, and in the lower and middle sections of the La

Meseta Formation (late Eocene-Oligocene":*, ,\ntarctica)

where R. aniarctica and R. newtoni have been found.

Retrotapes ninjasiensis, new species

Figures 15-18, 40

Marcia strialolamellata Frengueiii, 1926, not M striatola-

mellata Iliering, 1897.

Diagnosis: Shell thick, ovate to subrectangular. Lunule

deeply concave, nearly vertical, strongly inclined toward

opposite vaKe. Cardinal teeth straight, high, long, thick.

Median and anterior teeth sloping backwards, posterior

cardinal teeth horizontal. Pallial sinus triangular, mod-

erateK short.

Description: Shell thick, large, o\ate to subrectangular,

weakly convex, ranging from longer than high to nearly

equidimensional. Umbones small, at anterior quarter of

length. Dorsal margin moderately to strongly convex,

posterior margin truncated to weakK convex, ventral

margin nearly straight to w eakly convex, anterior margin

convex. Lunule concave, with longitudinal median lu-

nular groove, nearK' vertical, strongK inclined toward

opposite val\ e, bounded b\ deep groove and w ell marked

Claudia Julia del Rio, 1997

Page 83

.<.:/

^^ -

>iJv

\

-.V

#

15

J

S:^ "' - \^

;^

1 7

18

X i

/A**^

19

20

Figures 15-18. Rctrotapes ninfasiensis. new species. Puerto Madryn Formation, middle Miocene. 15. Interior and 18. exterior

views of right vaKe of holotype, CPBA 13.573, Cerro Prismatico. 16. Interior view of left valve of parat>pe, CPBA 15.110,

Fondeadero Ninfas. 17. Interior view of left valve of paratype, CPBA 15 090, Punta Norte Figure§ 19-21. Retrotapes lenticularis

(Sowerby), PU 165, Cfiile, Recent. 19. Exterior, 20. interior and 21. dorsal views of left valve. .\11 figures 1 x.

Page 84

THE NAUTILUS, Vol. 110, No. 3

22

23

/y

24

26

30

25

8W3'^

27

31

Figures 22-24. Retrotapes exallnda (Chemnitz) MACN 21.172, San Matias Gulf. Argentina, Recent Ail figures 1 x. 22. Interior

and 2.1. exterior views of left valve. 24. View of right hinge. Figures 2.5-28. Kateltjxia scalarina (Lamarck), PU 865-5. Victoria,

.Xustralia, Recent. All figures 2X. 25. Interior, 26. exterior, and 2K. dorsal views of left valve, 27. Interior view of right valve.

Claudia Julia del Rio, 1997

Page 85

ridge. Escutcheon on left \alve wide, long, inclining to-

ward right valve and only slightly sculptured with fine

commarginal ridges; escutcheon on right valve poorly

differentiated and ornamentated as remaining shell sur-

face. Hinge short, narrow , stronglv arcuate behind teeth,

with three high, thick cardinal teeth per vaKe (.Hgure

4). Anterior and median teeth strongly inclined back-

wards, posterior teeth nearK' horizontal. Right valve with

deepK groo\ed posterior tooth separated from nymph

h\ socket, followed b\ ridge, median cardinal bifid with

lamellar posterior section, anterior tooth thick, equal in

height to median tooth; socket for left median tooth w ith

radial lamella. Left hinge with strong, smooth, anterior

tooth, median tooth deepK' grooved, with both parts equal

in size, lower than anterior tooth. Posterior tooth lamellar,

slightl) arched, with superimposed ridge on posterior

face, separated from nymph by shallow groove. Anterior

adductor muscle scar o\ al, more deepK impressed, larger

than posterior muscle scar; anterior pedal retractor mus-

cle scar small, deeply excavated, placed above adductor.

Pallial sinus relativeK short, triangular, with horizontal

dorsal margin, straight, ascending ventral margin; ape.\

pointed. Exterior ornamented with wideK-spaced, thin,

lamellar commarginal ridges, much more numerous near

ventral margin; interspaces sculptured with ver> fine

radial threads.

Material examined: Holotype, CPBA 13.573, right valve,

Cerro Primatico, heiglit 66.6 mm, length 76.2 mm. Para-

tvpes, CPBA 15.090, left valve, Punta Norte, height 77.1

mm, length 81. 9 mm, CPBA 15.110, left valve, Fon-

deadero Ninfas height: 60,0 mm, length: 66,0 mm; 19

left valves, 16 right valves, one articulated specimen,

Punta Norte (PN 2): CPBA 15.087-15.089, CPBA 13.287-

13.288, Fondeadero Ninfas (F 9): CPBA 12.345, CPBA

15.1 1 0, Cerro Prismatico (N 2): CPBA 12.501, CPBA

11.646-11.650, CPBA 12.343. CPBA 12.502, CPBA

13.572, 13.574-13.575 (del Rio collection)

Stratigraphic and geographic distribution: Puerto Mad-

ryn Formation, middle Miocene. Valdes Peninsula, Chu-

but Province.

Remarks: Specimens of Retroiapes ninfasiensis have

erroneously been attributed to Mania striatolamellata

(Frenguelli. 1926). a closely related, late Oligocene spe-

cies that occurs in the Monte Leon Formation of Santa

Cruz Province. Retroiapes striatolamellata (Figures 33,

37-39, 42-47) ma>' be differentiated b\ its longer, more

convex shells, straighter dorsal margin, a lunule that is

more concave and not inclined toward the opposite valve,

less prominent teeth, a left anterior cardinal tooth that

is less inclined backw ards, being slightK arched forwards

in most specimens and higher or equal in height than

the median cardinal (Figure 4), and a pallial sinus that

is tongue-like and longer than in R. ninjasiensis. Retro-

tapes antarctica (Sharman & Newton. 1894. fig. 3). from

the La Meseta Formation (Antarctica. Eocene- early Oli-

gocene?), differs in having a trigonalK suboval outline

with the dorsal margin strongK sloping backwards and

rounding to the posterior margin, by the presence of a

shallow ly conca\e, narrow er lunule not inclined toward

the opposite valve, and a narrower, shorter pallial sinus

than R. ninjasiensis. Moreover, R. antarctica has nar-

rower teeth than R. ninfasiensis, the right anterior tooth

inclines forwards, the left anterior tooth is grooved and

higher than the median one, and the left median tooth

is deepK grooved, with both sections equal in width.

Retrotapes ninfasiensis is easily separated from R. na-

vidadis (Philippi. 1887. pi. 14. fig. 4. Navidad Formation)

because the Chilean Miocene species has a thinner and

smaller shell with an acuminate anterior margin,

straighter dorsal margin, more prominent beaks and a

narrower lunule that is not inclined to the opposite valve.

Cardinal teeth of R. navidadis are narrower and much

shorter than in R. ninfasiensis. the right anterior cardinal

tooth is vertical, but its anterior face inclines forward,

while the left anterior cardinal tooth is lightly curved

and also slopes forward. Retrotapes lenticularis (Sow-

erby) (Riveros & Gonzalez, 1950:fig.30) (Figure 19-21),

a species distributed along the Chilean literal from Co-

quimbo to N'alparaiso, and also present in the Pleistocene

outcrops in Central Chile (Herm, 1969:pl.l3, figs. 1-4).

has subcircular shells with straight to lightly convex dor-

sal margins, smaller lunules not inclined toward the op-

posite valve, and much narrower and shallow er grooved

cardinal teeth than R. ninjasiensis. The right posterior

cardinal tooth inclines backwards and is separated from

the nymph by a groove, the right anterior tooth varies

from being vertical to slightK inclined forward, and the

left anterior and median teeth are curved. The exterior

surface is covered by fine commarginal ridges, widely

spaced, deep commarginal grooves, and very fine radial

threads.

The Recent R. exalbida (Castellanos, 1970:250, pi. 22,

figs. 4-5) (Figures 22-24, 41) is characterized by subrec-

tangular and more inflated shells, with a shallower lunule,

lower and shorter cardinal teeth with a deeper, grooved,

left median tooth than in fi. ninfasiensis (Figure 4).

Retrotapes striatolamellata (Ihering, 1897)

Figures 33, 37-39. 42-47

Marcia itriatolamcUata Ihering, 1897:253, pi. 7, fig. 44; Ihering,

1907:305,

Mania navidadis Philippi, Ortmann, 1902:141, pl.27, fig.l2.

Marcia ortmanni Ihering, 1907:;504

Figures 29-31. Retrotapes scutata (Ihering). .\\\ figures 2X, Holot\pe M.^CN 429. Canadon El Lobo, San Julian Formation, late

Eocene. 29. Left hinge. 30. exterior and 31. dorsal views of left valve. Figure 32. Retrotapes fuegoensis, new species. 1 x. Exterior

view of left valve of holotype. PL' 356-12. Cerro Castillo. Carmen Silva Formation, late Oligocene-early Miocene.

Page 86

THE NAUTILUS, Vol 110, No. 3

Figure 33. Retrolapes striutolanwllata (Ihering) Interior view of left valve, CPBA 9359, Monte Entrada, Monte Leon Formation,

late Oligocene ( 1 X ) Figures 31— 36. Retrotapcs fiu'goensis. new species. Carmen Silva Formation, late Oligocene-early Miocene. 34.

Kniargement of right hinge, DNSG 16.500. f:stancia La Federica (2X), 35. Interior view of left valve of holotype PL' 356-12,

Cerro Castillo (IX). 36. Dorsal view of an articulated paratype DNSG 16.501, Estancia La Federica (IX). Figures 37-39. Rctrulapes

Claudia Julia del Rio, 1997

Page 87

Diagnosis: Shell thick, o\ate to subrectangular in outline,

longer, more convex than R. ninjasiensis. Lunule strong-

ly concave, not inclined toward opposite valve. Left an-

terior tooth arched (in most specimens), equal in height

to median cardinal. Pallial sinus tongue-shaped, longer

than in R. ninjasiensis.

Description: Shell thick, large, ovate to subrectangular,

moderatelv convex, longer than high. L'mbones small, at

anterior quarter of length. Dorsal and anterior margins

convex, ventral margin slightly convex, posterior margin

slightly convex to subtruncated. Lunule broad, deeply

concave, nearly vertical, not inclined toward opposite

valve, marked b\ deep groove and distinctive ridge; lon-

gitudinal, median, lunular groove impressed. Escutcheon

on left valve better defined than on right, consisting of

wide, flat surface facing opposite valve, sculptured with

very faint commarginal ridges. Escutcheon on right valve

narrower than on left, heavily ornamented like remain-

ing valve surface. Hinge with three narrow cardinal teeth;

right valve with cardinal sloping backwards, lamellar

anterior tooth, median cardinal narrow K and shallowly

grooved, as high as or higher than anterior tooth. Pos-

terior tooth broadly grooved, separated from nymph by

ridge; anterior tooth of left hinge lamellar, arched, ver-

tically or slightly inclined backwards, median tooth deep-

ly grooved, broad, as high as, or higher than anterior

tooth, narrow, straight or arched posterior cardinal tooth.

Muscle adductor scars equal in size, shallowK- impressed.

Pallial sinus deep, tongue-shaped. Commarginal sculp-

ture of thin lamellar ridges, much more closeK spaced

near ventral margin. Interspaces smooth, except for

growth lines.

Material examined: Holotvpe, MACX 437, right valve,

Yegua Quemada, height 67.5 mm, length 79.3 mm; Para-

tvpes, two left valves, Yegua Quemada, MACN 2639,

height 65.0 mm, length (38.0 mm. and MACN 2640,

height 65.0 mm, length 68.0 mm; two left and two right

valves from Monte Entrada, CPBA 9.359, PU429, Las

Cuevas, CPBA 9.39L and from Cerro Puntudo (Lago

Cardiel) CIRGEO-PI 2.513. (Ihering, Ortmann, Medina

and Camacho collections)

Stratigraphic and geographic distribution: Monte Leon

Formation (late Oligocene), from the mouth of the Santa

Cruz River to Yegua Quemada, and El Chacav For-

mation (late Eocene), Cerro Puntudo (Lago Cardiel).

Remarks: Retrotapes striatolamellata comes from out-

crops of the Monte Leon Formation at Monte Entrada,

Las Cuevas and Yegua Quemada (Figure 2). This species

has also been recorded from exposures of the El Chacay

Formation at Cerro Puntudo (Lago Cardiel) (Santa Cruz

Province). The type material of R. striatolamellata had

been collected b\ Carlos Ameghino at Yegua Quemada,

but this fossiliferous localitv has never been recognized

by subsequent authors. Ortmann (1902) placed fi. stria-

tolamellata in synonymv with R. navidadis (Philippi,

1887). Ihering (1907) considered Ortmann's specimens

to represent a new taxon that he named Marcia ortmanni

Ihering, 1907. However, a re-analysis of Ortmann's ma-

terial (1902, plate 27, figure 12) (Figure 47), reveals them

to be voung specimens of R. striatolamellata.

Material coming from Cerro Puntudo (Lago Cardiel)

(Figures 37-39) is limited to a poorly preserved right

valve with an eroded hinge. It is placed in R. striato-

lamellata because of the arrangement of teeth, shape

and size of pallial sinus, characteristics of lunule, es-

cutcheon and exterior commarginal ornamentation. The

onl) difference between this specimen and those coming

from the Monte Leon Formation is in the outline of the

shell. While the Cerro Puntudo specimen has a narrowly

convex posterior margin w ith an ovate-subtriangular out-

line, most representatives of R. striatolamellata from

eastern Patagonia have shells with outlines ranging from

subrectangular to ovate, with a broadly convex or trun-

cated posterior margin (Figures 33, 42-46).

Retrotapes navidadis (Philippi, 1887) is distinguished

from R. striatolamellata in having a thinner and smaller

shell with a straight dorsal margin, narrower and less

concave lunule, shorter and narrower teeth and more

widely spaced and homogeneousK' distributed commar-

ginal ridges than in R. striatolamellata. Retrotapes len-

ticularis has a subcircular shell w ith a narrower and less

concave lunule, lower and more shallowly grooved teeth

than in R, striatulaniellata. a vertical right anterior tooth,

right and left posterior teeth that slope backwards, and

a pallial sinus with an acute apex and concave ventral

margin. Retrotapes striatolamellata may be easily sep-

arated from R. antarctica (Sharman 6i Newton, 1894),

which has a trigonally suboval shell, slightly concave

lunule, narrower teeth, a right anterior tooth that is in-

clined forward, and a shorter and triangular pallial sinus

than R. striatolamellata.

Retrotapes scutata (Ihering, 1907)

Figures 29-31

Marcia scutata Ihering, 1907:303, pi. 11, fig.76.

Diagnosis: Shell, elongate-oval in outline, anterior mar-

gin convex. Lunule narrow, slightK' concave.

Description: Shell ot medium size, elongate-oval in out-

line, weakly convex, longer than high. Dorsal margin

striatolamellata (Ihering), CIRGEO-PI 2513 Cerro Puntudo, El Chacay Formation, late Eocene, All figures IX. 37. Exterior, 38.

interior and 39. dorsal views of right valve. Figure 40. Retrotapes ninjasiensis, new species, CPBA 15.087, Punta Norte, Puerto

Madryn Formation, middle Miocene. Dorsal view of articulated specimen (1 x ). Figure i^X. Retrotapes exalbida (Chemnitz), MACN

21 172, San Matias Gulf. .Argentina, Recent, Dorsal view of an articulated specimen (1 x ).

Page 88

THE NAUTILUS, Vol. 110, No. 3

^^Sa^

%

44

47

45

46

Figures 42-47. Retrotapes strialolamellata (Ihering), Monte Leon l'\irmation, late Oligocene. 42. Interior view ot riglil valve of

paratype, MACN 2639, Yegua Quemada (1 x ). 43. E.xterior, 44. interior, and 46. dorsal views of the holotype, MACN 4.37, Yegua

Quemada (1 x ). 45. Interior view of left valve of parat\ pe, MACN 2640, Yegua Quemada (1 x ). 47. P-429, Monte Entrada. interior

view of left valve of juvenile specimen illustrated by Orlniann (1902:27, fig.l2) (2x),

Claudia Julia del Rio, 1997

Page 89

steeply sloping donwards, rounding to posterior margin;

anterior margin narrowly convex. Lunule narrow, shal-

lowly conca\e, not inclined toward opposite valve. Es-

cutcheon on left \aKe strongly sloping toward right valve.

Left hinge narrow, with lamellar anterior cardinal tooth

sloping backwards, median tooth deeply grooved, pos-

terior tooth lamellar, horizontal, slightly curved. Exterior

v\ith lamellar commarginal ridges. Interior unknown.

Material examined: Holotype, NIACIN 429, left valve,

Canadon El Lobo, height 32,0 mm, length 3i>,0 mm.

(Ihering Collection).

Straligraphie and geographic distribution: San Julian

Formation (late Eocene) from Canadon El Lobo (Santa

Cruz Province).

Remarks: Rctrotapcs scutata is known only- from the

holot\ pe, a left valve whose interior is unknown and the

hinge partially covered with marl. The elongate-oval

outline, narrow, slightly impressed lunule and thin car-

dinal teeth, separate this species from fi. striatolamellata

and R. ninfasieuf^is. Retrofapes scutata and R. newtoni

(Wilckens, 1911: plate 1, figure 16; Zinsmeister, 1984:

figure 9 I, H, K, Eocene-early Oligocene? La Meseta

Formation, Antarctica), share similar features including

outline, ornamentation, characters of the lunule, and ar-

rangement of cardinal teeth. These similarities suggest

that these species are very closely related. A more thor-

ough comparision between R. scutata and R. newtoni is

not possible until more material of R. scutata is collected.

Retrotapes fuegoensis new species

Figure 32. 34-36

Eurhomalca '' cl juenzalidai . Malumian et al., 1978:278. pi. 3,

tig. 4, not Venus fuenzalidae Pliilippi. 1887

Diagnosis: Shell elongate-oval in outline, anterior margin

subtruncated, posterior margin convex. Cardinal teeth

lower, narrower than in both R. striatolamellata and R.

ninfasiensis, right anterior tooth straight, vertical, right

median tooth shallowly grooved. Lunule very narrow,

shallowly concave. Pallial sinus tongue-shaped, much

deeper than in R. striatolamellata.

Description: Shell large to medium sized, weakly convex,

elongate-oval, longer than high. Umbones at 1 /9 of dorsal

length. Dorsal margin slightly convex, rounding to con-

vex posterior margin; anterior margin subtruncated. Lu-

nule very narrow, shallowly depressed to nearly flat, not

inclined toward opposite valve, broader in left valve than

in right valve, with slighth marked median radial sulcus,

bounded b\ moderately deep lunular groove. Escutcheon

narrow, slightlv broader in left valve than in right valve,

with strongly sculptured commarginal ridges on both

valves. Hinge with anterior tooth vertical, median and

posterior teeth inclined backward. Left hinge with strong,

high anterior tooth, median cardinal rectangular, bifid,

with both parts equal in size, as high as anterior tooth.

posterior cardinal lamellar, curved. Right valve with thin

anterior tooth, narrow grooved median tooth, thin, deep-

ly grooved posterior tooth. Exterior with strong, rounded,

commarginal ridges, more crowded toward ventral mar-

gin.

Material examined: Holotype, PU 355-12, left valve,

Cerro Castillo height 62.0 mm, length 73.0 mm (Zins-

meister Collection); Paratvpes, articulated specimen,

DNSG 16.501, height 48.3 mm, length 58.8 mm, right

valve DNSG 16.502, height 32.8 mm, length 27.0 mm;

six articulated specimens, one left valve, two right (Mal-

umian Collection), and one fragment with left hinge

from Estancia La Federica and Cerro Castillo PU 357-

16, PU 355-13, DNSG 16.500, 16.503-16.509 (Zins-

meister and Malumian Collections).

Stratigraphic and geographic range: Carmen Silva For-

mation (late Oligocene-early Miocene), Cerro Castillo

and Estancia La Federica, Isia Grande de Tierra del

Fuego.

Remarks: Retrotapes fuegoensis comes from the up-

permost conglomerated beds of the Carmen Silva For-

mation's exposures at the quarrel of Estancia La Federica

and at Cerro Castillo, where it is associated with a highly

diverse and well preserved molluscan fauna previously

studied by Malumian, Camacho and Gorrofio (1978). The

outline of R. fuegoensis is the most distinctive character

that readily distinguishes this species from its congeners.

Based on examined material, the Chilean Neogene spe-

cies Venus colchaguensis Philippi, 1887 ( = V .fuenzalidai

Philippi, 1887) ma\' be also included in the genus Re-

trotapes. It differs from R. fuegoensis in having an acu-

minate anterior margin, a poorly defined escutcheon, and

commarginal sculpture that consists of wideK and uni-

forniK spaced lamellae. Retrotapes ninfasiensis may be

distinguished from R. fuegoensis by its ovate to subrec-

tangular, posteriorly truncated shell, a more concave and

broader lunule facing the opposite valve, much larger

cardinal teeth, median and anterior teeth that strongly

slope backward, and a triangular and shorter pallial sinus

than in R. fuegoensis. Retrotapes striatolamellata differs

from R. fuegoensis in having a shell that is ovate, more

convex, and more acuminate anteriorly, with a lunule

that is more concave and broader, and teeth that are

larger than in R.fuegoensis. Retrotapes scutata has a

more elongate and ovate shell, with a more concave

lunule than R. fuegoensis. Retrotapes antarctica is char-

acterized by a trigonally suboval shell with umbones

placed more posteriorly, an anterior margin that is not

truncated, and a smaller triangular pallial sinus than that

of R. fuegoensis. as well as a right anterior tooth that

inclines slightly forward. Retrotapes newtoni has an ovate,

smaller shell that is not subtruncated anteriorly. It also

has a more concave lunule, thinner teeth and a shorter

pallial sinus than R. fuegoensis. The Recent species R.

exalbida and R. lenticularis have subrectangular and

subcircular outlines respectively, deeper, concave lunules

and shorter pallial sinuses than R. fuegoensis.

Page 90

THE NAUTILUS. Vol. 110. No. 3

DISCUSSION

Fleming (1963) proposed that Neoaustral faunal ele-

ments originated in the low latitudes of the Pacific mar-

gins during the late Tertiary. However. Zinsmeister

(1982.1984) pointed out that at least some components

of this fauna would have originated during the early

Tertiar\ in high latitudes of circumpolar regions, and

subsequently radiated northward. Antarctic Eocene re-

cords of Aulacomya Morch, 1853, Gaimardia Gould,

1852, Gomphina Morch, 1853 and Eurhomalea Coss-

mann.1920. led Zinsmeister to consider these to be

Neoaustral taxa that originated in the higher austral lat-

itudes much earlier than Fleming suspected.

The transfer to Refrotapes of Antarctic species pre-

viously assigned to Eurhomalea (Zinsmeister. 1984; Stil-

well & Zinsmeister. 1992) limits Eurhomalea to strata

younger than the Pleistocene. Eurhomalea most likely

originated in Chile during the Pleistocene, later reached

the Peruvian littoral fauna, and is now restricted to the

eastern Pacific coast. Ret rot apes, however, is a Neoaus-

tral genus that appeared in Patagonian and Antarctic

regions during the early Tertiary.

According to present data. Retrotapes first appeared

in the southwestern Atlantic Ocean by the Eocene. It

occurs in the late Eocene San Julian Formation and the

El Chacay Formation of Patagonia, as well as in the

Antarctic La Meseta Formation of late Eocene-Oligo-

cene? age. By the early Miocene, Retrotapes expanded

northward into the middle Chilean region, reached

northern Patagonia b\ the middle Miocene, and attained

its northermost distribution (R. lenticularis and R. ex-

albida) during the Holocene (Figure 1). Fleming (1963)

and Zinsmeister (1982. 1984) agreed that cool seawater

temperatures were related to the presence of Neoaustral

taxa. Zinsmeister (1982) stated that the development of

Neoaustral faunal elements in the early Tertiary of Ant-

arctica reflects the cool water temperatures that existed

in that region during the late Eocene. He also considered

the absence of Neoaustral genera in earlier Tertiary de-

posits of South America to indicate the lack of suitable

climatic conditions for the development of this fauna.

The present study records a new. Tertiary and Ho-

locene. Neoaustral genus in southern South America, and

suggests that it has adapted to a wide range of water

temperatures that fluctuated from cool-temperate to

warm. Estimates of Antarctic late Eocene paleotemper-

atures range wideK among authors. Kennett (1977) cal-

culated Eocene sea-surface temperatures around Ant-

arctica to have been 9-12°C higher than today, while

Feldmann and Zinsmeister (1984) indicated that cool

temperate conditions occurred in the area. More re-

cently, Stilwell and Zinsmeister (1992) indicated that,

although cool to warm conditions may be inferred from

marine invertebrate taunas, the molluscan fossils indicate

warm-temperate conditions during the deposition of the

La Meseta P'ormation.

Despite discrepant inlercnccs ol paleotemperatures.

mollusks reveal that Antarctic conditions during the late

Eocene were cooler than those in southern Patagonia

during both the Eocene and Oligocene, as well as those

recorded during the middle Miocene of northern Pata-

gonia. Del Rio (1990, 1994 a,b), inferred warm temper-

atures for northern Patagonia during the middle Miocene

based on the presence of tropical and subtropical genera

as Amusium, Flahellipecten, Chionopsis, Antinioche,

Hexacortnila. Miltha. Egeta, Area. Dosinia s.str. and

Lucinisca. According to del Rio (1990. 1994 a.b), middle

Miocene temperatures in northern Patagonia would have

been similar to those of the tropical Panamic Molluscan

Province and the warmest regions of the Gulf and Ca-

ribbean Molluscan Provinces. Along the Patagonian li-

toral, temperatures decreased steadily, with cooler con-

ditions similar to those occurring today having been es-

tablished by the end of middle Miocene times. Almost

70% of the middle Miocene genera became extinct, in-

cluding all of the above mentioned tropical and sub-

tropical genera. Retrotapes. howe\er. continued to live

in this region adapting to the new climatic conditions.

Its present distribution is restricted to the warm tem-

perate Argentinian Province and to the cool-temperate

Magellanic Province.

Recent advances in our knowledge of Antarctic and

Patagonian molluskan assemblages (Zinsmeister & Ca-

macho, 1980; Zinsmeister. 1976.1981.1984; Camacho &

Zinmeister, 1986; Griffin. 1991; Stilwell & Zinmeister.

1992) show that the ocurrence of Retrotapes in Patagonia

and Antarctica, along with several other genera and sub-

genera, including Periploma (Ae/ga ) Slodkewitsch. 1935.

Pteromijrtea Finla\. 1927. Lahillia Cossmann. 1899.

Crassatella Lamarck. 1799, Au/aconij/a Morch. 1853, the

gastropods Struthiolarella Steinmann and Wilckens. 1908

and Eoscaphella Stilwell and Zinsmeister. 1992. as well

as nuculoids. arcoids (del Rio & Camacho. 1997) and

veneroids that are presently being studied, reinforces the

similarities in paleoclimates of southern South America

and the Antarctic continent during the Paleogene. even

after the final break-up of the Weddellian Province.

ACKNOWLEDGMENTS

1 wish to express my gratitude to G. Parma (Facultad de

Ciencias Exactas y Naturales of the liniversity of Buenos

Aires) for her assistance in the accurate stratigraphic

determination of fossil specimens from the San Jorge

Gulf area and to M. G. Harasewych (National Museum

of Natural Histor\. Washington. DC) and A. Oleinik

(Purdue Lhiiversity. Indiana) who improved the English

text. I am also indebted to W. Zinsmeister (Purdue Lhii-

versity). F. Medina (Centre de Investigaciones en Re-

cursos Geologicos. Buenos .Aires. Argentina) and N. Mal-

umian (Ser\icio (ieologico Nacional. Buenos .Aires. .Ar-

gentina) for allowing me to study their collections. Thanks

also to the curators of the Ortmann and Ihering Colle-

citons housed in Purdue L'niversity and in the Museo

Argentino Argentino de (Ciencias Naturales "Bernardino

Rivadavia respectiveK'. for permitting me access to the

material described in this paper. This research has been

Claudia Julia del Rio, 1997

Page 91

partialK' carried out through a Research Fellowship of

the CONICET (Consejo Nacional de Investigaciones

Cientificas y Tecnologicas) carried out in the Department

of Earth and Atmospheric Sciences of Purdue University.

C. Gambedotti took the photographs and Amalia Gon-

zalez (Centro de Investigaciones en Recursos Geologicos,

CONICET) drafted the figures.

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14.

THE NAUTILUS 110(3):94-96, 1997

Page 94

Iphitus robertsi (Epitoniidae: Nystiellinae), A New Species of

Deep-Sea Gastropod from the Gulf of Mexico

Bruno Sabelli

Dipartimento di Biologia

Evoluzionistica e Sperimentale

Universita d Bologna

Via S. Giacomo 9

1-40126 Bologna

ITALY

Marco Taviani

Istituto di Geologia Marina,

C.N.R.

Via Gobetti 101

1-40129 Bologna

ITALY e-mail; taviani

boigm2.igm.bo.cnr.it

ABSTRACT

Iphitus robertsi. new species, is described on the basis of nine

specimens collected in the northern Gulf of Mexico. This new

species, which differs from its six congeners in having a bicar-

inate shell with weakly canceliate sculpture, represents the first

record of this genus in the Gulf of Mexico. The co-occurance

of this species with deep-sea scleractinian corals suggests that

Iphitus robertsi may, like its New Zealand congener, may be

parasitic on corals.

Key words: Gastropoda, Epitoniidae, new species. Gulf of

Mexico, deep-sea.

INTRODUCTION

A recent survey of deep-sea cold seeps of the continental

slope of the Gulf of Mexico using the research submers-

ible Johnson-Sea-Link I has led to the discovery of a new

species of Epitoniidae from off Louisiana and Texas. The

specimens were collected in a vent-free area at a site

known as Green Canyon (Roberts et al., 1990), Lease

Block 189, during Johnson-Sea-Link I dive 3306. The sea

bottom at a depth of 184 m was composed of mixed

rocky and biogenic (silty-sandy shell debris) substrate.

The description of this new species is based on nine

empty shells in different stages of growth that were re-

covered while sorting three cubic decimeters of biogenic

sand under a dissecting microscope. The species diversity

of the molluscan fraction of this sample was very high

(n=197).

SYSTEMATICS

Family Epitoniidae Berry, 1910

Subfamily Nystiellinae Clench and Turner, 1952

(Jenus Iphitus Jeffreys, 1883

Iphitus robertsi nev\ species

Figures 1-4

Description: Shell small, relatively tragile, trochilorm,

hoiostomatous, umbilicate, with carinate, angular whorls

rapidiv increasing in diameter. Suture subcanalicuiate.

Protoconch I of 1 '/i smoothly rounded w horls. Protoconch

II (larval shell) of 3V2 whorls, strongly sculptured by axial

ribs and weaker spiral cords. Protoconch brownish, axis

inclined by about 10° to teleoconch axis. Teleoconch of

3'/2 whorls, with strong, spiral sculpture of widel\- spaced

carinae and strong spiral cords. Color w hitish, w ith few,

irregular, pale brownish blotches. First half-whorl some-

what rounded, subsequent whorls with two carinae, one

in upper third of w horl, the second suprasutural. Strong

spiral cords, one between suture and shoulder, one or

two between carinae, appear beginning with second

whorl. Finer spiral threads visible on body whorl of larger

specimens. Shell base with 5-6 spiral threads, innermost

bordering deep, wide, infundibuliform umbilicus. Sur-

face of umbilicus sculptured with thin, spiral threads.

Axial sculpture of densel\- spaced threads, producing

rectangular, canceliate pattern at intersections with spi-

ral cords, threads, and carinae. .\perture attached to pre-

ceding whorl along one face of thin, roughly pentagonal

peristome. Only one of nine specimens retained a com-

plete protoconch, the remaining specimens lacked em-

bryonic portion of the protoconch (protoconch I).

Type locality: Green C^anvon, Block 189, Gulf of Mexico

(27°46.48'N, 93°17.74'\V)'in 184 m.

Type Material: Holot\ pe. National Museum of Natural

History, Smithsonian Institution, Washington, DC, USNM

880185, Paratype USNM 888186; 3 paratypes, Museo di

Zoologia, Bologna lWii\ersit\-, MZB 11621 a,b, MZB

11622; 1 paratype Museum National Histoire Naturelle,

Paris; 1 paratype, Houston Museum of Natural Sciences,

HMNH 42544, all from the type localit\ .

Additional Material Kxamined: 1 shell (Figure 2, acci-

dentalK broken), from the t\pe localit>.

Etymology: This species is named after Dr. Harr> Rob-

erts (Louisiana State University), the Chiel Scientist of

the research crui.se and Scientist-in-Charge of scientific

operations during dive JSL-I-3306.

B. Sabelli and M. Taviani, 1997

Page 95

Figures 1-4. Iphitus robertsi, new species. Green Canyon, Block

Holotype, USNM 888185, Scale bar = 3 mm 2. Adult shell, scale bar

of holot\'pe. Scale bar = 3 mm.

189, Gulf f Mexico (27°46,48'N, 93°17.74'W) in 184 m. 1.

3 mm. 3. .Apical and 4. Lateral views of the protoconch

['age 96

THE NAUTILUS, Vol. 110, No. 3

Remarks: Bertolaso and Palazzi (1994) synonymized

Iphitus Jeffreys, 1883 with Stylotrochus G. Seguenza,

1876, which is unavaliable because it is preoccupied by

Stylotrochus Haeclcel. 1862. To date, oni\ six (four extant

and two extinct) species of the genus Iphitus have been

described. The Recent Iphitus tuberatus Jeffreys, 1883,

/. cancellatus Dautzenberg and Fischer, 1896, and /.

tuarshalli (S\ kes, 1925) inhabit the eastern North Atlan-

tic, while /. neozelanicus (Dell, 1956) occurs off New

Zealand. The fossil species Iphitus asperatus (G. Seguenza,

1886) and /. papillosocinctus (G. Seguenza, 1886) are both

from upper Pliocene-lower Pleistocene bathyal deposits

of Sicily (Palazzi & \illari, 1996). All of these species

have been well illustrated (Beu, 1978; Taviani & Sabelli,

1982; Bouchet & Waren, 1986, Bertolaso & Palazzi, 1994,

Palazzi & V'illari, 1996), and show little similarity to the

new species described here. Iphitus rohertsi superficially

resembles 7. marshalli. \\ hich also has strong carinae and

spiral cords, but these differ in number and arrangement

in the two species. Iphitus marshalli also differs in lack-

ing cancellate sculpture. Iphitus cancellatus. the only

species to be reported from the western Atlantic, differs

from /. rohertsi in having a much coarser reticulate sculp-

ture and a rounded aperture lacking carinae. The fossil

legacy of Iphitus is quite scant. The oldest record (as

Iphitus sp.) is from an upper Miocene coral-assemblage

in southeastern Spain (Barrier et ai, 1991). Iphitus as-

peratus (G. Seguenza, 1886) and /. papillosocinctus (G.

Seguenza, 1886) are known from Plio-Pleistocene de-

posits in the Messina region of SiciK' ( Bertolaso & Palazzi,

1994. Palazzi & Villari, 1996), while Iphitus tuberatus

occurs in glacial-Pleistocene deposits of southern Italy

(Rindone & N'azzana, 1989) and submerged last glacial

(Pontinian) tanathocoenoses of the Sicilan Channel (Tav-

iani & Sabelli, 1982).

Flabitat: .\vailable ecological data indicates that species

of Iphitus are parasitic on deep-sea scleractinian corals.

Beu (1978) discovered /. neozelanicus living embedded

in the calices of Goniocorella dunwsa (Alcock, 1902).

While there are no direct observations on any of the

Atlantic species, a relationship with deep-sea scleracti-

nian corals has been suggested by Taviani and Sabelli

(1982). Bouchet and Waren (1986) noted that I. tuber-

atus occurs with Lophelia and that an association is prob-

able. Bertolaso and Palazzi (1994) recorded one shell of

/. asperatus still associated with a piece of Lophelia from

the glacial Pleistocene of Calabria (southern Italy). A

similar habitat is inferred for /. robertsi. As supported

by visual observations (H. Roberts, personal communi-

cation, September 1992) and sample content, dead and

living scleractinian corals (e.g. Madracis sp., Caryophyl-

lia sp., Coenosmilia arbuscula Pourtales, 1874, and Bal-

anophyllia sp.: H. Zibrowius, in litt. 29 October 1993)

were abundant at the sampling site. Among the possible

hosts, the branching scleractinian Madracis. the com-

monest coral, is the most likeK candidate, although no

traces attributable to a secondary ectoparasite (Beu, 1978)

were visible in our material.

ACKNOWLEDGMENTS

We are very grateful to Dr. Harr\ Roberts for inviting

one of us (M.T.) on the research cruise, and for obtaining

the samples on site. The Captain, crew and scientists

aboard the R/V Edwin Link are gratefully acknowl-

edged for their help and cooperation. We thank P.

Bouchet , A. Waren and H. Zibrowius for criticalK read-

ing the manuscript. Scanning Electron Micrographs are

by A. Bonfitto. Thanks are due to H. Zibrowius for the

scleractinian determinations. This is Istituto di Geologia

Marina scientific contribution number 952.

LITERATURE CITED

Barrier, P., Zibrowius, H , Lozouet, P.. Montenat, C, Ott d Es-

tevou. P., Serrano, F and Soudet, H.J. 1991. Une faune

de fond diir du bathyal superieur dans le Miocene terminal

des Cordillers Etiques (Carboneras, SE Espagne). Mesogee

51:3-13.

Bertolaso, L. and S. Palazzi. 1994. /p/iifus Jeffreys, 1883, un

sinonimo di Stylotrochus G. Seguenza, 1876 (appunti di

malacologia neogenica:!). Bollettino Malacologico 29(9-

12):286-290.

Beu, A. G. 1978. Habitat and relationships of Iphitctia neo-

zetanica (Dell) (Gastropoda; Epitoniidae). New Zealand

Journal of Marine and Freshwater Research 12(4):391-

396.

Bouchet, P. and A. Waren. 1986. Revision of the Northeast

Atlantic bathyal and abyssal Aclididae, Eulimidae, Epi-

toniidae (Mollusca. Gastropoda). Bollettino Malacologico,

Supplemento 2:299-576,

Palazzi, S. and .\. X'lllari. 1996. Malacofaune batiali plio-

pleistoceniche del Messinese.2: Capo Milazzo. Naturalista

siciliano, ser.4, 20 (3-4):237-279.

Rindone, V. and A. N'azzana. 1989. Alcune specie di niolluschi

delle argille batiali del piano siciliano (Pleistocene inf.)

della cava di Archi (Reggio Calabria). Bollettino Mala-

cologico 25 (5-8): 233-240.

Roberts. H. H., P. Aharon, R. Carney, J. Larkin and R. Sassen.

1990. Sea floor responses to hydrothermal seeps, Louisiana

continental slope. Geo-Marine Letters 10:232-243.

Taviani, M. and B, Sabelli. 1982. Iphitus (Mollusca, Gastrop-

oda) a deep-water genus new to the Mediterranean sea.

Lavori della Societa Malacologiea Ilaliana, .\tti del \'°Con-

vegno della Societa Malacologiea Italiaiia 191-131.

Vazzana, A. 1995. Malacofauna batiale del Pleistocene in-

feriore del Vallone Catrica (Reggio Calabria, Italia). Bol-

lettino Malacologico 31 (5-8); 143-162.

THE NAUTILUS 110(3):97-101, 1997

Page 9"

Verification of the Specific Status of the Endangered Anthony's

River Snail, Athearnia anthonyi, Using Allozyme Electrophoresis

Robert T. Dillon. Jr.

Department of Biolog\,

College of Charleston

Charleston, SC 29-424 USA

Steven A. Ahlstedt'

Tennessee \'alley Authority

Clean Water Initiative

Norris, TN 37828 USA

ABSTRACT

Although nominally the single surviving representative of a

unique pleurocerid taxon, Athearnia anthonyi (Budd, in Red-

field, 1854) is so rarely collected that even its specific status has

been uncertain We used allozy me electrophoresis to compare

a population of A. anthonyi to the similar pleurocerid snail,

Leptoxis praerosa (Sa\. 1821), co-occurring with it in the Se-

quatchie River of Tennessee, and to a second population of L.

praerosa collected approximately 500 km distant. Observed

levels of heterozygosity offered no evidence of inbreeding or

unusually severe population bottlenecking in any of these pop-

ulations. Strikingly different allele frequencies at five of the

eleven enzyme loci examined, together v\ ith differences in shell

morphology especially apparent in young individuals, con-

firmed that A. anthonyi and L. praerosa are distinct species.

Their similarity at the six loci remaining supports previous

suggestions that Athearnia may be a subgenus of Leptoxis.

Key words: Athearnia. Leptoxis. Pleuroceridae, freshwater

gastropods, proteins, enz\ mes. Tennessee.

INTRODUCTION

The pleurocerid genus Atliearnia was proposed by Mor-

rison (1971) to include the two species, Anculosa an-

thontji (Redfield, 1854) and Anculosa crassa (Haldeman,

1842), previously but incorrectly included in the genus

Eunjcaelon (Goodrich, 1931). Some authors (Davis, 1974;

Burch, 1982) have subsequently considered Athearnia to

be a subgenus of the widespread genus Leptoxis, but we

follow Bogan and Parmalee (1983), Garner (1992), and

the Federal Register in retaining its generic rank. The

distinction between the two nominal Athearnia species,

A. anthonyi and A. crassa, has been confused for some

years. In any case, since A. crassa is generally considered

to be e.xtinct (Bogan & Parmalee, 1983), A. anthomji

' Present Address: U.S. Geological Survey, 1820 Midpark Drive,

Knoxville TN 37921 USA

appears to be the sole existing representative of this dis-

tinctive pleurocerid taxon.

Historicallv , populations of Athearnia have been re-

corded from the Tennessee River and its larger tribu-

taries upstream from Muscle Shoals, Alabama, including

the clinch, Powell, Nolichucky, Little Tennessee, French

Broad, Sequatchie, and Elk Rivers (Goodrich, 1940).

However, the impoundment of the Tennessee drainage

that began in the 1930's, improvement of the stream bed

for navigation, and general habitat degradation due to

farming, mining, and industry eliminated Athearnia from

the great majority of its former habitat. Bogan and Par-

malee (1983) feared that Athearnia "may be extinct

throughout its range except for a possible relic popula-

tion.

More recently , it has become clear that small but ap-

parently viable populations of A. anthonyi still inhabit

lower regions of the Sequatchie River in Marion County.

Tennessee, and Limestone Creek in Limestone County,

Alabama (Garner, 1992). These snails also survive in at

least one channel of the main Tennessee River down-

stream from the mouth of the Sequatchie (Jenkinson,

1994; Garner, 1994). In early 1994, A. anthonyi became

the first pleurocerid gastropod formally listed as "en-

dangered" by the United States government (Federal

Register 59:17994-17998). In addition to protecting ex-

tant populations, the A. anthonyi recovery plan (U.S.

Fish & Wildlife Service, 1996) calls for the future re-

establishment of at least a few populations into portions

of the snail's historic range.

.\lthough great interest has focused on the distribution

and abundance of A. anthonyi, questions remain re-

garding its ta.xonomic status. Adult A. anthonyi may be

confused with large, senescent Leptoxis praerosa (Say,

1821). a much more common pleurocerid with which

Athearnia often occurs. Juveniles of A. anthonyi have

not often been described. Given that trematode infection

may induce extreme and anomalous growth in gastropod

hosts (Sturrock, 1966; Hodasi, 1972), it has seemed pos-

sible that Athearnia may be parasitically castrated, gi-

gantic, L. praerosa.

Allozyme electrophoresis has become established as an

Page 98

THE NAUTILUS. Vol. 110. No. 3

important tool for measuring genetic di\ersit\ both with-

in and among populations of pleurocerid snails (Cham-

bers. 1978; 1980; Stiven & Kreiser, 1994). Published work

to date has. howe\er. concentrated e.\clusi\el\ on the

diverse genus Goniobasis, primarily an inhabitant of

smaller streams. Genetic variation seems to be unusually

low within Goniobasis populations, but unusually high

between them (Dillon & Davis, 1980). The levels of both

intra- and interpopulation gene flow also seem to be

unusually low (Dillon. 1988a). Significant gene frequen-

cy differences have been reported between samples of

Goniobasis taken at distances as short as 500 m (Dillon,

1988b). GeographicalK' distant Goniobasis proxima (Say,

1825) populations may share no alleles at as many as si.\

enzyme loci of seven studied, yet show no evidence of

reproductive isolation (Dillon, 1984; 1986; 1988a). Dis-

tinct species of Goniobasis generalK' share alleles at very

few enzyme loci (Chambers, 1980; Dillon & Davis, 1980).

So although no data have been published on divergences

in other pleurocerid genera to this date, there is reason

to e.xpect that if A. anthonyi is indeed different from L.

praerosa, differences in alloz\ me frequencies will be ap-

parent.

In the present work, we use allozyme electrophoresis

to compare a population of A. anthonyi from the Se-

quatchie River to a co-occurring population of L. pra-

erosa and to a population oi L. praerosa from the Duck

River 500 river kilometers distant (appro.ximately 350

km downstream, then 150 km upstream). In addition to

verifying the specific status oi A. antlionyi, our data also

bear on two ancillary questions: the systematic placement

of the genus Athearnia and the degree to which an im-

portant population of A. anthonyi may be inbred or

bottlenecked.

MATERIALS & METHODS

Athearnia anthonyi and Lcptoxis praerosa were col-

lected from a site on the Sequatchie River in Marion

County, Tennessee. Leptoxis praerosa were most com-

mon in riffle areas, while A. anthonyi were more com-

mon on larger submerged objects (rocks, snags, etc.) in

the transition areas between rifiles and pools. A second

sample of L. praerosa was collected from the Duck River

at Lillard Mill, Marshall County, Tennessee. Voucher

specimens of A. anthonyi have been deposited in the

Ohio State University Museum of Biological Diversit\

(catalog number 19820). Individuals were transported

alive to Charleston, S.C, where the shells were cracked

and the digestive glands examined for parasites. Tissues

were then frozen in a tris-phosphate bufter at — 70°C

and examined electrophoreticalK within one week.

Horizontal starch gel protein electrophoresis was per-

formed on whole animal homogenates as has been pre-

viously described (Dillon, 1985; 1992). We initially

screened 15 A. antlionyi and 14 Secjuatchie River L.

praerosa for variation in 10 enzyme systems: aspartate

aminotransferase (AAT), esterases (EST a-napthyl ace-

tate as substrate), glucose phosphate isomerase (GPI),

alcohol dehydrogenase (ADH, hexanol as substrate),

mannose phosphate isomerase (MPI), octopine dehydro-

genase (ODH), phosphogluconate deh\ drogenase

(6PGD), phosphoglucomutase (PGM), sorbitol dehydro-

genase (SDH), and superoxide dismutase (SOD). Simple

Mendelian inheritance of codominant alleles has been

demonstrated at the 6PGD and PGM loci in Goniobasis

floridensis (Reeve, 1860) by Chambers (1980) and at the

GPI, ODH, and ESTl loci in G. proxima by Dillon (1986).

It should be noted that a great many esterase allozymes

are generally detectable in pleurocerids. but that to date

only the strong, slowly-migrating products ot the "ESTl"

locus are genetically interpretable.

An effort was made to examine alloz>me phenot>pes

in as wide a range of buffers as possible. The following

gel buffers were emploved: Tris-Cit 6.0 (for PGM, GPI,

ODH, and MPI), AP 6.0 (for 6PGD), TEB S.O (for EST

and 6PGD), Poulik (for SDH, ODH, and GPI), and TEB

9.0 (for SOD, ADH, AAT, PGM, and SDH). Recipes for

most of these buffers and stains were obtained from Shaw

and Prasad (1970) or Harris and Hopkinson (1976), mod-

ified for agar overlay in many cases.

Also included in the initial survey for allozyme vari-

ation were C. proxima standards (population SL'GR of

Dillon, 1984). Leptoxis and Athearnia bands were la-

beled by their mobilities in millimeters relati\e to this

population in a standard buffer. As was the case in 1984,

some "hidden variation was detected at the ODH locus

- bands not resolved b\ the standard Tris-Cit 6.0 buffer

were detected with a Poulik buffer. These isozyme classes

were labeled "S". "F", and "VF" for their migration in

Poulik gels.

After the initial screening, we concentrated our in-

vestigations on the five putative enzyme loci apparently

varying in our comparison of A. anthonyi and L. pra-

erosa. Ultimately we examined 37 A. anthonyi, 29 L.

praerosa from the Sequatchie River, and 25 indixidual

L. praerosa from the Duck Ri\er. Gene frequencies and

observed heterozygosities were calculated using BIOSYS

version 1.7 (Swoftord & Selander, 1981).

RESULTS

Figure 1 compares representati\ e shells from Sequatchie

populations of A. anthonyi and L. praerosa. Athearnia

anthonyi is distinguishable b\ the higher shoulder of its

w horls, beginning as a pronounced keel (or "carina ) but

becoming less distinct with age. The oldest A. anthonyi

are quite smooth, and come to resemble large L. pra-

erosa.

\er\ little evidence of parasitism was detected in ei-

ther Sequatchie pleurocerid population. OnK 2 oi 29 L.

praerosa showed obvious trematode infections, while none

of the 37 .4. anthonyi appeared to be infected. Thus the

large sizes attained by individual A. anthonyi do not

appear to be due to parasitic gigantism.

R. T. Dillon, Jr. and S. A. Alilstedt, 1997

Page 99

.#^

a

I

10 mm

Figure 1. Growth series for Athi-anna nntlionyi (abose— live specimens) and Lepioxis praerosa (below) from tfie Sequatchie River,

TN.

No variation was detected between populations of A.

anthomji and L. praerosa at AAT, ADH, 6PGD, SDH,

SOD, or at a faster-migrating PGM locus. Divergence

was striking, however, at the five loci listed in Table 1.

The two svmpatric populations share no alleles at ESTT

GPI, MPI,' or at the slower PGM locus, 'PGMS". And at

the ODH locus, A. anthomji appears to be fixed for an

allele present at a frequenc\ of onl\ 0.345 in Seciuatchie

/,. praerosa.

Although spatially separated by over 500 km, the two

L. praerosa populations seem to have remained rather

similar genetically. Table 1 shows that the differences in

alloz) me frequenc\ at the GPI, MPI, and ODH loci are

significant, but not fixed. The Sequatchie population ap-

pears substantially more polymorphic.

Rather low levels of genetic variation were detected

within these three populations of pleurocerids, as has

been commonly reported in the past, jutlging b\' the 95%

criterion, onl\ four of the 15 loci show n in Table 1 could

be considered polymorphic. The fit to Hardy- Weinberg

expectation at these four loci was verv good, however.

DISCUSSION

It is clear that previous misgivings about the specific

status of A. anthomji were unfounded. Although not as

divergent as typical species of Goniobasis, the data pre-

sented in Table 1 show clearly that A. anthomji and L.

praerosa are reproductiveK isolated in the Sequatchie

River. Their shell morphologies are distinct. The greater

size attained by A. anthomji appears to be a natural

Table 1. .Allele frequencies and observed heterozygosities (H)

at variable loci in populations of Athearnia anthomji from the

Sequatchie River. Leptoxis praerosa from the Sequatchie River,

and L. praerosa from the Duck River, Sample sizes 37, 29, and

25, respectively.

Page 100

THE NAUTILUS, Vol. 110, No. 3

feature of development, not a consequence of parasitic

gigantism.

The level of intrapopulation variation in A. anihoniji

is low compared to that of most other organisms, but

comparable to that seen in L. praerosa and in other

pleurocerids. We did discover one highly polymorphic

locus in A. arithonyi, GPl, showing genotype frequencies

not significantK different from expectation under Har-

dy-VVeinberg equilibrium (Yates-corrected x' = 2.53).

Thus we are unable to detect evidence of inbreeding or

an unusually severe population bottleneck.

Previous electrophoretic investigations of Goniobasis

populations have generally detected higher levels of di-

vergence than we report here between Sequatchie River

and Duck River populations of L. praerosa. Multiple

fixed differences are often observed between G. proxima

populations isolated in small creeks at distances as great

as 500 km. The less dramatic differences between con-

specific Leptoxis populations reflected in Table 1 may

be a consequence of their adaptation to larger rivers.

Populations of Leptoxis ma\ have inhabited occasional

rocky shoals down the length of the Tennessee River

before its impoundment, connecting such tributary pop-

ulations as those of the Sequatchie and Duck Rivers in

stepping-stone fashion.

Previous studies have shown greater divergence be-

tween pleurocerid species than uncovered here. Dillon

and Davis (1980) and Chambers (1980) reported that

typical species of Goniobasis rarely share any similarity

at any allozyme locus. However, L. praerosa and A.

anthonyi were indistinguishable at six of the eleven loci

initially screened, and seem to share one fairly common

allele at a seventh locus (ODH). This constitutes some

support for the nomenclature ot Burch (1982) and Dillon

(1989) in which Athearnia is placed as a subgenus of

Leptoxis. Regardless of whether they represent a unique

genus or a unique subgenus, the populations of A. an-

thonyi now restricted to just a tew rivers of central Ten-

ne.ssee are a valuable resource for evolutionary study,

and warrant protection at the highest levels.

ACKNOWLEDGMENTS

Charles Saylor assisted in collecting the snails, Vivian

Johnson delivered them to Charleston, S.C., and Amy

Wethington was a great help running the gels. Technical

reports were provided by John Fridell.

LITERATURE CITED

Bogan, A., and P. Parmalee. 1983. Tennessee's rare wildlife,

volume II: The mollusks. Tennessee Wildlife Hesnurces

.Agency, Nashville, TN. pages 81-84.

Burch, J. 1982. North Auierican fre.shwater snails; identifi-

cation keys, generic synon\ my, supplenicntai notes, glos-

sary, references, index. Walkerana -}:l-'36.5.

Chambers, S. M. 197S. An elfitidpiiorelically delected siljimg

speeiesof " Coniolmsis floi ulcnsi.s" (Mesogastropoda: Pieu-

roceridae). Vlalacologia 171.57-162.

Chambers, S. M. 1980. Genetic divergence between popu-

lations of Goniobasis occupying different drainage sys-

tems. Malacologia 20:11.3-120.

Davis, G. 1974. Report on the rare and endangered status of

a selected number of freshwater gastropoda from south-

eastern U.S.A. U.S. Department oi Interior, Fish and W'ild-

life Service, Washington, DC. 51 p.

Dillon, R. T., Jr. 1984. Geographic distance, environmental

difference, and divergence between isolated populations.

Systematic Zoology 33:69-82.

Dillon, R, T,, Jr 1985, Correspondence between the buffer

systems suitable for electrophoretic resolution of bivalve

and gastropod isozymes. Comparative Biochemistry and

Physiologv 828:643-645.

Dillon, R. T , Jr. 1986. Inheritance of isozyme phenotype at

three loci in the freshwater snail Goniobasis proxima:

Mother-offspring analysis and an artificial introduction.

Biochemical Genetics 24:281-290.

Dillon, R. T., Jr. 1988a. Evolution from transplants between

genetically distinct populations of freshwater snails. Ge-

netica 76:111-119.

Dillon, R. T., Jr. 1988b. Minor human disturbance influences

biochemical variation in a population of freshwater snails.

Biological Conservation 43:137-144.

Dillon, R. T., Jr. 1989. Karyotypic evolution in pleurocerid

snails: 1. Genomic DNA estimated by flow cytometry.

Malacologia 31:197-203.

Dillon, R. T., Jr. 1992. Electrophoresis IV, nuts and bolts.

World Aquaculture 23(2):48-51,

Dillon, R, T., Jr., and G. M, Davis 1980. The Goniobasis of

southern Virginia and northwestern North Carolina: ge-

netic and shell morphometric relationships. Malacologia

20:83-98.

Garner, J 1992. .-V survey for .Anthony's River Snail, Athear-

nia anthonyi ("Budd" in Redfield, 1854) in tributaries of

the middle reaches of the Tennessee River, .\quatic Re-

sources Center, Franklin, TN 17 p

Garner, J. 1994. Survey of mollusks, Tennessee River mile

412.1, .Aquatic Resources Center, Franklin, TN, 2 p.

Goodrich, C. 1931, The pleurocerid genus Eurycaelon. Oc-

casional Papers of the Museum of Zoology, University of

Michigan 223:1-9,

Goodrich. C, 1940, The Pleuroceridae of the Ohio River

system. Occasional Papers ol tlie Museum of Zoology, Uni-

versity of Vlichigan 417:1-21,

Harris, H., and D Hopkinson. 1976. Handbook of enzyme

electrophoresis in human genetics. Elsevier, New York.

Hodasi, J. K. M. 1972. The effects of Fasciola hepatica on

Lymnaea Iruncatula Parasitology 65: 359-369.

Jenkinson, J. 1994 PYeshwater mollusk survev at CSX railroad

bridge, near Bridgeport, .Alabama, Tennessee River Mile

414.5. Tennessee \'alley Authority, Chattanooga, TN. 15 p.

Morrison, J. 1971. Athearnia a new name for a genus of

pleurocerid snails. The Nautilus 84:110-111.

Shaw, C. R., and R. Prasad. 1970. Starch gel electrophoresis

ot enzymes — a compilation ot recipes. Biochemical Ge-

netics 4:297-320,

Stiven, A , and B, Kreiser. 1994. Ecological and genetic dif-

ferentiation among populations of the gastropod Gonio-

basis proxima (Say) in streams separated by a reservoir in

the Piedmont of North Carolina, Journal ol the Elisha

Mitchell Scientific Society I10:.53-67,

Sturrock, IV M, 1966, The intluence ol inlection with Schis-

tosoma mansoni on the growth rate and reproduction of

R. T. Dillon, Jr. and S. A. Ahlstedt. 1997 Page 101

Bioniphalana pjeiffch. Annals ot Tropical Medicine and United States Fish & Wildlite Service. 1996. Technicai/Agen-

Parasitology 60: 187-197. cy Draft Anthony's Riversnail Recovery Plan. Atlanta GA.

Swofford, D. L.. and R B Selander. 1981 BI08YS-1; a FOR- 31pp.

TRAN program for the comprehensive analysis of elec-

trophoretic data in population genetics and svstematics.

Journal of Heredity 72:281-283.

TliE NAUTILUS 110(3);102-106, 1997

Page 102

Induced Metamorphosis of Freshwater Mussel Glochidia on

Nonhost Fish

Sheila C. Kirk

James B. Layzer

National Biological Service

Tennessee Cooperati\e Fishery

Research Unit

Tennessee Technological University

Cookeville, TN 38505 USA

ABSTRACT

Intraperitoneal implants of Cortisol, an immunosuppressant,

suspended in liquid cocoa butter were administered to nonhost

fish species. Fish were then infested with glochidia of fresh-

water mussels to determine if transformation \\ ould occur after

immune system manipulation. Glochidia ot Venustacoitcha sima

(Lea, 1838) {Villosa iris complex) transformed on orangethroat

darters, Etheostoma spectabile (Agassiz, 1854), after injection

of Cortisol at concentrations ranging from 0.005 to 0.040 mg

per g of fish weight. Banded sculpins, Cottus carolinae (Gill,

1861), transformed glochidia of Villoaa taeniata {Conrad, 1834)

following Cortisol injections ranging from 0.005 to 0.020 mg-

g. Creek chubs, Scinolilus atromaculatus ^Mitchill, 1818), failed

to transform glochidia of either mussel species after Cortisol

administration. No juvenile mussels were collected from non-

injected or sham injected fish during any experiment. Cortisol-

induced immunosuppression facilitates metamorphosis of glo-

chidia on some nonhost fish species. Refinement of this tech-

nique could provide an alternative means for propagating

freshwater mussels, especial!) for those endangered species that

utilize unknown hosts.

Key Words. Cortisol, inmiunosuppression, transformation, glo-

chidia, implants, Linionidae.

INTRODUCTION

The obligate parasitic stage in the life cvcie of freshwater

mussels on specific host fish ma\ limit reproduction of

many populations. Because adequate biological inven-

tories were not conducted before impoundment of most

river systems, many fish species were extirpated before

identification of glochidia! hosts could occur (Schindler,

1989). Hosts are known for onl\ one-quarter of the Nortli

.American species of mussels (Hoggarth, 1992; Walters,

1994). Although glochidia of a few mussel species have

laeen transformed in a sterile culture medium (Isom &

Hud.son, 1982), metamorpliosis of other species has been

less successful. Consequently, artificial propagation of

many freshwater mussel species that utilize unknown

hosts has not been possible.

Other than the use of known hosts and artificial media,

few advancements have occurred in methods of artificial

propagation of freshwater mussels. With the relatively

recent threat of the zebra mussel, Dreissena pohjmorpha

Pallas, 1771, concern for the fate of our native fauna has

increased efforts to sustain diversity and protect existing

populations, especially endangered or threatened species

without identified hosts. Efforts include assessment of

relocation into areas with little risk of zebra mussel in-

vasion and the use of mussel refugia such as hatcheries

(Shannon ct ai. 1993).

Host specificity of freshwater mussels is believed to

have an immunological basis (Reuling, 1919; .Arey, 1932;

Meyers et at.. 198U). Isom and Hudson (1982; 1984)

demonstrated that components in fish blood necessary

for initiation of glochidial transformation are not species

specific and can be found in the blood of all fish. More-

over, horse and neonatal calf serum are also suitable

media for glochidial transformation (Keller & Zam, 1990).

The iiuplications of these studies suggest that the inuiume

response of the host is the determining factor in mussel-

host specificity.

Immune responses of both host and nonhost fish to

glochidia of freshwater mussels ha\e been documented

(Reuling, 1919; Arey, 1932; Meyers et at., 1980; Bauer

& Vogel, 1987). Responses of host fish consist of host

tissue proliferation which encysts the glochidium within

a few hours after attachment (d'Eliscu, 1972; Waller &

Mitchell, 1989). In host and nonhost species, humoral

and cell mediated responses of the immune sy stem occur.

These immune responses include the presence of specific

antibodies to the parasites (Reuling, 1919; Meyers et ai.

1980; Bauer & \'ogel, 1987), an increase in eosinophils

and other leukocv tes around glochidial c>sts (Are) , 1932),

hyperplastic sloughing of host epithelial tissue (Meyers

et ai. 1980), and in some cases, cytolitic destruction of

the glochidium (Reuling, 1919; Arey, 1932).

C^ortisol, a major corticosteroid in teleosts (Idler &

Truscott, 1972) is released during stress (Bennett & Wolke,

1987; Schreck, 1990) and produces immunosuppressive

S. C; Kirk and J. B. Layzer, 1997

Page 103

effects. Cortisol can be administered to produce a dose-

dependent elevation of plasma Cortisol and immuno-

suppression (Pickering & Duston. 1983). Cortisol reduces

K mphocNte concentrations circulating in the blood, re-

duces KmphocNte participation in inflammation and re-

duces other inflammator\ cells (Pearson et al.. 1978). We

h\ pothesized that the effects of Cortisol might inhibit the

abilit> of nonhost fish to respond to glochidia b> de-

creasing the number of antibodies and reducing inflam-

mation around glochidial cysts, thus allowing glochidia

to attach, enc\st, and metamorphose into juveniles. In

this paper, we e\aluate the effects of Cortisol, adminis-

tered in a vehicle of liquefied cocoa butter, on glochidial

metamorphosis.

MATERIALS & METHODS

Collection of fish and mussels: For our experiments,

we selected tw o brad\ ticitc species of mussels based on

their narrow host specificity and availability. Gravid

Veniistaconcha sima (Lea. 1838)', were collected from

the Collins River in Grundy County, Tennessee. Gra\ id

Villosa taeniata (Conrad, 1834) were collected from the

Roaring River in Overton County, Tennessee. Rockbass,

Ambloplites rupestris (Rafinesque, 1817), is the only

known host of V. taeniata (Gordon et al., 1994), and

banded sculpin, Cottus carolinae (Gill, 1861), is the only

known host of V. sima (unpublished data). To avoid

premature expulsion of glochidia, mussels were kept at

approximately 4°C in pans of shallow water until needed.

Laboratory established nonhost fish species were used as

experimental fish, with the exception of creek chub, Sf-

motilus atromaculatus (Mitchill. 1818). The suitabilit\

of creek chub as hosts for ^'. sima has not been previousK-

tested in the laborator\-; howe\er, examination of wild

fish did not re\eal an\ infestations (unpublished data)

A backpack electrofishing unit was used to collect fish

from the East Blackburn Fork, Putnam County, Ten-

nessee. Populations of banded sculpins, creek chubs,

rockbass, and orangethroat darters, Etheostoma spect-

ahile (Agassiz, 1854), are abundant in this stream. No

mussels occur in the East Blackburn Fork, and an im-

passable waterfall downstream prevents any possible up-

stream movement of fish that ma\' have been exposed to

glochidia. Fish were transported back to the laborator\',

acclimated to laborator\ conditions, and held in 38 liter

aquaria

Implantation and infestation: Pure cocoa butter was

melted at a temperature of 40°C. A predetermined

amount of Cortisol was dissolved in ethanol. Ethanol was

used at a rate of 10^7 the volume of cocoa butter. The

'This species is generally considered a member of the Villosa

iris complex but was elevated by Gordon (1995). In this paper,

we use Venustaconcha sima to clearly identify which of the

two members of the Villosa iris complex occurring sympatar-

icaii\ in the Collins River were used in our experiments

dissolved Cortisol was suspended in the liquid cocoa but-

ter and allowed to mix in a water bath. Average fish

weight was used to calculate the amount of Cortisol per

gram of fish weight needed to obtain the desired con-

centration within the fish. Orangethroat darters received

50 m1 implants and creek chub and banded sculpins re-

ceived 100 /ul implants.

Infective glochidia were obtained from gravid females

and exposed to salt to determine maturity (Zale & Neves,

1982). After fish were anesthetized with tricaine meth-

anesulfonate (MS 222), they were placed on a wet paper

towel to reduce the removal of mucus during the injec-

tion process. The liquid cocoa butter implant was injected

into the peritoneal cavit\ of the fish with a 1 ml tuber-

culin hypodermic syringe. The liquid cocoa butter solid-

ified within the cavity of the fish and acted as a solid

implant during the experiment. PresumabK, Cortisol

leaked slowK from the implant, but the kinetics of this

transfer were not addressed, Pickering and Pottinger

(1985) observed maximum mean plasma Cortisol levels

of 9 and 15 ng/ml for brown trout, Salmo triitta (Lin-

naeus, 1758), that received 10 and 20 mg Cortisol re-

spectively. ImmediateK after Cortisol injection, glochidia

were pipetted onto the left gills of each fish. Fish were

then transferred to fresh water and revi\ed. Each ex-

perimental group was kept in a separate aquarium.

Experimental design Treatments varied among exper-

iments, but generalK' included a known host control group

to measure glochidial viabilit\. The no-injection (NI)

group \erified that the test species did not normally serve

as a host. Sham injected fish received injections of pure

cocoa butter only, and were used to determine if the

injection process or cocoa butter had an effect on glo-

chidial transformation. Cortisol injected fish received

concentrations that ranged from 0.005 to 0.040 mg/g.

Preliminary experiments determined the feasibility of

working with these concentrations, which we initially

based on those used b\ Pickering and Duston (1983) and

Pickering and Pottinger (1985).

Aquaria were siphoned daily and the siphonate was

examined under a dissecting microscope at lOX to 20X

magnification. The number of glochidia and juveniles,

water temperature and fish mortality were recorded.

Criteria for recognizing juvenile mussels included the

presence of tw o adductor muscle scars, closed valves, and

movement w ithin 24 hours of collection. If these criteria

w ere not met, organisms w ere considered untransformed

or partially transformed glochidia. Experiments were

terminated when no juveniles w ere collected from host

fish for one week after the last juvenile was collected, or

examination of the gills revealed no enc\sted glochidia.

.\11 fish were used for onl\ a single infection experiment,

RESULTS

Metamorphosis of I enustaconcha sima on nonhost fish:

.\ total of 58 juvenile Venustachona sima were trans-

formed on Cortisol injected orangethroat darters. Trans-

formation occurred only on orangethroat darters injected

Page 104

THE NAUTILUS, Vol. 110, No. 3

Table I. NumlKTi ot ju\eiiiles collected troni Cortisol injected

orangethroat darters infested with glochidia of Venustaconcha

sirna in each experiment. Dates indicate v\ hen fish were injected

and infested. Numbers in parentheses are the average number

of juveniles transformed per fish (NI = No-injection)

Table 2. Nuinljers ol ju\ eniles collected from Cortisol injected

banded sculpins (October and May) and orangethroat darters

(December) infested with \ illosa taeniata in each experiment.

Dates indicate when fish were injected and infested Numbers

in parentheses are the average number of juveniles transformed

per fish (N! = No-injeclioii).

* Fish mortalitv was 2:88'

in March, May, and June (Table 1). Glochidia from V.

sima transformed on fish injected with 0,005 to 0.040

mg, g Cortisol. Metamorphosis never occurred in sham

injected and no-injection (NI) treatment groups during

any experiment. Glochidia did not transform on either

orangethroat darters or creek chubs injected in August.

Juvenile transformation varied among experiments.

For instance, the average number of juveniles trans-

formed per orangethroat darter injected with 0.005 mg/

g Cortisol ranged from 0.00 to 1,89 among experiments

(Table 1 ). There w as a positive correlation between Cor-

tisol concentration and metamorphosis within experi-

ments; however, this relationship was confounded by

high mortality in some treatment groups. Mortality var-

ied within and among experiments; most deaths occurred

soon after injection, and were likely the result of physical

injury.

Metamorphosis of \ illosa taeniata on nonhost fish: A

total of six juvenile X illosa taeniata were collected from

banded sculpins injected with Cortisol (Table 2). Trans-

formation of V. taeniata glochidia did not occur on

orangethroat darters or creek chubs injected with Cortisol.

In one experiment, nonhost species sloughed glochidia

quickly after the infestation. After examination of their

gills revealed no remaining glochidia, the fish were rein-

fested 1 1 days after the initial infestation. Although creek

chubs and banded sculpins retained glochidia longer after

the second infestation, metamorphosis did not occur.

Metamorphosis of glochidia on host fish: Transfor-

mation of glochidia of Venustachoncha sinia on the host,

banded sculpins, varied between experiments Banded

sculpins transformed 35 juvenile mussels in March and

six juvenile mussels transformed on banded sculpins in

May. When used as a host control, all banded sculpins

were held in one aquarium during each exi)eriment, and

individual variation in juvenile transformation could not

be asse.ssed.

Because of the aggressive nature of rockbass, the host

of Villosa taeniata, individuals were kept in separate

aquaria. Transformation of glochidia of V. taeniata on

rockbass varied greatly among individuals and among

experiments (Table 3). Ju\enile transformation on in-

dividual fish ranged from 0 to 13 juveniles in one ex-

periment, to a range of 7 to 262 ju\eniles in another

experiment. There was no clear relationship between the

number of glochidia used to infest host fish or water

temperature and the number of juveniles recovered from

individual rockbass.

DISCUSSION

Cortisol affects leukocyte circulation, influences immune

effector mechanisms in lymphocytes, modulates acti\i-

ties of inflammatory mediators, and modifies protein,

carbohydrate, and fat metabolism (Tizard, 1988). In fish,

this results in immunosuppression and an increased sus-

ceptibility to infectious diseases such as furunculosis and

bacterial fin-rot (Pickering & Duston, 1983; Pickering &

Pottinger, 1985).

The present study has shown that cortisol-induced im-

munosuppression can facilitate glochidial metamorpho-

sis on nonhost fish species. Presumably, Cortisol sup-

pressed the humoral and cell-mediated responses ob-

served in glochidial infestations by Arey (1932) and Mey-

ers ct al. (1980). .Although the strength of fish immune

responses after Cortisol administration was not measured,

transformation occurred on experimental nonhost species

only when the immune system was compromised. These

results are consistent with the hypothesis that host spec-

ificity of freshwater mussels is immunologically con-

trolled.

Glochidia of Venustachoncha sima transformed on

orangethroat darters in three of four experiments at the

lowest concentration (0.005 mg, g) used. Transformation

of glochidia of Vtllusa taeniata occurred on banded scul-

pins during two experiments. For this species, fewer ju-

veniles were collected from fish injected with the same

concentration used for orangethroat darters. Creek chubs

failed to transform glochidia from either mussel species

S. G. Kirk and J. B. Layzer, 1997

Page 105

Table 3. Total It'iigth. luunber ot glochulia reco\erecl, niinilier ot ju\eniles recovered, metamorphosis period, and water temperature

(±S.D.) for iinlix iiliial rockl>ass infested with glochidia ot \'illosa tacniata. Dates indicate when fish were infested.

when injected with Cortisol concentrations of 0.005 and

0.010 mg/g Cortisol, suggesting that responses to (or up-

take of) Cortisol varies with species. Threshold levels of

Cortisol in the blood may exist that allow glochidial trans-

formation, and these levels ma\' not have been reached

in creek chubs.

Fish experience periods of natural elevation ot plasma

Cortisol (Idler & Truscott, 1972). Spring and siunmer

elevations of plasma Cortisol and other circidating cor-

ticosteroids coincide with migration, smoltification of ju-

venile anadromous salmonids, se.xual maturation and

spawning of salmonids (Thorpe el al.. 1987; Pickering

& Pottinger, 1983; Pickering & Christie, 1981; Anderson,

1990). Cortisol concentrations used in this stud\' should

have elevated Cortisol le\els to the ph\siological range

of fish undergoing stress or periods of natural Cortisol

elevation (Pickering & Duston, 1983; Pickering & Pot-

tinger, 1985; Thorpe et al., 1987), Periods during the life

cycle of fish where Cortisol elevation has been observed

may correspond with the presence of some species of

host fish over mussel beds during the discharge of glo-

chidia (Farzaad, 1991). Perhaps mussels evolved to take

advantage of weakened immune responses of host species

during spawning or other periods of natural immuno-

suppression.

Host and nonhost fishes exhibit humoral responses of

similar strengths, but host and nonhost species reject glo-

chidia at different rates (O Connell, 1991 ). Although Cor-

tisol injected fish transformed relati\el\' few juveniles

during these experiments, transformation per fish was

comparable to hosts in some cases. For instance, 0,020

mg/g Cortisol injected orangethroat darters transformed

an average of 0.50 juvenile \'. sirua per fish in Ma\ . Host

fish (banded sculpins) onl\' transformed 1.00 juvenile per

fish in the same experiment. Individual variations within

and among experiments in glochidial transformation of

\'. tacniata on host fish may also reflect a seasonal aspect

in the susceptibility of the host (e.g., host spawning sea-

son) or the ability of glochidia to be infective at times

of the year other than the normal discharge period. We

have observed similar variation among trials and indi-

viduals in the numbers of juveniles transformed per host

for other mussel species. Variation of an order of mag-

nitude or more in the mean number of juveniles trans-

formed per fish has been reported in other studies as well

(e.g., Zaie & Neves, 1982). Although the number of glo-

chidia attaching to fish is difficult to control and may

contribute to some of the variation in the numbers of

juveniles transformed, there was no relationship in the

apparent numbers (glochidia + juveniles recovered) of

Villosa tacniata glochidia attaching to individual rock

bass and the numbers of juveniles produced,

.Additional development of the cortisol-induced im-

munosuppression of nonhost fish is needed before the

technique can be applied on a large scale as a culture

method for freshwater mussels. Nonetheless, refinement

of this technique could have widespread management

implications, especially as a means for propagation of

endangered species. This method could also be used to

reestablish populations of nonendangered mussel species

into new and reclaimed habitat. Application of Cortisol

immunosuppression techniques to host species could per-

haps increase juvenile transformation for reintroduction

studies. Also, application of this technique to non-host

fish would provide an alternative means of propagating

mussel species that have rare or endangered hosts.

The need for further research on the ecological re-

quirements, including identification of fish hosts, habitat,

and ph\sicochemical information of mussel species can-

not be over emphasized. These ecological requirements

hold the answers to developing self-sustaining popula-

tions of freshv\ater mussels. Because survival and fitness

of juvenile mussels produced b\' means of artificial prop-

agation has not been determined (O Connell, 1991), ar-

tificial propagation methods should be used with caution.

Any long-term artificial propagation could alter the ge-

netic integritv of mussel species, and possibh' prove more

harmful than beneficial (Kennedy, 1975; O Connell,

Page 106

THE NAITILUS. Vol. 110, No. 3

1991 ). As a short-term solution to the immediate problem

of endangered species, the technique developed in this

stud> offers hope for propagating many species for which

hosts have not been identified.

ACKNOWLEDGEMENTS

Funding for this project was provided by the Upper

Mississippi Science Center of the National Biological Ser-

vice, and the Center for the Management, Litilization,

and Protection of Water Resources at Tennessee Tech-

nological University. We thank Michael J. Redding for

his suggestions and comments on the study, and the stu-

dents and staff of the Tennessee Cooperative Fishery

Research Unit for their assistance in the field.

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to malacologij.

NOV 1 7 1997

EDITOR-IN-CHIEF

Dr. M. G. Harasewych

Division of Mollusks

National MuscMiin of

Natural Histor\'

Smithsonian Institution

Washington, DC 20560

MANAGING EDITOR

Dr. Jose II. Leal

Bailey-Matthews Shell Museum

.3075 Sanibel-C^aptiva Road

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CONSULTING EDITORS

Dr. Riidiger Bieler

Department of Invertebrates

Field Museum of

Natural Histor\'

Chicago, IL 60605

Dr. Arthur E. Bogau

Freshwater Molluscan Research

36 Venus Way

Sewell, NJ 08080

Dr. Robert T. Dillon, Jr.

Department of Biolog\'

(College o( Charleston

Charleston, SC 29424

Dr. William K. Emerson

Department of Living Invertebrates

The American Museum of Natural

History

New York, NY 10024

Dr. Eileen H. Jokinen

Institute of Water Resources

Uni\('rsity of Connecticut

Storrs, CT 06269-40 IS

Mr. Richard I. Johnson

Department of Mollusks

Mnseiun of Comparative Zoology-

Harvard University

Cambridge, MA 02138

Dr. Aurele La Roctjue

Department of Geology

The Ohio State University'

Columbus, OH 4.3210

Dr. James H. McLean

Department of Malacology

Los Angeles County Museum of

Natural History

900 E.xposition Boule\ard

Los Angeles, CA 90007

Dr. Arthur S. Merrill

% Department of Mollusks

Museum of Comparative Zoology

Harvard University

Cambridge, MA 02138

Dr. Paula M. Mikkclsen

Department of Living Invertebrates

The American Museum of Natural

History

New York, NY 10024

Dr. Donald R. Moore

Dixasion of Marine Geology

and Geophysics

Rosenstiel School of Marine and

Atmospheric Science

University of Miami

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Miami. FL 33149

Dr. Gusta\' Paulay

Marine Laborator\'

University of Guam

Mangilao, Guam 96923

Mr. Richard E. Petit

RO. Box 30

North Myrtle Beach, SC 29582

Dr. Edward J. Petuch

Department of Geolog)'

Florida Atlantic University

Boca Raton, FL 33431

Dr. Garv Rosenberg

Department of Mollusks

The .\cademy of Natural Sciences

1900 Benjamin Franklin Park^xax

Philadelphia, PA 19103

Dr. David H. Stansber)'

Museum of Zoology

The Ohio State Universit)'

Columbus, OH 43210

Dr. Ruth D. Turner

Department of Mollusks

Museum of C]omparati\e Zoology

Haivard Univi'rsitv

Cambridge, MA 02138

Dr. Geerat J. Vermeij

Department of Geolog)'

University of California at Da\is

Davis. CA 95616

Dr. G. Thomas Watters

A(juatic EcologN' Laboratory

1314 Kimiear Road

Columbus, OH 43212-1194

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TH Et7NAUTI LUS

CONTENTS

Volume 110, Ntiinbcr 4

November 6, 1997

ISSN 0028-1344

Arnold G. Eversole Cianu'tom'iu'sis oi Mcrcciunia nicrcciiarid. M.

r(iiiii)ccliicn\i\ and Their Ihlirkls 107

James L. Theler The Modern Terrestrial Gastropod (Land Snail) P\iiina ol

\\'('stern Wisennsiii's Hill Prairies Ill

Edward J. Petuch A New (Gastropod Fauna From An Oliiiocene Back-Reef

Lagoonal Environment In West Central Florida 122

-i rt.jifc ■•.Jo.,c:r'OQrapr,,- Ir

NOV 1 7 1997

Notice

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THE NAUTILUS 110(4):107-11(), 1997

Page 107

Gametogenesis of Mcrcenaha mercenaria, M. ciinipechiensis and

Their Hybrids'

Arnold G. Eversole

Department of Aquaciilture, F'isiieries

and U'ilcllife

Clenison Uni\"ersit\

Clenison. SC: 29634-0362 USA

ABSTRACT

N(in-nati\t' stocks ot A/crcc/K/n't/ nwrtciiunn, M (V/i/i/iic/dc/isiv

and their reciprocal Inbrids were enltured in Sontli (Carolina.

Histolog;ic-al examination of the gonads revealed a himodal ga-

nietogenic pattern with a major spring spawiiing peak that con-

tinued into smnmer, followed by a smaller spawning peak in

fall. The spring spawning peak of A/, iiiciniinriii and its ma-

ternal Inhnd ocenrred one month earlier than that ot M. cam-

pccliicnsis and its maternal hybrid. The length of the spring

spawaiing peaks of the hybrids were hvo months shorter and

less intense than those obsen.ed for M. lueiTctuiria or M. aiiu-

prcliii'iisis.

Kfi/ aorils: M. iiu'rcriinriti, M. crimpeclufusis. gametogenesis.

h\brids. (juahogs. reprodncti\e C'\cle

INTRODUCTION

Lod.saiioll (1954) was the first to suceesslulK Inhridize

Mcnriuihii iiwiwitaiia {Linnueiis, 175S) and A/, ciiin-

pcchicnsis (C-inelin. 1791). Progeny of intra- and inter-

specific niatings were cultured in Virginia (Ha\"en & An-

drews, 1957), North Carolina (Chestnut el al . 1957) and

Florida (Menzel, 1964). In general, M ((unpci-lnciisis

grew the fastest, hvbrids were internietliate and A/, incr-

cctKiria was slowest, hut M. cainpccliieiisis and its hy-

brids had lower sunival. These early studies confirmed

that there were some aquacultnre lienefits ot M nur-

ccitiiria and M. cauipcchiensis Inhridization.

The distributions ol M iiwrcciunid and A/, canipt-

chii'H.sis mcrlap along the southeastern United States

(Abbott, 1974). Dillon and Manzi (1989) found that hy-

!)rids composed as nuich as 88*^ oi 179 clams collected

from Indian Ri\er, Florida. Soniewiiat lower hybridiza-

tion rate (31%) was obsened Irom a larger sample (n =

1,141) of clams collected from four sites in tlie Indian

River by Bert et ill (1993). In Hamlin Creek, South Car-

olina, pure A/, (-(uitpvchiensis were present at a irequen-

c\' of O.l'yf in large lieds of A/. iniTirnari/i with negligible

'Technical Contribution No. 4291 of the Siiutli tlarolina Ex-

periment .Station

Inliridization (Dillon, 1992). This suggested that repro-

ductixe isolation lietx\een the hvo species may be more

pronounced in Soutli Carolina than in Florida.

Dalton and Menzel (1983) docinnented the gameto-

genic cycle ol' soung male progeny from laboraton

crosses of A/, lucircii/iria and A/, raiupeclucniis cultm-ed

in .alligator Harbor, Florida. The bimodal spermatogenic

cvcle of the female A/, campecluensis X male M. mcr-

ccnaria progeny was similar to the A/, caiupcchiensis pa-

rental group, whereas, the spermatogenic pattern of the

reciprocal hxbrid was similar to M ineiveuaria parents.

Hesselnian li id. ( 1989) reported a bimodal gametogenic

cvcle for adult quahogs collected from Indian River,

Florida; howexcr, no differentiation of taxa was attempt-

ed in that stucK.

Information on the ganietogeuic cycle ol Mercriiiiriii

species is limited and there are no studies on adult (jua-

hogs of known pedigree. Also, nothing is known ol the

gametogenic cvcle of A/, ciinipcciiicnsis as far north as

South Carolina. The objectixe of this study xvas to com-

pare the gametogenic cvcle of adult clams of the species

M. lucrcciuiria. M. ciiinpn-liicnsis. and reciprocal cross-

es.

MATERIALS AND METHODS

Mcncniirid iiurciii/niii troui Millcn-d, Connecticut, and

M (Y/»i/)(y7»V/i.s(s IroTu Port St. joe. Florida, were

spawnecl in November 1985. Januan' 1986 and Februarx

1986 at Waddell Mariculture Center, South Carolina.

Crosses included: A/ incrrrniiiiii lemale X A/, inercc-

naria male (MM); A/, ciiiupcrhiensis lemale X A/, cam-

pcclticitsi.s male (CO; M iiurccnaiiii lemale x A/, cinn-

l>cclui'n.sis male (MC); and A/, cuntpechivmis female x

A/, nwrceiwria male (CM). Indixiduals from spawns and

crosses xvere held separately in recirculating sxstems im-

til the tjuahogs xx'ere mo\ed to a nurserx' at Folly Rixer,

South Carolina. Quahogs from the three spawnis were

combined for each cross on 5 May 1986 and maintained

in recirculating systems mitil planting (10 October

1986). Quahogs of each cross were planted in the Folly

River at about 960/m2 in separate protected trass. Ax-

Paw 108

THE NAUTILUS, \ol. 1 10, No. 4

eragc shell icii^tli.s (SD <il ])l.iiiliii<4 \\cr<' 10. .'5 nini, 17,2

mm, 15.fi nun and 15,0 nnn lor MM, ilC. \Kl and CM,

rt\s[H'fti\('K' (Manzi & Hadlcw mij)nl)l. data). Samples

were collected montliK Irom September 19S7 through

Ntnemher 1987 and [aunan 1988 through October

1988 and presened in lO'/f hnlTered lormalin. The qna-

hogs from the (XI tra\s were not sampled in Fehruan'

1987 and (juahogs Irom MM tra\s were not sampled in

October 1987 and Septembei' 1988 becanse of poor sur-

\i\al. A total of 113, 117. Ill and 78 qnahogs were sam-

pled Irom the CC, CM, MC and MM crosses, respi-c-

ti\el\.

(Jonailal tissue was dissected li'om (lie niublateral por-

tion ol the \iseenil mass, dehx'drated in an alcohol series,

cleared in xylene and embedded in paraplast. Sections

were cut at 8-10 |i,ni, stained with Harris' hematoxvlin

and counterstained with eosiu.

Slides tor each qiuiliog gonad were microscopicallv ex-

amined and gametogenic stages were identified accord-

ing to the descriptions of E\ersole ct al. (1980), Dalton

and Menzel (198.3), Ropes (1987) and Hesselman et al

(1989). Eight arliitran' stages ol male and lemali' gonail-

al dexelopnient were used: undillerentiated (lollicle wall

thicker than in later stages and lew, it any, gametogonia

near the periphen' of the lumen): early developing

(small oocytes and priniai'v and secondaiy spermatogonia

with a few spermatocytes found near tlie lollicle wall):

developing (larger oocytes adhering to the lollicle wall

b\ a peduncle and spermatocv+es and spermatids in the

lumen): acti\'elv de\'eloping (all stages ol gametogenesis

Irom spermatogonia to spermatozoa and oogonia to ma-

ture oocytes present); ripe (lumen filled with large ma-

ture oocytes or radiating bands ol s[)eriuatozoa); spawn-

ing (spermatozoa and large mature oocvtes still be abun-

dant in the Inmen, but gametogenic la\'er next to the

lollicle wall noticeabK reduced): partially spent (lollicle

w:ill contracted and a lew mature oocvtes and sper-

matocvte in the lumen): and spent/inactixc (lew ;ind of-

ten phagocytized gametocvtes in lumen suriouuded b\

a veiT thin lollicle wall). To huilitate eomp:irison among

the parental and hvbrid qnahogs, :i gonadal index was

calculated b\ assigning xalnes to each dexclopmental

stage. These xalues were undillerentiated = 0, earlv de-

veloping = 1, developing = 2, acti\eK <le\eloping = 3,

ripe = 4, spawning = 3, parti;ilK spent = 2, and spent/

inactive = 1. MontliK' gonadal index values wer(> com-

puted loi- e:ich pedigree b\ sumiiiing :ili the dexelop-

inental stage values lor male and lem:iie i|iuihogs, and

dividing this figure bv llie number ol i|uahogs in the

sam]ile. This procedure has been snccesslnllv uscmI In

others to sludv the gametogenic cvele ol M. lucrcouniri

in New York (Kassner & Malouf, 1982) and in (ieorgia

(Hel'fernan rl al.. 1989: Walker & Hefferuim, 1994).

These montliK gonadal index v:ilnes were then fitted i)V

spline inter])olations to follow liie g:mietogenie cvele

over aTi annual cvele.

KKSUITS

CJonadal index eunes lor progenv Irom each cross are

given in I'"ignre 1. These results indic:ite a bimodal I'a-

metogenie I'vele lor the progenv ol all lour crosses: how-

ever, the timing ol g:unetogenesis dillers among the loui'

progcTiv. .\ rapid decline in gouiulal index v:ilues indi-

cated :i spring S[)awning tluit contimied into summer lor

M cainpci-h'wnsis ((KJ) and its materiuil hvbrid (('M).

Average gonadal index values lor (X," and (^M (juahogs

were 3.62 and 2.28 in April, and 0.50 in August and 0.22

in [nne, respectivelv. A second smaller spawning peak

Wiis obsened in the fall when average gonadal index val-

ues lor (X: and CM decreased li'om November (3.50

and 1.30) to |;iuuan/Februan- (2.50 and 0.fS7).

The pattern ol gonadal index values exhibited bv A/.

1 1 writ' Italia (MM) and its maternal hvbrid (MC) also in-

dicated a decrease in spring, but roughlv one month ear-

lier than CC, and CM (|ualiogs. .Average g()n;idal index

values for MM and MC decri'ased Irom M;n'cli (2.94 and

3.25) through August (0.64) and |nne t 1.30), respective-

lv. The decrease in gonadal index values obsened Irom

September through November also indiciited a l:iil

spawning. Although the trend of gametogenesis in the

fall for MM and MC was not as clear cut as that of (X,'

and CM, it appears that spawning occurred 1 to 2

months earlier in MM and MC.

The patterns in gonadal index values loi' tlie hvbrids

:ip[)e:n' to tr:ick tluit ol the m:iteiiKil parent, but at re-

dncetl levels and for shorter periods. For ex:nnple, the

decreases in gonackil index values lor spring and tail

sp:ivvning periods were 3.12 and 1.00 lor C(>' compared

to 2.06 :ni(l 0.63 for CM (|uahogs and 2.30 and 1.57 lor

MM comp;n'e(l to 1.95 and 0.50 lor the M(,' hvbrid, re-

spectively. The lengths ol spring spawning periods (i.(>.,

tim(> of declining gouatlal index values) were approxi-

mately 5 mouths lor (X" ;uk1 MM comp:ned to .'■) montlis

lor CM and MC (juahogs.

DISCUSSION

The sjiring domin;int bimodal ganietogenii- cvele ob-

seiAed in this study has been reported loi- Wciri-iuniii

t:L\a in Noith ( iiU'olina (Porter, 1964). South (Carolina

lEver.solee/ f//., 1980: M;mzi <>/ r;/., 1985), (;eorgia ( llef-

fernan .■/ ai. 1989: Walker & Heffern:m. 1994) and Flor-

ida (Diilton & Menzel, 1983: Hes,selm:ui rl al . 19891. l,,

contrast to Diiltou :uul Menzel's (1983) findings, tlie tim-

ing of the sp;iwning pe;iks for M iiicrcciiaria. M. cam-

jiciliiciisis and the recipicnal hvbrids Wiis not the s:ime;

the spring sp:iwiiiiig pe.ik ol .\/ inirccnaria :ind its nui-

teru:il hvbrid oecniied one month earlier than IIkiI ol

M. lainpccliirii.sis :ind its maternal hvbrid. n:ilt()n :nid

McMizel's (1983) lailnre to detect a dillerenee betxveen

the two Mrrrciiaria species :iud e.ich ol the spe( ies ma-

lernal hvbiid m.iv have resulted in [lart li'om using siiuill,

voimg. and mostlv 111. ile qiKihogs lor histi liogK'.il evalu-

ation. Mcrciiiaiia undergo a juvenile male sexual ph:ise

(I,oos:molf. 1937) before :icliieving sexiuil maturitv (I'a-

ersole ,7 al . 19801. Sex nitio (1.0 : 1.1, M : F) :uid av-

erage SL (40.5 + 6.4 nun. SD) indicate that tlu' sanijiled

qiiahogs had com|)lete(l the juvenile nuile phase (F\er-

si>\v (I al. 1980; Knaiib \- Fversole 1988).

A. G. Ex

1997

Fa;

109

X

0)

■D

_C

■o

CO

c.

o

O

3.5

2.5

1.5

0.5

3.5

2.5

- 2

- 1.5

- 0.5

10

12

2 4

Month

8

10

Figure 1. Me

ctiinpcclticnsis

livbiid (CM).

an gdiiLulal inik'\ with oiu' staiidaixl crnir liar fur crosses of M iitci'iriiiirid friiialc ■ ,\/ inircnunid male (MM), A/.

female X A/, camprchiiiisis male (CC). A/, inercciitnia female ■' A/ rdinpcrhiiiisis male (MCJ) and the reciprocal

Line fitted b\' spline inteq«)lation.

Hcsselmaii ct a]. (1989) encoiiiitc^n'd eoiisidcralilc

variations in tin- f^anieto<4eiiic cxiAv ainont; samples of

(jiiali(iij;s eolli'ctecl li'diii Iiuliaii Rixcr. Florida. Clixfii that

liotli species of Mercenaria occih' in Indiaii Ri\cr, Hfs-

selnian et nl. (19S9) speculated one source ol the \ari-

alion coiikl he related to genetic ditlei'ences in repiii-

dncti\c acti\it\- among the taxa. Since then, i5eit el nl

(1993) \'erifii'd tiiat Indiaii River contained hotli Mcr-

ceiKiria species and their hvhrids. Ditficiilts associated

with defining a ganietcjgenic cvcle wonld he expected il

tlie reproduction cvcle is imder some genetic coiitiol

and if closely-related species coexist and liNhridization

occurs. Currently, there is exidenct' that dillereiit stocks

of A/, nwrccnarid and Cnissostrca lin^inica (Cimelin.

1791) exhibit geiieticallv distinct cxries o( gametogenesis

aiid spawning when cultured in a common enxironment

(Knauh & ENersole. 198S; Barber c/ ,il . 1991). Bai-ber

and his colleagues (1991) suggested th.it osster stocks

had geneticalK' different emii-oinnental re(|nirenients

necessarx' lor initiating gametogenesis and excntualK' lor

cuing spawning.

nillcin and Manzi (19S9) sugg<'sted that spawning

time and habitat separation could he possible mecha-

nisms inaintaininsi the genetic inteirritN' ol the two Mer-

cenaria species. In the Indian Ri\ei', Floi'ida, where Mer-

crnaria coexist, S.S7f and llWc of 179 and 1,141 collected

(juahogs were natnralK occurring hxbrids, respecti\t'l\'

(Dillon & Manzi, 19S9'; Bert et at.. 1993), Farther north

in Hamlin CJreek, South Carolina, oulv about 0,1% of

tfiousands oi tjuahogs examined were M. antipecluensis.

and Inbrids wei'e i-arer still (Dillon, 1992). Although the

timing ol the gametogi'uic cvcles ol introduced stocks of

A/, mercenaria and M. cantpecltiensis differ in the

spi'ing, sufficient o\erlap inav exist for hvbi'idization of

native Mercenaria in South C'arolina. The fact that Mer-

cenaria hybrids were encountered less frequently in

South Carolina than in Florida indicates that ,spatial sep-

aration or somi- other fact may he luore important for

I'epioductixe isolation than the tenipoi'al diffei'ences in

spawning. It is interesting that the M. campechiew>is and

h\bi-ids identified bv Dillon (1992) were collected from

a latlier uniijue South Cai'oiina habitat, one character-

ized by ocean-dei'i\ed high-saliiie water, firm substrate

covei'ed by deep water and a comnimiitx' of stenohaline

biota (e,g,, ('.liinne cancellala Fimiaeus, 1767). A suncx'

of M. mercenaria habitat in South Carolina rexealed that

most ol the areas with clams were mud and sand covered

In sh.illow mixohaline waters (Anderson et iiL. 197S). It

Page 110

THE NAUTILUS, \ol. 110, No. 4

is .uiticipatfd tliat Miriiiitiriii li\ lnidizatioii will hccome

a rarer exent in higlici' l.ititudcs ol tlic Atlantic coa.st as

contignous habitats nicctinii; the sp( 'tics-specific require-

ments ol both species become less available.

Earlv stndies indicated a growth advantage ol M nwr-

cenaria and M. campccliiciisis hxbridization (Haven &

Andrews, 1957; Chestnut c/ tiL. 1957; Menzel, 1964)

possiblv because tlie hybrids Ivdvc a shorter and less in-

tense spawning period than tlie hvo species. However,

Eversole and Heffernan (1995) also obsened that go-

nadal neoplasia occurred more frequently and in a more

advanced stage in tlie hvbrids tlian in M. mercenaria and

A/, nnnpcchiensi.s. ,\(juacultnrists need to be aware of

both the favorable and unlavorable attributes associated

with liybridization [irograms when trxing to improve

(juahog performance.

ACKNOWLEDGMENTS

The author thanks Nancv II. Hadlev and her fellow

workers lor the clam husl)andn. Special thanks go to

John Hudson who assisti'tl with the dissection, Yvonne

Bobo for help with histological preparation and Chris

Kempton helped with the illustration. Connnents by

Hob Dillon and Randv' Walker on an earlier dralt greatly

improved this manuscript. The research was supported

by S.C. Sea Grant Consortium and S.C^ Agricultural Ex-

periment Station.

LITERATURE CITED

Abbott, R. T. 1997. Aiiiericaii Scasliclls, 2ik1 cd Win Nostrand

Rlieinliold, New York.

Anderson, \V. D., W. ]. Keith, F. H. Mills, M. E. Bailey & J

I,. Steinuiever. 1978. A snnev of South Carolina hard

clam resources. South Carolina Marine Resources Center.

Technical Report Number .'32, Charleston, South Carolina.

Barber, B. J., S. E. Ford & K. N. Wargo. I99I. Genetic vari-

ation in the timing ol gonadal maturation and spavvaiing

ol the eastern oyster, ('mssostrcti r/rg/ii/rr; (Cmelin). Bi-

ological Bnllctiii' 1SI:2I(>-22I.

Bert, T. M.. D. M. llcssclnian, VV. S. .\rnold, W. S. Moore, II.

Cruz-Lopez & D. (;. Marelli. 199.3. High Irequencv ol

gonadal neoplasia in a hard clam (Mcnciuiiia spp.) hybrid

zone. Marine Biologs 117:97-104.

Chestmit, .\. F, W^. E. Fahv ^■ 11 ] I'orter. 1957. Growth ol

vonng Wntis mcnrniirm \iiiiis r{iiiii>crhicii'<is. and their

livhrids. Proceedings ol the National Sliellfislieries As.so-

ciation 47:50-.56.

Dalton, H. & VV. Menzel. 19S.'?. Seasonal gonadal changes ol

young lahoratorv-spavvned .southern (A/, (•ainpcchii'iisis)

and northern (A/, nwrcfnnriii} (jualiogs and their recip-

rocal hybrids in northwest Florida. Jomnal ol Shellfish Re-

search'3:1 1-18.

Dillon, R. T. 1992. Minimal hvhridizalion between populations

ol the hard clams, MciTi-niirid imrccndria anti Mcncnaiia

(■(iiiipccliicnsis. co-occurring in South Carolina, Bulletin ol

Marine Science 50:411—416.

Dillon. R. T. 6c J. J. Manzi. 1989. Genetics and shell nior-

phologv in a hybrid zone between the hard clams, Mer-

cenaria mercenaria and M. canipechiensis. Marine Biologv

100:217-222.

Eversole, A. G. & R B. Heffernan. 1995. Gonadal neoplasia in

northern Mercenaria mercenaria (Linnaeus, 1758) and

southern M. campechien.sis (Gmelin, 1791 l (|uahogs and

their hybrids cultured in South Carolina, jmniial ol Shell-

fish Research 14:3.3-39.

Eversole, A. G., W. K. .Michener & R J. Eklridge. 1980. Re-

productive cycle of Mercenaria mercenaria in a South

Carolina estuary. Proceedings of the National Sliellfisli-

eries Association 70:20-.30.

Haven, D, 6c J. D. Andrews. 1957. Sunival and growth ol Ve-

nus merceimria, Venus campecltiensis. and their hvhritis

in suspended travs and natural bottoms. Proceedings ol

tlie National Shellfisheries Association 47:43-49.

Heffernan, R B., R. L. Walker 6f J. L. Carr. 1989. Gametogenic

cvcles of three bivalves in Wassavv Soimd, Georgia: I. Mer-

cenaria mercenaria (Linnaeus, 1758). Journal ol Shellfish

Research 8:51-60.

Hesselman, D. M., B. J, Barber 6c N. J. Blake. 1989. The

reproductive cycle of adult hard clams, Mercenaria spp.

in the Indian River Lagoon, Florida. Journal ol Shellfish

Research 8:4.3-49.

Kassner, J. 6^ R. E. Maloul. 1982. ,\n evaluation ol spawner

transplants' as a management tool in Long Island's hard

clam fisherv'. Journal of Shellfish Research 2:16.5-172.

Knauh, R. S. 6c A. G. Eversole. 1988. Reproduction of different

stocks oiMercenaria nwrcenaria. Journal ol Shellfish Re-

search 7:.371-.'376.

Loosanoff \'. L. 19.'37. DevelopiiUMit ol the priiiiaiv gonad and

sexual phases in Venus nu reenaria Limiaeiis. Biological

Bulletin 72:.'589-405.

Loosanolf V. L. 19.54. New advances in the stuilv ol bivalve

lanae. American Scientist 43:607-624,

Manzi, J. J„ M, Y Bobo & V. G. Bunell, Jr. 1985. Gameto-

genesis in a hard claui, Mercenaria mercenaria, population

in North Santee Bay, South Carolina. The \'eliger 28:lS(i-

194.

Menzel, H. W. 1964. Seasonal growth ol nortlieiii .uid soiitliern

i|ualiogs, Mercenaria mercenaria and M. eainpecliiensis.

and their hybrids in Florida. Proceedings ot tlii' National

Sliellllslieries Association .53:111-119.

I'oilei, II. 1964. Seasonal gonadal changes ol ailnlt clams, Mer-

ciiiiiria mercenaria (L,) in North Carolina, Proceedings of

the National Shellfisheries As-sociation .55:3.5-.52.

Ropes, j, 1987. Age and growth, reproductive cv'cle, and his-

tological tests for heavA' metals in hard clams, Merceniiri<i

mercenaria. from Raritan Bav 1974-75. F'isherv Bulletin

85:65:5-662.

Walker. R, I„ 6v P B. Heffernan. 1994. Temporal and sjiatial

elleets ol lid.il evposure on the gaiiietogenic cvcle ol the

iioillieiii c|ii.ilioi;, Mrreenand nurceninia ( Liiiii.ieus.

1758), 111 eo.islal ( a'Oigia. |oiiiii.il ol Slielllisli Hesearch

1:5:479-486.

THE NAUTILUS 110(4):1 1 1-121, 1997

Faer 1 1 1

The Modern Terrestrial Gastropod (Land Snail) Fauna of

Western Wisconsin's Hill Prairies

James L. Thcler

Department of Sociol(>i;;\ ami

Arcliaeolog\'

llni\ersit\' of Wisconsin-La Crosse

La C^rosse. Wisconsin 54601

ABSTRACT

This pajx-r describes the first snncv and (|uantified unaKsis ol

the terrestrial gastropod fauna associated with \eric "hill prai-

rie" and related dr\ habitats in western W'isconsin's Driftless

Area. A region of 35,000 km- in the Midwestern U,S.A.. the

Driftless Area was surrounded on three sides, but ne\er co\-

ered b)' late Pleistocene glacial ice. In all, 44 separate \egeta-

tion detritus samples, each with a xohune of 0.5 to 2.0 liters,

were collected in nine western Wisconsin counties. These .sam-

ples contained 10,900 gastropods, with 29 ta\a represented.

The sampled hill prairies were found to ha\c localized, high-

densit\' gastropod populations characterized bv a small mmiber

of abimdant t;L\a. One species, Qaatrocoptii proccra (Gould.

1840) is currently listed as Threatened in Wisconsin. This spe-

cies occurs at disjunct prairie habitats along the dissected \alle\

of the Mississippi Ri\er that mav represent a relict distribution

for this species

Ai'i/ words: Laiifl snails. biot;eograpli\, Midwestern L!.,S..\.

INTRODUCTION

This report describes tile results oi a liascliue sunev to

assess the living terrestrial gastropod lauiia assnciatt'd

with the xeric "iiil!" prairies (Curtis, 1959:275-276) ol

Wisconsin's Driftless Area. The studv provides the first

.systematic assessment of gastropod abuudaui-e and spe-

cies diversitv' at lull prairie and associated dn loi-ations

in nine western Wisconsin counties. From the stand-

point of terrestrial gastropod distril)ution, this area is one

of the least known regions in eastern North America

(Hubricht, I9S5:2), although some researeli lias been

undertaken in the past (I. P. E. Morrison, 1929; Levi &

Levi, 19.50).

The phvsical setting of western Wisconsin is varied

and complex. The heavily dissected hill coimtn- of this

region has been assigned to the Western Upland phys-

iographic province iiv Martin ( 19fi5:42-43). Southwest-

ern Wisconsin is within the Driftless Area (Martin, 1965:

82-8.3), located primarily within the Western Upland,

with smaller portions extending into the adjacent states

of Iowa, Illinois, and Minnesota. The Driltless .Area, as

its name implies, lacks evidence ol glaciation during the

later Plei.stocene (MickeLson it r;/. 1982:155-169), and

covers an area of 35, ()()() km-, about 75% of which is in

southwestern Wisconsin (Roosa, 1984:43). The Driftless

Area's topography is characterized by steep-sided, stony

valleys dissecting the uplands with dendritic patterns of

small stream development. The narrow upland ridges

and escaqinients of the larger valleys e.xhibit up to L50

ni of relief. The two prominent rivers crossing the region

are the Mississippi, which lornis the western boundan'

of Wisconsin, and the Wisconsin River, which drains

much of central and southwestern Wisconsin.

Western Wisconsin is situated between the tall-grass

prairies to the west and the deciduous forest to the east.

The southw-estern portion of the state has been assigned

to the "prairie-forest province" King to the south of the

Horistic tension zone of Curtis (1959:15) and is charac-

terized by its mosaic of vegetation communities. The

structure and position of tliese connnunities have re-

sulted from a variety of factors, including topography,

climate, peiiodic biu-ning of prairies, and underlying

soils and bedrock. At the time oi Enro-.'Vnierican settle-

ment, four distinct vegetation connnunities were pre-

dominant: prairie, oak savanna, southern oak forest, and

southern mesic forest. Tlie most extensive prairie com-

munitv was the mesic prairie tvpe, dominated by grasses

such as big bluesteni {Aiulropo^on gcrardi). Indian grass

{Soriihristnini iiut/ms). and forbs. Prairies bv definition

support fevvi'i- tli;m one tree per 0.4 ha. In western Wis-

consin, mesic prairii'S iormed large tracts on gentiv roll-

ing uplanils, particularlv south ol the W'isconsin River in

Iowa, CTrant, and Lafayette counties. Nineteenth-centun,-

travelers who journeyed from forested regions in eastern

North America often described the extensive tall-grass

prairies they first encountered in western Wisconsin.

One of the best-known mesic prairies was the Militarv'

Ridge Prairie, which ran Irom the town of N'erona in

Dane County, to the northwestern corner of Cwant

Countv- (Curtis, f 959:262-264).

Xeric prairie coTimumities in the Driftless Area were

locally conspicuous but spatially circumscribed (figures

I, 2). These xeric hill or "goat" prairies occupied steep

south- f)r southwest-facing slopes and made up only a

Pa2e 112

THE NAUTILUS, Vol. 110. No. 4

J^

I \

Figure 1. Brack's Bliitt liill prairie in Trempealeau Couuh', Wisconsin. Arrow points to the saniplini^ location (l'\\.\-3()2(ii shown

in figure 2. \'iew is toward the north, with the Mississippi River in the foreground.

.small traction ot tlie Diiltlcs.s Areas vc^ctatioiial laiul-

scapc. Hill prairies dexelopi'tl (jii the tliiii soils o\erKing

(loloiiiite outcrops with exce.ssive ruuoH. These locales

lia\'e a nniforiiilv high calcium couteut and a pH of S.O

or more (CJurtis, 1959:275-276). The four most common

plant species on these hill prairies, in order of abun-

dance, are little bluestem {:\iitln)j)<i<i<i}i sc(ij)iiiiiis). side-

oats gania grass (Boulclona /■iiiliixiiiliilii), big bluestem,

and silky aster (Asler scrirciis). (hiMall, Asteraceae make

up 27.5%, Poaceae 13.7%. Kabaceae 5.3%', Rosaceae

4.6%, and Asclepiadaceae 4.6% ol the hill pi.iiiie com-

numity (Curtis, 1959:2691.

Tlie upland mesic prairie comnnmities in western

Wisconsin originated during the mid-Holoeene warm/

dr)' .Mlantic climatic episode (Bartleiu & Webb, 19S2:

80; Maher, 1982:1.32). In later lloloeene times, most of

the region's prairie stands were maintained b\ freijuent

burning, which is well documented l)\ both historic ac-

counts (.Anderson, 1954:15-22; Martin, 1965) and the

presence of charcoal in regional pollen profiles (Davis,

1977:204-213). .Soils that formed under mesic prairie

conununities, partieularK Tama and Dodge\ille silt

loams (Hole. 1976:140, i'73, Plate 7) are widely distrib-

uted along the upland ridges of western Wisconsin. The

mesic prairies are gone from western Wisconsin todav.

replaced bv agricultural fields.

MATERIALS AND METHODS

In 1979 the autlioi- collected two vegetation detiitus

samples from ,i hill prairie in Crawford Couut\, Wiscon-

sin. These s.nupies contained 16 gastropod ta\a, includ-

ing one species not prexionsK' recorck'd in Wisconsin,

Gastroropdi proccni (Ciould. IS40>. Gtistnu-oplii procira

is a species associated with prairie/savanna vegetation

connmmities in regions with a frost-free season of 160

ckivs or moie (Reigk', 1963; Baerreis, 1980:108-109). In

1985 two addition. il hill prairie vegetation detritus sam-

ples were collected in La Oosse Countv. Wisi-onsin.

One of these sam|)les .ilso contained G proccrii. Per-

sonnel at the Wisconsin Chapter of The Nature Cou-

senancv' (TNC) and the Wisconsin Department ol Nat-

ural Resources (WDNR) showed interest in de\elo|)iug

aTi iu\entor\ of hill prairie gastro[)od connnmnlies, lead-

ing the autlioi to (olleit and analvze 10 .uldilioii.ii s.im-

ples from western Wisconsin hill pi.iiii<'s and rel.ited

habitats during 19.S6 and 1987.

Hill pniiries sampled hn gastropods nicliided .i \,uiet\

J. L. Theler, 1997

Paw 113

Figure 2. BraiK's BIiitT hill prairir in rreiiipeiileaii Coimtv, Wisconsin. Sampling location l!\\A-o()2H is at tlic iipsiope side ot the

ilftachcd lK]nkler. Photo looks west, with the Mississippi Ri\er on the lett.

of TNC, WDNR, antl pri\atel\ (iwiicd \cric scttiiiy;s in

nine western Wisconsin counties, eight of uliicli l)order

the Mississippi Ri\er; Ironi soutli to nortli, thi'\' are

Grant. Lafayette, CJrawfonl. N'crnon, La (^ros.se, Trem-

pealeau, Buffalo, Pepin, and Fierce (table 1). Whenex'er

possible, TNC anil WDNR propertii'S were emphasized,

under written agreement with those agencies. In all, 35

samples were taken and anaKzed at 19 separate hill prai-

rie settings (figure 3).

Specific sampling locations were selected 1)\ searching

each hill prairie to identii\' areas with living gastropods

or fresh shells. The most prodiicti\-e locations occurred

where relativeh dense stands or clinnps of prairie grass-

es aiiutted isolated bedrock exposures, forming "pock-

ets" that would hold accunuilations of \egetation detri-

tus, .Althougli some of the sampled hill prairies (e.g..

Hush (Ireek South in X'ernon Countv) coxcr st-M-ral

acres, the gastropods were primariK limited to discrete

concentrations within the larger areas. Tlii'se localizt'd

gastropod populations, often restricted to less than a

sfjuare meter, seem to represent small, isolated com-

mmiitii's within the hill prairie setting. Clixen this distri-

bution pattern, random sampling would liaxc resulted in

the recoveiT of fewer species and lower individual snails,

in addition to the hill prairie settings, nine sampU-s

were taken in four counties (Cirant, Lafavette. CJrawford.

and N'ernon) from what were consitlered to be "iiigh-

stress" habitats for gastropods (table 2). The habitats tiiat

are referred to as high-stress appeared to represent the

driest locations that could be expected to support lixiiig

gastropod commmiities. These locations geueralK lacked

xeric prairie vegetation and consisted of a thin layer (<2

cm) of organic detritus on dolomite bedrock exposm^es.

The\' were olteu adjacent to some canop\ co\er from

wood\- \egetation (birch, cedar, or oak), but were otli-

en\ise exposed to the extremes of Wisconsin's climate.

One to four samples, eatli with a xolume of 0.5 to 2.0

liters, were taken at each hill prairie or high-stress hal)-

itat. Each sample consisted of vegetation detritus and

approximateK' 1.0 cm of underKing soil taken among

prairie plants adjacent to or against rock exposures. Sam-

pled locations were marked on IISGS topographic maps

and usnalK photograpiied. Each sample was measured

In solnme in the laboratorx' and then water-screened,

with the gastropods aiul other material retained in geo-

logic siex'es (T\'ler #40. mesh size 0.425 mm). The

screen-captured residue was then dried and .systemati-

cally se-arched for gastropods under a low-power (lOX)

binocular microscope. .^11 gastropod shells and poten-

tialK identifiable shell hai'inents were remo\ed from the

Page 114

THE NAUTILU.S, Vol. 110. No. 4

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Page 116

THE NAUTILUS, \\>\. 1 10. No. 4

Table 3. (;;istn)|)iKl Li.s.s('inl)l;ii.ii'S troni wcstcni Wisconsin liill ])rairies.

Lula-

Crant vette C'lawtorcl X'enion

Counh : J

Dew- M- Lcitner Battle

ev Ziin- Mo- (Jran (Jrac Hollow Rush Oeek BluH X'icton

Sariiplf location; Hts. nier untl

UVVA:3051 3014 3040 2547 2576 3055305fS 303530363058 30433053 296229633037

Taxon: Sample volume (1.): 1.1 0.75 1 NR NR O.S 0.7 0.8 0.8 1 2 1.8 1 1 0.75

Stcnotrema harlwtiiin (G.H. Clapp, 1904) 2 — — — — — — — — — — — — — —

Mesodon claiisns (Say, 1821) 1 — — — — — — — — — — — — — —

Euconiilws /uIdus (Mullen 1774) — — — — — — — — — — — — — — —

Ghipluinlinia hulcntnta (Say, 1823) 2 2 6 _ 3 — — 7 14 ^ 60 7 1—2

Scscnitrca ricciriiia (Gould, 1841) — — — — — — — — — — — — — — —

Hawaua minnsada (Binney 1840) — 7 — 2 4 2 14 1 1 1 7 12 3 6 9 3

'Ainitoidi-s nrhorciis {Sa\, 1816) — — — — 13 — 2 2 S — 5 — — 1 1

Sirifiturii iiiiliuiii (E,S. Morse, 1859) — — — — — — — — — — — — — — —

Striiiturii sp. — — — — — — — — — — — — — — —

Deroccras laeve (Miiller, 1774) — — — — — — — — — — — 1 — — —

Anugimpira (dtenmta (Say, 1816) — — IS — — — — — — — — — — — —

Hrl tod isms pamlMus (Sav, 1817) 9 — — 4 35 1 29 — 16 8 79 26 — 3 20

Hclicodisrus sin^lciinnus (Pil.shn. 1890) 2 — — 1 — — 5 — 23 4 4 36 3 — 12

Piiiictuiii iiiintifissiiiiiiiii (I. Lea, 1841) 4 6 — — 2 — — — 6 1 8 1 10 4 5

Snccineidae 3 — 2 — 1 — 2 1 1 — — — — — —

Strohdops lahtp-iiithira (Sa\-, 1817) 9 6 — 9 14 4 2 13 45 2S — — 3 3 2

Sirohdops (iffiitis Pil.shiT, 1893 6 8 — ____________

Strohdops sp. ■ _ 17 _ _ _ _ :3 1 42 5 — — 2 1 —

Gastrocopta amiifcra (Say, 1821) 11 32 IS .39 27 12 98 2 3 59 34 11 4—2

Gastrocopta contrmia (Sav, 1822) 5 2 6 1 — — 5 2 16 — — 1 2 11

Gastroroptti Iwlzinocri (Sterki, 1889) 39 9 5 101 14—2 13 27 34 30 33 69 37 53

Gdstrocopia pcntodon (Say, 1821) 7 8 — 7 29 — — — 9 13 23 12 16 21 3

Gdstrocoptii corticaria (Say, 1816) 3 — 1 2 1 — 3 — 1 — — 1 — — —

Gastrocopta proccra (Gcnild, 1840) 10 13 — 2 5 14 4 3 7 1 21 12 8 — 10

I'lipoidrs albdahris (C.B. Adams, 1841) 4 10 2 — — 13 22 — 6 4 51 9 — — —

('oliimella cndcittida iDrapamaud, 1805) — 1 — — — — — — — — — — — — —

Valloiiia pidchclla (Muller, 1774) — — — — — — — — — — — — — — —

Valloiiia costata (Muller, 1774) — — — — — — — — — — — — — — —

Vallouia panula Sterk-i, 1893 _ _ _ 75 14 2 29 2 5 27 51 33 1 6 10

Valloiiia pcrspcctiia Sterki, 1892 1 — — 1 — — — — — — — — — — —

Vallouia sp, 20 — 7 108 15 7 53 1 6 54 68 64 5 3 13

('ochlicopa Inhrica f Muller. 1774) — — — — — — — — — — — — — — —

Juwnilcs ,30 15 4 NR NR 20 28 5 24 35 58 33 56 46 61

NR = Not recordeil.

(Ictritus and held lor identification. Spceinicn.s were later

identified, fatalo<led, and stored in tflass vials with iden-

tifieation/provenance laliels. The nonienelatnre used in

tiiis paper follows Turneon <■/ al. (I9.S8). Voucher spec-

imens ol each t;L\on collected were deposited in the

Field Mnsenni of Natural iiisloa-.

HESLLTS

The 44 samples taken duriiii; this suncN contained

l(),9()() ga,strop()ds or gastropod shells representing 29

species. Of these, the 35 hill prairie samples contributed

8,664 gastropods with all 29 taxa represented (table 3).

The three most connnon and widespread specic^s found

at the hill prairie sites were C,aslroa>])t(i armifcra (Say,

IS21), Gastrocopta hohJuocri (Sterki, 1889), and Gns-

tfocoptfi })ciito(loii (Sav, 1821). Each of these species

contributed more than 11% of the total number of snails

and was present in 30 or more ol the 35 samples: iwn

ol the three were represented in all nine counties sam-

pleil (table 4). Each ol the next 12 most .ibnndant t,L\a

represented 1.3'^ to S.T'/r ol the total number ol snails,

occurred in 14 to 2S ol the 35 samples, and was lound

in at least six ol the counties suncved. Together, these

15 most abundant taxa Tuack' up oxer 96*^ ol the hill

prairie gastrojiods. The niajoritx of the snails rejiresent-

ed are taxa that UpicalK occur in prairie nv sav.uina hab-

itats. The rem, lining 14 species made up less than 4'"'f

of the gastiopods, and most occmri'd in rel,iti\el\ lew

sam|)ies.

The nine sam|)les taken Irom high-sticss h.ibitats con-

tained 2.236 gastropods of 20 taxa it.ible 5). All of lliese

J. L. Theler. 1997

Page 117

Tabli- 3. Kxtciulrd.

Iniii-

pe-

La Crosse aleaii Pepin Pieree

7. ■ i n Buffalo 77. 777, , , ,

hA]ieniiieiital Bra- hi\e-.\lile Maid-

Fanii Hixtiiii Forest Bin! (l\-'s Landfill Rd. Stock- l^rairie Fk Pepin en Rusli Hajj;er(:it\

Bliitf Bluff liolm Rock Riv. ^ ~

29S7 3005 :50()fi 3029 3030 3042 3057 2V}W :)020 3024 3025 3023 3031 3()4f 3()4S 3049 3047 3046 3044 3045

1 1,5 1 1 f.2 f,f O.S OS I 1 1 1.5 1,1 f 1.5 ().(i5 1 1.7 1.1 0.6

_______ __ !________ — _

___ 2— 4— — 1 4 1 — 1—— !____

— 5 2 11 22 3 1 4 5 12 — 3 5 15 11 11 3 20 — —

_______ ______ !______

_ _ 1 27 1 3 — — 1 — 2 5 — 4 89 36 27 42 6 43

13 1 5 2 1 — — 2 20 1 — — — 5 3 1 — — 2

_______ ____________ 4

_______ !____________

_______ _____!_______

— S 12 61 5 6 — — 19 41 5 4 41 9 40 4 10 5 1 1

9 — 2 22 20 23 23 — 12 36 — — 31 20 13 24 23 14 4 —

_ _ _ 36 1 14 ] — 3 4 — 1 — — — 2 3—12

1—1 2 2 — — — — 1 1 4 10 23 24 14 5 2 — 12

6 — — 15 4 — — — — 5 — 1 59 31 55 101 — 1 3 13

_ _ 29 5.S 4 141 11 5 15 11 16 42 1 4 13 7 13 80 56 97

1 _ 1 3 3 7 3 _ _ IS — 1 17 .33 6 7 4 16 1 1

;54 2 63 — 24 3 11 33 5 — 10 31 — — 19 45 23 53 27 4

23 5 13 79 25 9 12 1 U 4S — 13 51 23 76 53 11 41 22 22

_____ 7- ___2— — — — — 8 — — —

1 _ 10 14 — 1 5 _ 14 _ 1 _______ 6 3

_ _ 22 28 — 9 — 1 6 2 12 13 3 — 9 12 3 45 15 10

_______ fi____________

— — ——— — 1 71 3 — 19 32 — — 9 19 8 60 12 —

_______ _________ i__4

_ _ _ _ _ 54 — 480 6 — 42 43 — — 10 19 — .54 41 21

_______ ______ 5______

41 4 53 83 10 102 44 78 15 55 21 48 31 16 31 77 21 79 45 44

tiLxa were also represented at hill prairie sites; however,

tiie rank order of the species was different in tlie liigli-

stress samples. The first- and second-ranked species at

hill prairies, Gastrocopta amiifern and Gastwcopta liol-

zinaeri, ranked second and fifth in the hiifji-stress sam-

ples and contrihnted 3S.5% of all snails (talile 6). Three

other species, Gastrocopta roiiicaria (Sax; ISlfi), Valloii-

la perspectiva Sterki, 1S92, and Vallonia cii.\tala (Miiller.

1774) contributed 52.5% of the gastropotls in the lii'j;h-

stress samples, ranking first, thiril, and fourth respec-

tively, wliilc ranking sixteenth, nineteenth, and twentx-

first in tile hill prairie samples. Together, these fixe most

ah\mdant forms represented 91.19?- of the species re-

coxered in the high-stress habitats. The difference in

species composition between the assemblages probablx

reflects the differences between the sampled habitats.

DISCUSSION

The hill prairies of xx'estern Wisconsin host a fairly di-

xerse connnuuitx' of terrestrial gastropods, with some 15

species being relatixelx' abundant and xxddespread. A

comparison of these sunex' results xxith Hubricht's

(1985) countx' listings for eastern North America re-

xealed that the hill prairie surxey contributed 1.30 new

countx^ records, and eiglit species that had not been re-

corded prexiously in Wisconsin (table 4).

One taxon xx'orthy of special comment, Gastrocopta

proccra. is currently listed in Wisconsin as a state

Threatened species. Gastrocopta procera finds its pre-

ferred habitat in prairie and sax'anna settings. This spe-

cies does not lixe in the closed-canopy moist xx'oodlands

of xx-estern Wisconsin (Theler, 1992). The nortliernmost

Page 118

THE NAUTILUS. \ol. 110, No. 4

LANDFORMS OF WISCONSIN

Gaological and Natural History Survay

Q*or9a F Hsnkon. DirvcWf and Sut* Oaoloaist

uwiviaairf cxtcnsion. uKiviasmr or Wisconsin

1971

Hager Cit

Five-Mile Prair

Rush Riv

Maiden Ro

Lk. Pepi

jtoc'kho

Landfill R

Brady's Blu

Bird Blu

Hixon Fore

Experimental Far

Cedar Valle

Vicfo

Battle Blu

Rush Cre

Leifner Hollo

Gran Gra

Zimme'

Dewey Hts. -*'

M-Mound

KiHiirc 3. Saiiipliiiii liK.itioii in wi-sUiii \\ iscoiisiii. I Biisi- map n\i(l ii illi jicriiiissioii aj llir W isroiisin Crolo^ical iiiid Xahiriil

llisliin/ Siiirfijl.

iiiodcni range ol Cdsli'niojiln pnitcni in North .Vnierica

is represented hv tlie small popnlations lonnd at hill

prairie locations in western Wisconsin described in this

paper. These hill prairies are ii innanls ol more extensive

njiland prairies and sa\'annas that c<)\ered ridgetops and

well-drained sonth- and west-lacing exposures in much

ol southern and western Wisconsin al the time ol Euro-

American .settlement (Anderson. 1954; Curtis, 1959:

262-264). These prairies became established during die

mid-1 loloccTie warm/dr\' Atlantic climatic episode, al)out

S.OOO to 5.()()() years ago (Wendland, 197S:27S: Wend-

land (S: iiiAson, 1971). The Atlantic episode witnessed

the eastward expansion ol the wedge ol prairie .iiid sa-

vanna often called the Prairie Peninsula (Wright. I96S).

The ilnnalic episodes that lollowed the Atlantic were

cooler and/or moistcr. and the prairie/savanna vegetation

conmuuiities were reduced (Wendland. I97S) to stands

that were either fire maintained or situated on topo-

graphicalK controlled droughtx exposures, such as hill

prairies. The cessation ol prairie fires with the onsel ol

Enro-Ameiican seltleineut dining lln' ninitiintii <en-

tun' led to the loss ol most lire-maintained |)rairies in

Wisconsin (Cmlis, 1959:295-')()5). The Mississip|)i Riser

\alle\ has an ameiioialmg clim.ilic ellecl e\lcii(lmg the

J. L. Tlieler, 199";

Patie 119

Table 4. Hank mdcr and assi'ni!)la<;c rrc(|nci]t\ (it siinnneil tfrrestrial i^astrnpods Ironi 35 wi'stfni Wisconsni Inll prairif samples.

Irdst-tice period ikji tliward aloiii; tlic xallcx eoiridoi".

Hill prairies along this eorritlor ari' the oiiK kuowii umd-

ern locations in Wisconsin snpportiu<j; (^/isl nxoplii pm'-

rcra populations.

The large nmnher of state and counts records estab-

lished during this baseline suncx indicates the lack ol

prior stud\ on the small-sized snail hunia in western Wis-

consin. At least one species. Gastrocoplii proccra. now

classified as Threatened in Wisconsin, is beliewd to rep-

rest'ut a relict population frcun the niid-Holocene warm/

(\y\ Atlantic climatic episode. The land snail faunas of

western Wisconsin's hill prairies are onlv one component

ol a complex c()mmnnit\' of animals and plants associated

with this striking landscape (Imla\, 197.3). The continued

existenci' in Wisconsin of Gastrocoj}!// pniccr/i. and per-

haps (jf other members of this biotic ccnnuuinitw de-

pends on the continued e.xistence ol tlie hill praiiie hab-

itats.

ACKNOWLEDGMENTS

1 would like to express mv special thanks to Dr. Kath-

erine Ste\enson for her application of her editorial skills

to this paper. I would like to thank KelK' Kearns and

Thomas \Ie\er at the Wisconsin Department of Natural

Rescjurces. and KariMie Moti\ans, f(.)rmerK" at the Wis-

consin Chapter of the Nature Consenancw for assis-

tance with \'arious aspects of this sur\e\'. Fimding for a

portion of this research was pnnided h\ the Center for

Climatic Research at the Unixersib,' of Wisconsin-Mad-

ison. I thank Drs. M. G. Harasewxch, T. A. Pearce and

one anouNinons reviewer for offering insightful editorial

connnents. Tlie late Dr. Da\id A. Baerreis initiated my

interest in stndxing modern gastropod distributions to

aid in inteqirefation of shells recovered from ancient

sediments. I am also gidtelnl to Dr. Arthur Bogan for

his connnents on aspects ol this paper, and to Dr. Rii-

Pasre 120

THE NAUTILUS, \.

10. No. 4

Tabic 5. Terrestrial (gastropods recovered Iroiii nine high-stress liahitats in western \\

isconsin.

CouiitN:

Sample Location:

UWA:

Taxon: Sample volume:

c:raiil

De\ve\ Hts.

Lafa\ette

M-.Vlomul

(.'rawtord

\i 1 11

Rush C'reek

Cledar \'all<-\

3050

1.0

.3052

Eticoiuilus ftlllU.S

Glijpliii<ilinia indcntata

Hawdiid iidnuscula

TlmiloUlrs (irhorcus

Ait'j^uispira iillcnwtii

Hfliri idisciifi parallchis

Hclirixlisciif^ siiifi^lcytinus

Fiiitctiuu iniiiiilissimuni

Succineidae

Strohilops hihyrinthicii

Strohiloi)s iiffinis

Strohil»i>\ sp.

Qastrocoptii annifcni

Gastrocoptd contractu

Gastrocoptii holzingeri

Gastrocopta pcntodun

Gastrocopta coiiicaria

Pnpoich's alhiliihhs

Viilltiititi citstiilii

Vdllonid pdnidu

Xdlloniii pcrspi'cthd

\'dll<>nid s]).

Iinc'iiilcs

13

NR = Not recoriled.

diger Bieler at tlie Field Miisciiiii ol Natural llistiii-\ lor

ollcring coiiiinents on this paper and accepting \'oiicher

specimens. I would also like to thank Jerenn' Nienow for

help on the tables and figures. An earlier xcrsion ot this

paper was presented at ati annual meeting oi the Wis-

consin .\cadeni\' ol Sciences, Arts, and Letters, held in

Platte\ille, Wisconsin, on ,\[)ril 21. 1090.

litp:h,ature cited

.'\ndersoii, O. 1954. The pliNtosociologN ol dn liiiie prairies of

Wisconsin. Pli.D. Dissertation, Universilx ol W Iseonsin at

Madison.

Bartlein, I'. ]., and 1'. Wcl.h 111 I0S2. Ilolortne eliniatic

change estimated Irom pollen <lata Iniiii the northern

.Vlidwest. Ill: J. (). Knox, !,. Claston and I). M. Mickehson

ieds.i, Quaternarv historx of the Driltless .Are;!. I'leldTrip

Guide Book .Numher 5. (leological and Natural lliston

Surxew l.^iuN'ersitx' of Wi.sconsin-l''Alension, Madison, pp.

67-82.

Baerreis, 1). .\. 1980. llahilal and Cliiiiatie inlei-])retations

from Terrestrial (iastropods at the Cherokee Site. In: D.

(;. .'Xnderson and II S. Seinken, Jr. (eds.). The Cherokee

excavations: l!oli)een<' ecologx' and human ada])tations in

northwestern low.i. .Vcadeniic Press, NY. ])]). 101-122.

Cautis. J. T. 19.59. The \cgetation ol Wisconsin: an ordination

of [)lant conninmities. Universitx of Wisconsin Press.

Madison. 657 p.

Davis, A. M. 1977. The prairie-deciduotrs lorest ecoloiie in

the Upper Middle West, .\nnals ol tlie .American .Associ-

ation ol American Geographers 67(2):204-2l3.

Hole, F, D. 1976. Soils of Wisconsin. UniviTsitx ol Wisconsin

Press, Madison. 223 p.

Huhricht. L. 1985. The distribution ol the native land niol-

lusks ol the eastern United States. Fieldiana Zoologv, New

Series, No 24, Field Museum of Natural Histoiy. 191 pp.

lnila\. M. C. 1973. The case for a Driftless Natitinal Park.

Bulletin ol the .American M;ilacological Union. Inc.. i).7-S.

Levi L. R.. and 11. W. Levi. 19.50. New records ol l.ind snails

from Wisi-onsin. The Nautilus 63(4):131-1.5S.

.M.iher, L. J., Jr. 1982. The paKiiologx of Devils Lake, Sauk

(^ountv, Wisconsin. In: J. C. Knox, L. C^lavton and D. M.

Miekelson (eds.), yuaternan Historv ol the Drittless

Area, Field Trip (aiide Book Numher 5. (ieological and

Natural Histoid Siinev. Universilv ol W isconsiii-Fxten-

sion. Madison, p. 1 19-13.5.

Miniin. 1.. 19(55. The phvsieal geographv ol Wisconsin. The

Universilv ol W iseonsin Press. Madison (iOS p.

Miekelson. I)' M.. j. C. Knox, and 1.. Ci.ivtoii. I9S2. Gl.Kia-

tion ot the Driltless .Area: an (.'valuation ol the evidence.

In: ]. (.'. Knox. L. (;lavton and D. M. Miekelson (eds.),

Quaternan lliston' of the Driltless .Area, Field Trip (aiide

Book Numher 5. Geological and Natural lliston Suncv,

Ihiiversitv of Wisconsin-Extt'usion, .Madison, p. 1.5.5-169.

Morrison. |. P. F. 1929. On the occurrence ol llcndirsonid in

Crawford Conntv, Wisconsin. The Nautilus 43(2):41-45.

Reigle. N I 1963. Northern records of C.;«.v/n)(i)/)/« ;»■<)(■<■/•«.

" The Naulilns 77iU:l(>-18.

J. L. Tlieler, 1997

Page 121

Table (i. Hank order ami asseinl)lai;e irequencs ol suiiiined

terrestrial t^astropcids iroiii nine liii;li-stress luiliitats in western

Wisconsin.

Roosa, D. M. 1984. Xalnral heritage protection in the Dritt-

less Area. Proceedings ol tlu' Iowa Acadenn ot Science

91(11:42-46.

Theler, J. L. 1992. The status of three Vei-fioo species in Wis-

consin. Report to the Wisconsin Department of" N'atnral

Resources, Bureau of Endangered Species, Mathson, Wis-

consin. 42 pp.

Turgeon, D. D.. A. E. Bogan, E.. V Coan, W. K. Emerson, W.

G. Lvons, W. I,. Pratt, C:. F. E. Roper, A. Scheltema, F,

G. Thompson, and J. D. Williams. 1988. Common and

scientific names of acjuatic in\ertel)rates from the United

States and Canada; mollusks. American Fisheries Societv,

Special Publication 16. Bethesda, Mar\land. 277 p.

VVendland, W. M. 1978. Holocene man in North America: the

ecological setting and climatic backgromid. Plains Anthro-

pologist 23:273-278.

Wendland, W. M., and R. A. Br\son. 1974. Dating climatic

episodes of the Holocene. Quatenian Research 4:9-24.

Wright, H. E., Jr. 1968. Histon of the Prairie Peninsula. In:

R. E. Bergstorm (ed.). The Quateniar\' of Illinois, Special

Report 14, College of Agriculture, Universit\- of Illinoi,s,

Urbana. p. 78-88.

THE NAUTILUS 1 1()(4):122-13S, 199']

A New Gastropod Fauna From An Oligocene Back-Reef

Laeoonal En\dronnient In West Central Florida

o

Edward J. Petuch

DfjKutineiit of Gt'olos^'

Florida Atlantic Uiii\f'rsitv

Bota Raton, Florida 33431

utkI

Departnjent ol Pakontoloin,-

Graves Miisfinn ol Arcliacoloi^x ,uul

Natural History

481 SoutI] Federal Ili<ili\\a\'

Dania, Florida 33004

ABSTRACT

T\vent\-one new species of i^astropods and three new gastro-

pod genera are described from a ncwK -recognized back-reef

lagoonal facies of the Oligocene Snwannee Formation. The

new species, which were collected in a lirnt'stone quarry in

westernmost Polk Count\', Florida, inclnde: Astraca (Astral-

iiiin) polkcn.sis n. sp. (Turbinitlae), Piimzi.sinus kentlmvi n. sp.

(Potaniididae). Prisiiuircritliiiiiii prisma n. sp. (Cerithiidae),

Paclii/troinminm iliilli n. .sp. and Paclujcrommium inansfiehli n.

sp. (i)otli .Naticidae), Ct/i)nir()rl>is ki'iuhvici n. sp. (Cvpraeidae).

Spiniftilf^^iir i^c'inmulatiiin n. sp. (Bus\conidae), Soh'no.steira sit-

iciiiiiifcnsis n. sp. (Buccinidae), Vasuin siiwaiincensi.s n. sp.

(Turl)inellidae), Ftilsih/ria kciidrcui n. sp. (\'olutidae), Con-

oiiiitra ki-iiilnwi n. sp. (N'olntomitridac), D<'nliiiuir<j,ti dalli n.

sp., Persiculri (hickin/i n. sp., Per.siculii nuieneili n. sp., Persi-

cula suuditni-cnsis. n, sji,, PniiiiiDi friaic n. sp., Pniimiii je.s-

sicac n. sp., Pniniiiii sdiuhyu- n. sp. (all Margin<-llidac). Coiiits

lAsprcllin kcndrciLt n. sp. (Gonidac). Plciii'dfiisid (hiwliitfl,! n.

sp. ('Inrridael, and Siiaaiiiirsriiplui liiiiliu- n, sp. (Cylichnidae).

The new genera, whicli ajipear to be irslrirled to Oligocene

coralline enxironnicnts, include: ('I'stiiiiici rilliiiiiii n. ij;cii, and

Prisiiiacciifhiniii n. gen. (( !crilliiidae) and Siiiiaiiiu'siaplia n.

gen. ((A'liehnidac). In tolal, seventv-two species of gastropods

were collected from h)iir diiierent back-reel lagoonal biotopes,

including biolicruis ol the branching coral Sti/ltiplmm iiiiiiutis-

siiiui, open-bottoni sand areas, sea grass beds, and inlertidal

mud flats with o\ster bars.

Kci/ Udifls: Oligocene, I'lorida. Suwarniee Fornialion, tA)ral

Biohenn. (Gastropoda.

INTRODUCTION

The early Oligocene (Latdorfian) gaslri)|)<)il lanna ol'tlic

lower Mississippi Ri\cr X'aJlev and soutlieastern United

States has long i)eeii known to he one- ol the nio.st spe-

eies-rieh in the Eastern North American (leno/oie. This

tropieal-snhtropieal laniia is ol particular interest to exo-

hitionan paleontologists and hiogeographers in that it

ehi

N(

contains the liist-knowii t'.xaniples ol main (.■lassie .\eo-

gene Eastern American endemic genera and is the cUi-

cestor ol the Miocene, Pliocene, and Pleistocene C>a-

loosahatehian Mollusean Provinces (Petuch, 1982; 19S8).

As presently understood, the richest and best-preserved

Oligocene assemblages are found onlv in southwestern

Mississippi State (MacNeil & Docken', I9S4), and this

area is considered the arehetvpe tor the earlv Oligocene.

To date, three hmidred and ninet\-se\en species and

subspecies of gastropods are known from the \icksburg

Group (MacNeil & Docken-, 1984).

In the earlv part of tlie Twentieth CyenttuA, N'icksburg-

aged gastropod faunas were discovered along the Flint

Ri\er and Blue Springs areas near Bainbridge, Georgia

(Dall, 1916). These were found to be associated with the

rich and extensive Oligocene coral reefs that were de-

scribed from the same area bv Vaughan (1900). Later,

Mansfield (1937) described an e\en richer Floridian

Vicksburg-aged gastropod fauna, primarilv from (|uarries

in Pasco and Hernando Counties (west central Florida^

and showed that several Flint Ri\er species occurred

that far south. Although eontaining some elements ol the

classic X'ieksbnrg lanna ol the .Mississippi Embasment.

these two so\ithern loral reel-associated lamias also

housed numerous characteristic endemic elements, in-

chidiiig the stidinbid Chiliilttlax, the \olutid Fiilsilijrui.

and a distinctixe assemblage ol o\er filtc'en species ol

cerithiids and fi\e potamidids (combiiu'd Flint Ri\eraiid

Suwannee Foriii.ilions). Since Mansfields work in 19.37,

liowe\er, no new eompreliensi\e stiuK ol the s\steni.ities

ol the Suwannee gastropods has bei'ii published.

l^eeeiilK. an e.vposme ol a pre\ ioiisK -miknow ii, higlllv

lossililercnis hu'ies ol the Suwannee Fcnination has been

louiid in a limcslone (|uaii\ in extiemc western Polk

CoiiuIn, west central i'lorida (The Terramai' Pit on l''lor-

ida Highwav 54. approximatcK' 5 miles east ol '/eph\r-

hills. i'a.sco CoimtN) (E. X'okes, 1992). This localit\ rep-

E. J. Pftucli. 1997

Paw 123

resenti'il a liatk-rci-i laij;()(inal t'n\ iinniiifiit that liad

lonned lii'liiml a icct tract to the west and cciiitaincd

the largest miinlier oi gastropod species vet loiiiid at aii\

single Suwannee exposure. All specimens ot corals and

niollnsks ironi the Terraniar site are silicified (siliceous

pseudonioqihs), much like the Flint Hi\-er niaterial, and

are generalK" well-presened. Throu<4li the iiitensi\t' col-

lecting of Mr. Eric Kendrew ol X'alrico, Florida, and

througli additional collecting hv niyseli, se\'ent\■-t^\•o spe-

cies of gastropods (listed in Appendix 1 ) are now knowTi

from the Terraniar Pit, more than douhling Mansfield's

total of tlnrt\-one species. Nine of the Suwannee species

reporteil In' Mansfield (1937) and one h\- E. N'okes

(1992) were not collected at the Terrainar site, and these

are listetl in Appendix 2. When Mansfield's and N'okes'

additional species are combined with the list of those

from the back-reef lagoonal enxironnicnt. the total num-

ber of known Suwannee gastropods increases to eightv-

two, less than one third of the numlier of X'icksbvu'g spe-

cies.

Of the se\c'nt\-tx\'o species collected at the Terraniar

Pit, txvent\-one are new to science. Three new genera,

all of which appear to be endemic to the coralline en-

\ironments of the Flint Rixer and Suwannee Forma-

tions, are also new. Following a discussion of the paleo-

ecologv of the Terramar Oligocene site, tliese new spe-

cies and genera are described.

PALEOECOLOGY OF THE RAf:K-REEF

LAGOOxNAL ENVIRONMENT OF THE

SUWANNEE FORMATION

The stnd\ material from the Teiramar Pit was collected

either as perfectlv-present-d spt'cimeus looseK embed-

ded in a residual unconsolidated calcilntite or as speci-

mens emiiedded in blocks of a dense, partialK-silicified

limestone. These specimens were often remoxed in a

fragmentary state. Judging from the biotic assemblage

collected in both lithofacies. the entire area represented

a shallow (less than 5m deptli), quiet-water carbonate

lagoon. This lagoon formed behind a still-mistudied cor-

al reef tract that had developi'il farther to the west. E\-

idence for this unmapped Oligocene ri'cf tract, including

the presence of massixe coral heads and reef niollnsks

such as the bixaKes Lima luilcitsis Dall. 191fi and Clianui

sp. and the stromboidean gastropod Teirlielliim luniaii-

docnsis Mansfield. 1937. has been nnco\ered in quarries

in central Pasco and Hernando (bounties (Mansfield,

19.37). The Terramar lagoonal emironment, existing be-

txveen this reef tract and the Oligocene western coast of

Florida, contained four main substrate txpes: 1. bio-

herms composed of Sti/lnpliorn corals: 2. deeper lagoon-

al, open-bottom areas; 3. sea grass beds, and: 4. \er\

shallow-water o\ster beds and intertidal mud flats.

Of the four biotopes that existed within the Terramar

back-reef lagoon, that of the Sti/lopliora coral bioherms

xvas apparentK the most extensive. Composed almost

entirelv ol the small branching coral Sti/Iophora iiiiiiu-

tissiiiui N'autilian, 1900 these bioherms formed denscK

iiitermeshed beds that sheltered a wide \arietx' of gas-

tropods. Manv blocks of limestone examined contained

literallv hundreds of small pieces of this branching coral

antl loose fragments were the most aiiundant component

of the residuum facies. As exidenced bv haxing been

collected in silii in Stt/h>phora-r\L-\i limestone blocks, a

mmiber of Suwannee endemic niollnsks appear to ha\e

been associated with these biolierms. Some ot the most

characteristic gastropods inclndi- Calliastoma siliciitiiiti

Mansfield, 1937 (figure 19) Cypriicarhis kcmlrcwi n. sp.

(figures 22, 23), Salmosteira .siiwannccnsis n. sp. (figures

33, 34), \'ri\uiii Mtuiiiinccnsis n. sp. (figures 48, 49), Fal-

silipia kviidrcwi n. sp. (figures 37, 38), and Conus (As-

prcU(i) kciiilrcwi n. sp. (figure 72). Sexeral of the large

Suwannee cerithiids. including Ccsliniircritliiitin liroohs-

lillciisis (.Mansfield. 1937)(figures 3. 10) Ccsfiintn-ri-

lliiiiiii pdscot'iisi^ (Mansfield, 1937)(figure 2), and Pii\-

macerithuun prisma n. sp. (figures r>, fi, 11) were also

associated with the Sti/lophora bioherms. Iiiterestinglv,

sexeral specimens of Paclujcrommiiuit mansficldi n. sp.

(figure 30) x\cre also collected in Sti/lophora-rich lime-

stone, possibly indicating that this naticid may haxe lixed

on, or in close proximitx to, the coral bioherms. The rare

muricid Foiricria (Diillimiircx) nifinipicolus (Dall. I91R)

(figure 31), wliii'h was prexiousK' known onlx' as an in-

ternal mold from the Flint River Formation, was also

found to be associated xvith this biotope. Small heads of

the star corals Montastrea haiiihridf^cnsis (\'auglian.

1919) and Antiffiastrea siliccusis X'anghan. 1919 were

also present, although uncommoiiK. throughout the Stt/-

Icphora bioherms.

Interspi'ised betxxT^en the coral bioherms in deepi-r

water (approximatelx' 4-5 m) areas were large patches of

open, fine-grained carbonate sand bottom. This biotope

supported a rich gastropod fauna of xx'hich the Sux\aunee

endemics Oliva (Omot^i/mna) hrotih.siillcnsis (Mansfield,

1937) (figure fi5) and A/.)/r(//c/ Iniwciiae (Mansfield. 1937)

(figure 13) were the most abundant. A large \'icksbnrg-

aged marginellid fauna also was associated xsith this bi-

otope, and included species such as Persicida dackcn/i

n. sp. (figures 52,53), Pninitm cricac n. sp. (figure fi4).

Pniuiim jcssicac n. sp. (figures 56. 57), and Pniniun saii-

dnic n.sp. (figures 58, 59). The strombid OiihauUix lirr-

iiandofii'iis- Mansfield, 1937 (figures 20, 21) which is also

present in the coral reef enxironments of the Flint Rixcr

Formation, was abundant on these open bottom areas,

fixing in large colonies much like Strnmlms s.s. species

do in the Recent. Sex'eral of the new Suwannee species

described here, such as Pachi/rnimittiiim dalli n. sp. (fig-

ure 26). Sj)inifnl<s^tir ff'inmiddtiim u. sp. (figure 47), Cim-

oinitra kciidrcui n. .sp. (figures 44, 45) and Pleurofusia

dowlinoi u. sp. (figure 68) also appear to be associated

with this biotope. All of the widespread \icksburgian

gastropods found in the Terramar Pit (,\ppeiuli.x 1, des-

ignated b\' the letter "Y"), including such characteristic

species as Scmiieiia^tis mcnthajontis MacNeil. 1984

(figure 7), Ficus mississipjiivusis Conrad. 1848 (figure

24), Taliti/phis mifisis.sippicnsi.s (Gertmann, 1969)(fignre

29), Pnllaci-ni i-asviji (MacNeil, 184S), Chnolithcs licks-

Page 124

THE NAUTILUS, N'ol. Ill), No. 4

Figures 1-19. 'rrocliidac. Tiirbinicluc, Folaniididac, Ccritliiidac, and \l()(liili(l,ic lidni tin- .SiiwaiiiK'c hack-iVfl lagoDii.s. 1. Cirilliniiii

(new genus?) cookci Dull, 1916. lengtli 36 nun. 2. Ccstiiinccrilliiuin /xiscocnsis (Manslicld. 1937), .smooth variant, length 37 mm,

(t\pe of tlie new genihs Ccstiimccrilluiim). 3. Ccstuiiurritliiiiiii hmoksi illcnsis (Mansfield. 19:571, length 24 mm. 4. Crstuinccrithiuin

liveoakensis (Man.sfield, 1937), length 24 mm. 5,6. Prisnitici ritliiiiiii jirisimi Pclnili, nsp.. liololvpe (UF 75978)(fragnientar\. missing

E. J. I'rtiRli. 1997

Page 125

l)iir<ii-iisis (Conrad, lS4'S)(figuri' 46), Fiisiniilra i-oiKjiiis-

ta (Conrad, 184S)(figure 50), and TcnhnilTrnhnlliini i

divmim Conrad, 1848. were a.s.soeiated with tliis open-

liottoin area. The Flint Ri\er species Si-iuivcii(i<ius sili-

cium (Dall, 1916)(figure 17), Ccritliiiiiii (new geTius?)

cookci Dall, 1916 (figure 1), Aiujiiilliiiop'^is fliiitcnsi.s

(Mansfield, 1937) (figure 25), and I'dhihirui iiuiii.sficldi

(Dall. 1916)(figin-es 39,40), also oecurred in this hiotope.

The classic Suwannee iiide.x echinoiil Rlii/iiclinliiiiipiis

gouldii (Bouve) was abinidant on tliese open-bottom ar-

eas and niav ha\e been preNed-npon h\ the widespreatl

V'ieksburgian Pluiliuni ctichituni (Conrad, 1S48)( figure

27).

Also interspersetl between the large Stijlophoni bio-

herms were beds of sea grass, most probalil\- Thalassia.

Evidence for these Thalassia lieds is seen in tlie pres-

ence of characteristic indicator gastropmK. all of whicii

ha\e T/irt/rz.v.v/rt-dwelling descendants in the Recent. Li\-

ing on the sea grass blailes were a numln'r of small al-

givorous species, including Astraca ( .\sli-nliiim > polhnisi.s

n. sp. (figure 18). Ccrithitim insulatuin Dall, 1916, (fig-

ure 12) and Modulus liveoakensis Mansfield. 1937 (fig-

ure 16), and a large fauna of small, TlialassiaAovin^ mar-

ginellitls such as Dcutiinan^o dalli n, sp. (figures 62,63),

Hi/aliiKi silicifiuvia (Dall. 1916)(figure 51), Pcrsiculn

uiacncili n, sp. (figures 54,55), and Fcrsicula suuainiccii-

sis n. sp. (figures 60,61). Living within the grass beds

were larger gastropods such as Cerithioclava culcxtilc

(Dall. 1916)(figure 9), the Suwannee endemic Tiirhiiirlla

suutninciisis (Mansfield, 1937)(figure 42, 43), and a host

of small bivalves, priinarilv Cdtione haiul>rid<^cii.sis Dall,

1916, Pliacoidcs hcniandoensis Mansfield, 1937, and

Ghjci/iiicris suwauucusis Mansfield, 1937. .\ living ana-

logue to the Suwaimee grass Ix'd connnunitv is seen in

the Recent along Roatan Island, Honduras, where spe-

cies of Ccritliioclaia. Turhhulla. Astraca. Modulus. Pii-

sirula. Hi/aliiia. and Phacoidvs all co-oi'cur in 'Phalassia

beds. The unusual Suwannee endeniii.- bubble shell, .S'/(-

uauiu:scaj)l>a liudac n. sp. (figures 70,71 ). max also have

been associated with the sea grass beds.

Faimal evidence also points to the existence of shallow

intertidal mud Hat enviromnents within the Terramar la-

goon system. This is seen in the presence of a tlistinctive

potamidid fauna composed of tA\'o species of TelcscDpi-

uui, [T. hlrukwatereusis (Mansfii'Id. 1937) figure 15. and

T Itcniandiicusis (Mansfield. 1937) figure' S]. and Pi/ra-

zisiniis kciidrcwi n. sp. (figmv 14). bi the Recent bido-

Mahusian region, the amphibious genus I'lLscopiuui

lives on tin' highest mud Hat areas near mangroves and

is ex^50secl to air lor long periods of time. Often living

along with Telescopiuui on these exposed flats is the ge-

nus Tcrchralia. which is closelv related to the Floridian

endemic genus Pipazisiuus and probabK occupies the

same ecological nic-he as did its extinct New World coun-

teqjart. The ovster Ostrca jjaroxis (Lesueur) Dockeiy,

1984 is conmionlv encountered in large masses in some

blocks of limestone, indicating that small ovster bars also

grew on these intertidal nuid flats. The melongenid

Mipislira cn/ssicdinula (Conrad. 1848)(figures 35, 36)

and the muricid Plcii/uctus propeposti (Mansfield,

1937)( figure 64) were associated with these ovster bars

and were probably the major predators on Ostrca pa-

roxis. A Recent analogue to this Oligocene mud flat and

ovster bar enviromnent is seen in northwestern Australia

along the .Aralura Sea. where Ptcn/uotus iPhcdualli).

Tcrchralia. and Telescopiuui species occur togetiier on

nmd flats and oyster bars near mangrove forests.

systematic; section

The holotvpes ol the new species described here are

deposited in tlie lossil invfrtebrate collection of the

Florida Museum of Natural Historv. University of Flor-

ida, Gainesville, Florida, and bear UF numbers. Para-

tvpes are depositeil in the research collection of the au-

thor, as is a voucher collection of each species collected

at the Terramar site. All specimens are siliceous pseu-

domoiphs. .\ vouciier collection is also deposited at the

Cu'aves Museum of Archaeology and Natural Iliston.

Dania. Florida.

Class Gastropoda

Superfamilv Trochoidea

Familv Turbinidae

Suiifamily Astraeinae

Genus Astraca Rikling. 1798

Subgenus Astraliuiu Link. 1807

Astraca (A.straliuut) jxilki'itsis ni'w species

(figure 18)

Description: Shell small for gemis and suligenns. con-

ical, witli elevated spire whorls: suture indistinct; pe-

riphen of body whorl flattened, bladelike, romided. with

faint serrations; peripheries ol earlv wiiorls and spire

whorls edged with 15-20 small, roimded knobs: whorls

strongly ornaniented with 5 strong, beaded spiral eords;

appro.xiniatelv one-halt of spire vvliorls). length 45 nun. Itvpc iit the new genus Pri.\iii(ici htliimiil 7. Sciiiiiciidtiiis iiicnfliafi'iitis

MacNeil. 1984, length 15 mm. 8. Tricscitpiiiiit luriuniiliuiisis (Mansfield. 1937). length 4VJ nun. 9. d ritliiochica cutcxlih- (Dall,

1916), leni^tli 57 nnn. 10. Ccstuiiiiiiitluiiin l>niiik\i illnisis i Mansfield. 19.37). heavilv-sculptnrt'd variant, length 29 nun. H. Pris-

iiKicirithiuin prisma Petnch, n.sp.. paratvpe (spire vvliorls. missing part of body wliorl ), length 19 mm. 12. Ccrithium iTlicriciuin)

insulainrn Dall. 1916. length 22 nnn. 13. Apiciila hmccnac (Mansfield, 1937), length 39 mm. 14. Pt/razisiiuis kciulrcici Petnch,

n.sp., Iiolotvpe (UF 75977). length 62 nnn. 15. Telescopiuui hhiekieatcrensis (Mansfield. 1937), length 43 nnn. 16. Modulus live-

Odkensis Mansfield, 1937. height 15 nnn. 17. S<'iiiii(iiaiJ,us silieiion (Dall. 1916). length 15 nnn. 18. Asfrrien lAstmlium) polkensis

Petncli, n.sp.. Iiolotvpe (UF 7.5976). lieii^lit I I iniii 19. ('ulliosUniiii silientiou Mansfield, 1937. lirit;lit 14 luiii.

Page 126

THE NAUTILUS. N'ol. 110. No. 4

I to 3 c'.xtrc'tiieK fiiic tlircadlikc sccoiulan spiral cords

present between 5 main coids; hasi' oi shell Hattened,

ornanienteil witii 10 strong spiral cords: nnihilical region

sliglitK' depressed.

Material examined: 1 1( )L()TVFE— Height 11 mm.

diameter (incomplete) 13 mm. in hack-reef lagoonal fa-

des of the Sii\\'annee Formation. Tcrramar Pit, western-

most Polk Count\, Florida, UF 75976.

Etv'mologj': Named for Polk Countx. Florida, localit)

ol the Tcrramar Pit.

Discussion: This new species is the first A.striica

known from the Eastern North American Oligocene. Of

the kncmii Cenozoic American A.stralinin species, A.

polkcnsis is closest to the late Eocene (Jackson Stage)

Astraea (Astralium) withlacoochcnsis Palmer, 1953 from

the Inglis Formation of the Floridian Oeala Group. The

new species differs from this prol)aiile Eocene ancestor

in ha\ing a smaller shi'll with a Hatter, less trochitorm

shape, stronger spiral scnlptnri' on the whorls, and a re-

duced, nonprominent peripheral keel.

Snperiamilv Cerithioidea

Faniilv Potamididae

Genus Pt/razisinii.s Heilprin, 1SS6

Pt/razisimis kciulrcivi new species

(figure 14)

Description: Shell of average size for genus, elongat-

ed, cerithiform; whorls ornamented with large, promi-

nent. e\enl\ -spaced knobs; knobs distinctly angled at

shoulder. a\eraging 10 per whorl; whorls scnlptureil with

5 large, prominent, raised spiral cords; base ot shell flat-

tened; suture impressed, slightly canaliculate; apertural

region of unique holotvpe broken, so exact natine o(

adult lip is not known.

Material examined: HOLOTYFE — Length 62 nun,

width 28 nun, in nnid flat and ovster bar (acies oi the

Snwamiee Formation, Terramar Pit, westernmost Polk

County. Florida, UF 75977.

Etvinologv': Named for Mr. Eric Kendrew ol N'alrico.

Florida, who coll(>c'ted the holotvpe in the Terrainiu' l^it.

Discussion: Pi/riizi'-iiniis koitlnu i is the olilcst-known

member of its endemic Floridian gemrs. The ni'w spe-

cies is most similar to the A(|nitaniau Miocene P. cor-

imttis (Ilcilprin. 1.SS6) from the Tampa Nh'mberof the

Arcadia Formation, but diliers in being ;i narrower more

elongated shell with more numerous knobs per whorl.

in having an angled shoulder area and angled knobs. ;uid

in having a much more scnlptuicd shell, with (i\c dis-

tiTict spiral cords.

Familv Cerithiidae

Gemis ('/■•ihniiccrilliiiiin new genus

Diagn<)sis: ('eritli shells ol average-to-large size (25-

45 nun), slender and elongated: sutures indented, pio-

dncing tnrreteil spire whorls: shell stiilptnre character-

istically composed oi hvo distinct sections, with early

whorls ha\ing a reticulate sculpturi' composed oi strong,

oiten beaded, spiral cords intersected In longitudinal

ribs, and with tlie later whorls and bod\' whorl being

smootli, with fi\'e or six \ei-v large, ver\' prominent belt-

like spiral cords: on adults, belt-like cords e.xtend beyond

edge of lip, producing strong dentate, serrated appear-

ance (shown here in figure 2); siphonal canal short, stnb-

In-.

Ty|Je species: Ccnth'uiiii pascoerms Mansfield, 1937,

Suwannee Formation. earl\ Oligoceue of Florida (figure

2).

Other species in Cestumcerithitim: Ccritliiiiin

brooksiillctisis Nhiusfield. 1937 (figures 3.10), Cerithiiini

liveoakciisi.s Mansfield, 1937 (figure 4). and Ccritliiiiin

siiwannccnsis Mansfield, 1937. all from the Suwannee

Formation. I'arK Oligoceue oi Florida, and Cfiithiinn

vd'^iiKilinti IDall, 1916, Flint Ri\er Formation. earK Oli-

goceue of southern Georgia and .Alabama, and the Su-

wannee Fornuition of Florida.

Etymologv: A combination oi the Latinized Greek

"ccstii.s" (belt) and "Ccritliiiiin" (little horn), pnuhiciug

"belted little horn", in reference to the [ii-omiuent and

characteristic cords on the bodv whorl.

Discussion: With the exception of the Flint Ri\er and

Suwannee (\:stiiiiiceiitliiiiiii vaginatiinr all other knowni

species oi this new genus ha\e been iound only in the

Floridian Oligoceue. Here, tlie\' appear to luiM' been

closelv associated v\ith the extensive coral reels and coral

bioherms that dominated Latdorfian western Florida.

Four oi'the fi\e known species are restricteil to tlie Su-

wann(X' Formation and can be considered classic index

fossils lor the Floridian earK Oligoceue. The genus aji-

pears to ha\e been restricted to the Oligoceue. as no

known members ha\i' been collected in the o\'erl\ing

Tampa Member of the Arcadia Formation (Aquitaniau

Miocene) or irom the underKing Eocene formations.

The amount ;md degree of the characteristic l)elt-like

corded sculpture appears to be \ariable. sometimes co\-

eriTig onK the hist two whorls and at other times co\-

ering most ol the kiter whorls ol members oi the same

species. This is particnhirK noticeable oti the t\pe spe-

cies, Ccstiiincciitliiiiin pascocn.sis. which can be hea\il\-

belted (like the holohpe. Mansfield. 1937. plate 7, figure

4) or relativcK smooth, with onk' the bod\ whorl being

belted (like the specimen illustnited here in figure 2, or

like Mansfield's "Ccrilliiiiin s]i. afi'. iii<iiiialiiiii Diill",

plate S. figure 4). Specimens of (.'. pascociisis with belted

scnlptni'e tluit iuteigrades betA\een these two moiplio-

logical extremes h;i\(' been collet led at the Ternnnar Pit.

demonstrating thtir cous[)e(ificit\. This sanu' degree oi

sculptural variabilit\ is also seen in (' hrooksvillcn.sis.

with .some specimens ha\ing at le:isl hall the whorls belt-

ed (liki- figure 10 shown here or like Mansfield's ( 1937)

figure 3 on pkite 7' and otlii'rs having the belled sculp-

E. J. Petuch. 1997

Page 127

Uuv link III! till' liDiK wiioii (like figure o shown lu're

or like Mansfields figure 5 on pkite 7).

Ccstitmccrithinin is iiM)st similar to the s\inpatric ge-

nus Thcricium hut differs in lia\ing tlie liodv whorl, antl

sometimes the later whorls, seulptured with the promi-

nent heltlike eords and in ha\ing a strongK' dentate edge

to the adult lip.

Genus Fhsintuiiithiiiiit new genus

Diagno.sis: ("eritli shells ol laige size, axeraging (SO

mm in length, elongatedk p\raundal m outline: shells

higliK' inflated. e.xtrenieK' thin and fragile; shoulder slop-

ing, suture impressed, produeing turreted spire whorls:

whorls ornamented with 8 low. e\eul\-spaeed narrow ax-

ial swellings, producing inidulating. faceted appearance:

undulating varices become ohIiqueK' angled on l)od\

whorl; entire shell surface smooth, laintK' sculptmcd

witli 24-26 extremeK fine. e\enl\-spaced incised s[iiral

threads; aperture o\al in shape, with adult !i[) almost

nonexistent, onl\' faintK' thickened.

Type species: Prisiiiiirciifliiiiin jihMiid Petuch. n.sp..

Suwannee Formation. earK Oligoeene of Florida (fig-

ures 5.6.11 ).

Etymology: A combination ol the Latinized Cw'eek

"prisiiia" (sawed or faceted) and "Ci'iilliiinn" (little

horn), in reference to the faceted appe:nance ol the shell

whorls.

Discussion: With its large papei- thin. higliK inllatetl

shell :md faceted appearance. Prisitwccritliitiiii stands

out as one of the most distinctive members of the rich

eerithiid faima of tlie Suwannee coral reefs. Because of

its extreme fragilitx. this new genus is almost alwaws col-

lected as broken fragments. Judging from the ainmdance

of shattered pieces in the limestone iilocks within the

Terramar Pit, howe\er, Pri.sinaceriihiiiiii must lune been

a conniion component of the Sti/Iopli<irii lnolieiiiis.

Although ha\ing the large size of other Suwannee eer-

ithiid genera such as Thcriciuni. Ccritliiochiid. and Ces-

tnmceritliiiiin, Prisuiacerithiuin more cIoseK resembles

an immenselv-oversized Ahilxi. Botii genera, although

differing greativ in size, share several moqiliologieal

characteristics, such as the higliK' inflated, thin shell, dis-

tinct rounded axial swellings, and a shell ornamentation

of finely incised spiral threads. Further rese:ir(.'h and

larger collections of better specimens ma\' show that

Prisinan'rifhiiiiu bekjngs. along with Alaha mu] Al/ihiiia.

in the snbfamiK' Diastomatinae.

At present, PrisnuiccntJilitm is known ouK liom the

lagoonal faeies of the Suwannee FormatioTi. It bears no

similarities to anv other known huge eiinteiii])orani'ous

Eastern American eerithiid.

Prisinaci'ritliitiin prisDui new species

(figures 5, 6. 11)

Description: General shell shape, lorm, aiitl size as

for ilescription of genns.

Material examined: 1 lOLOTYPE— Length i frag-

mentary', missing approximatek' one-half of spire whorls)

45 mm. widtii 24 mm, in back-reef lagoonal faeies (SY(/-

liijilidrii bioherm) of the Suwannee Formation. Terramar

Pit. wi'sternmost Polk Gonnt\. Florida. UF7597S:

PARATYPE — lengtli (spire whorls missing, part of bod\'

whorl 19 nnn. same loealitv as holotxpe. in the researcii

collection of the author.

EtyTnology: Named in reference to the faceted ap-

pearance of the shell.

SuperiamiK N:iticoi(l(M

FamiK Xaticidae

SnbfamiK' (;lobnlariiii;ie

Genus Piirlniiroiiniiliiin Woodring. 1928

Pticlnicroiiniiiuiii ihilli new species

(figure 26)

Description: Shell small lor gemis. elongated, with

rounded bod\' whoil and high, stepped, scalariiorm

spire; sutures higliK indi'nti-d, producing wide, shallowiv

canaliculate subsntural border: edge of canaliculate bor-

der sliglitlv raised, faintK carinated: aperture o\al: no

mubilicus present.

Material examined: HOLOTYPE — Length 18 nun,

width 13 nnn, in back-reef lagoonal faeies of the Suwan-

nee Formation, Terramar Pit, westernmost Polk Count\'.

Florida. UF 75979

Etymology: Nanu'd in honor of tlie great pionet-r pa-

leomalacologist. William Heale\' Dall, who described the

Flint River gastropod fauna.

Discussion: l^efore its formal description liere. this

small Oligoeene naticid hatl been illustrated bv both Dall

(1916. plate 88. fig. 11) and Mansfield (1937. plate 9.

fig. 7) but ne\er correctK nametl. Dall referred it to

"Amauropsis ocalana Dall"', a Jackson-aged Eocene spe-

cies from the Floridian Ocala (w'oup (a miineii dubhun —

see Mansfield, 1937. p. 177). Mansfield later referred

this species to "Aindtimpsis afi. A. hiinisii jtieridiDnalis

Pilsbn". a related but dillerent shell from the Miocene

Baitoa Formation of the Dominican Republic, .although

similar-appearing, both the Eocene and Miocene species

differ from P. dalli in being larger, more inflated shells

with lower, less protracted spires, and in ha\ing deeper

and wider subsntural channels. To date, P dalli has only

been collected in the Flint Rixer and Snwamiee For-

mations.

PncJiiicn>}uiiiiniii iiiiiii\fiildi new species

(figure 30)

Description: Shell of ;i\crage size for genus, inflated,

globose, with lounded bod\ whorl: spire protracted, sca-

lariform; spire whorls relatively straight-sided; sutiu'es

higliK' indented, producing narrow, \en' tleepK' canalic-

ulate subsutnr:il border; edge of canaliculate bolder

Page 128

THE NAUTILUS, Vol. 1 10. NO. 4

Figures 2()-34. Strombidae, Cvpraeidae, Ficidae, Ca.ssidae, Naticidae, Muricidac, and Hnccinidai- fniin llic .Siiwaiiiirc liack-rcei

lagoons. 20,21. Ortluiuinx Iwrnamlornsis Mansfield, 1937. lenf^lh 70 nnn. 22,23. Cijprdcorlns kcndnui rclucli. n.sp., liolotApe

(UF 75981), length 26 mm. 24. Fiats mississippicnsi.s Cdniad, IS4S. lengtli 17 mm dragmentan ). 2.5. Amptillinopsis fiintcnsis

(Mansfield, 1937), lengtli 27 mm. 26. Pticliiicroniiiiinm iliilli Petucli, n.sp., Iiolotype (UF 75979), Icngtli 1^ imti. 27. Phnlium

awlatura (Conrad, 1848), lengtli 29 mm. 28. ricnjnolns piopeposti (Mansfield, 1937), length 35 mm. 29. Tdlitijpliis iiii^sissippicnsis

E. J. Petiicli, 1997

Faec 129

liiij;lil\ raised, proiluciiig sliarp, tlistiiict (.■aiiiia: iipcitiirc

(i\al; Tio uinhilicMis prest-nt.

Material examined: HOLOTYPE — Length 27 imii.

widtli 21 iiini, ill hack-rt'el lagoonal iacics (in association

with Sti/lophora liioherins?) of the Suwannee Forma-

tion. Terraniar Pit. we.sternniost Polk (Jonnt\'. Florida.

UF759S(): P.\R,ATYPE— length 24 nnn. same locality as

liolot\])(', in the research collection ol the author.

EtMiiologv: Named in honor oi Wendell (-'. Mans-

field, the pioneer paleonialacologist who first descrihed

the Suwannee gastropod ianna.

Discussion: This new Pachifcroiiiiiiiiiiii closeK resein-

hles, and is probabK' ancestral to. /' jiiuUland (Dall.

1892) from the .-Xquitanian Miocene Tampa .Member of

the Arcadia Formation. The new species differs from P.

jioriddiui in ha\ing a less-inflated shell with straighter

sides to the whorls, a more protracted spire, and a much

wider and deeper subsntmal channel. ,\t ])resent, P

luaiisfichli is known onl\' from tlu- coral leef lagoonal

facies ol the Suwannee Formation.

SuperfamiK' C\praeoidea

Family Cyjjraeidae

Subfamily Cypraeorbinae

Genus Cypmeorbis Conrad. 18fi.5

Cyprncorbis kcmlreui new species

(figures 22. 23)

Description: Shell of a\erage size for genus, ovate,

slightly subcviindrical: base flatteni'd: aperture narrow,

widening slighth' toward anterior end, almost straight,

recuning slightK at posterior end; labial teeth propor-

tionalK' large, e.xtending onto base ol lip. 23 in munber:

columellar teeth smaller than labial teeth. 21 in number:

extremities reduced, projecting onlv slightK hom main

shell bod\': spire and posterior extremits covered with

broad, low callus: fossula poorK'-de\eloped. supported

In single large tootli along anterior edge.

Material examined: HOLOTYPE— Length 2rs mm.

width IS nun, in back-reef lagoonal facies {Sti/lnpliurd

bioherm) oi the Suwannee Formation, Terraniar Pit,

westernmost Polk Countv; Florida, UF759S1; Parat\pe-

lengtli 26 mm, same localit\' as holotspe, in the research

collection of the author.

Etymology: Named for Mr. Eric Kendrew of X'ahico,

F"lorida, who collected the holotvpe in the Terramar Pit.

Discussion: Ct/praeorbis kciidnwi. one ol the last

species of the Cypraeorbis Hneage, is most similar to C.

vcnfiipotcns (Cossmann. 1903) from the [ackson Eocene

.Moodvs Branch I'^ormation ol .Mississippi. The new s[)e-

cies differs honi this older cowrie in ha\ing a iimch

more c\iindrical, less iTiilati'd shell witii straighter sides,

more rountled. less projecting e.xtremities. a straighter.

less recin"\ed aperture, and more munerous and finer

apertural tei'th. Tlie \'icksburg Oligocene cowrie listed

as "Cypmeorbis all. i'. icutripotens (Cossni.in. 1903)"

by MacNeil and Docken (19S4, plate 16, figs. 1().12.]5)

is veiv similar to C kciuln-ui, !)ut iliffers in being a more

inflated, rounded shell with a slightlv more recuned ap-

ertiue. F'urther stniK' ma\ show that the Mississippi shell

(from the Mint Springs Foiination) belongs to a north-

ern iioimlation ol (' kciKbiui.

SuperfamiK Hneciuoidea

FaiiiiK' Busxcouidae

SubfamiK Busx'couiuae

Genus Spinifiili^iir Petnch. 1994

Sjiiiiifiil^iir ^cimiiuliilniii new species

(figure 471

Description: Shell ol average size for genus, with dis-

tinctly sloping shoulders and high pagodiform spire;

edge of shoulder shaipiv angled, bordered bv promi-

nent, thick, undulating carina; undulations on carina pro-

duce small, rounded, evenlv-spaced knobs, numbering

between 14 antl IS per whorl; suture highly indented,

causing carina on previous whorl to overhang subse-

quent whorl; sloping spire whorls heavilv ornamented

with 8 strong spiral threads between which are interca-

lated verv fine secondan' threads; bodv whorl (portion

exposed in matrix, just anterior of shoulder carina of ho-

lotvpe) ornamented with 5 (extrapolated from holohpe)

ven' large, thick, beaded spiral cords: aperture and si-

phonal canal unknown.

Material examined: 11()L()TYPE — Length (frag-

mentan'. missing portion oi body whorl and siphonal ca-

nal) 24 mm. width 16 nun, in back-reef lagoonal facies

of the Suwannee I'ormation, Terramar Pit, westernmost

Polk Colmt^■, Florida, UF75928: PARATY'PE-spire only,

width 16 nun, same localit\ as holotvpe, in research col-

lection of the author.

Etymology: "Gennned". in reference to tlie charac-

teiistic gemlike knobs on tlie shoulder c;nina.

Discussion: Sj)iiiijiil^iir ij^cininulatnin. although

known onK honi Iragnieutan specimens, is so distinc-

tive that its lormal description seems warranted at this

time. The new species is similar to slope-shouldered var-

iants ol the N'icksburg Oligocene Spinifulffir spini^cr

(Conrad, 1848) (such as the tmv illustrated bv MacNeil

(Gertiiiaiiii. 1969), lerigtli 24 inni. 30. Pnrlnirniiiiiiiiuiii iiumsficUli Petnch n.sp,. holotvpe (UF 75980), l<-iigtli 27 iiiiii. 31. Pcuriiria

(Didlii)uuTx) nifinipicolus (Dall. 1916), tragniciitaiy spccinieii, length 21 mm (missing portion of bodv whorl and siplional canal).

32. Clui-«r(u\ stctopus (de Gregorio, 1890), length 32 mm. 33,34. Solenosteira suutnutecnsis Petucli, n.sp.. holotvpe (UF 75983).

leiiirtli 27 mm.

Page 130

THE NAUTILUS. \ol. 110. No. 4

m^i

Figure

35,36.

IciiHtli

42,43.

Iciiiltli

s 3.5-49. Mcloiigeriidae, BiLsycoiiidue, X'olutichie, Volutoniitridac. uiid Turhim-llidae from the Siiwiuiiiff back-reef lagoons.

Mijrislica cra\sironuita (Conrad. 1848). length 49 nnn. 37,38. luilsili/ria kciidmvi Petuch, n.sp.. hoIot>pe {VF T59S5),

37 nnn. 39,40. FuUihjriu munsfiddi (l)all, 1916), length 32 mm. 41. Coiwiiiitm cirnulatn Dockeiy. 1984, length 21 nnn.

Tiirhinclla suwtiniicnsis (Mansfield, 1937), length 107 nnn. 44,4.5. Conoiiiitm kviulivui Petneh. n.sp.. holot\7x- (UF 75986),

14 inni. 46. Clriiiolitlics vickshuixen.sis (Conrad, 1848). length S9 nnn. 47. Spinif'uU^ur Hfiniiiiildtiiin I'etneh. n.sp.. holot\pe

E. J. Petuch, 1997

Paec 131

.nul Doeken, 19^4. plate 54. fifj;.H'. Iiut dirirrs in lae-king

distinct spines, having, instead, low ronnded knobs ar-

ranged in an undulating pattern. Spinijidour ^riiiiuiila-

tint) also differs from S. spinii^cr. and an\- other known

Sphiifulfiiir spi'cies. in being liea\ily sculptured with

wide, thick spiral cords on the boiK whorl. At present,

this highlv ornate species is known onK lidin the Su-

wannee Formation.

FamiK l-^uccinidae

SublamiK Buccininae

Genus SolcniKtcira Dall. 1890

Solcnostririi sinLdiiiiccn.'iis new species

( figures 33. 34)

Description: Shell of average size for genus, liroadiv

fusiform, inflated, with liigh. protracted spire; shoulder

angled, spire whorls sloping; bodv whorl with 7-S \er\

large, rounded, e\enl\-spaced axial folds; folds become

largest at shoulder producing large, rounded knobs at

shoulder periphen'; sutme prominent uutlulating as it

overlaps .slioulder knobs of spire whorls; bodv whorl or-

namented with 12 large, raised spiral cords; spire whorls

ornamented with 5 spiral cords; siphonal canal well-tle-

veloped, broad, ornamented with 3 large spiral cords;

pseudonmbilicus present, partially covered by parietal

shield of siphonal canal: aperture wide, oval in shape.

Material examined: HOLOTYPE — Length 27 nmi.

width f7 mm, in back-reef lagoonal facies (Sti/lo))lu>r(i

bioherm) of the Suwannee Formation, Terramar Pit,

westernmost Polk Count), Florida. UF75984; PARA-

T\TE — length 25 mm. same localitv' as holotv|ie. in the

research collection of th

ithor

Discussion: Solowsteiiyi fnnvdiiiiciiisis is the oldest-

known member (jf its genus to be found in Eastern

North America. The new species is most similar to S.

iiiiiniiilii Dall. 1890 from the Aquitanian Miocene Tam-

pa Member of the Arcadia Formation, but differs in iiav-

ing a lower, less protracted spire, in ha\ing a nnich

coarser shell sculpture composed of strong spiral cords,

and in having more numerous and smaller axial folds and

shoulder knobs.

Superfamilv Volutoidea

Family Tnrbinellidae

Subfamily \'asinae

Genus \'asiiin Roding. 1798

Vasiiin siiurinnceusis new species

(figures 48. 49)

Description: Shell small for genus, with broad, shaip-

Iv-angled shoulder; spire stepped, proportionally low.

with spire whorls slightK sloping; suture indented, bor-

dered by broad, nndnlatiug subsutnral collars; slioulder

ornamented with 10 large, evenly-spaced, open spines:

body whorl sculptiued with 8 large primarx' spiral i-ords:

smaller secondan spiral cords present betxveeu some

priman,- cords: spire whorls sculptured with 3 largi' spiral

cords; siphonal canal well tlexcloped. ornamented with

single row of veix' large, opi-n spines tliat correspond to

the spacing of the shoulder spines: siphonal canal sculp-

tured with 5 large spiral cords, 2 posterior of row of

spines and 3 anterioi of spines; aperture open, flaring;

columella with 3 \cr\ huge, promini'iit plications; on lio-

lotx'jje, a louitli Ncix laint plication is present between

anterior ;md media! plic;itious.

Material examined: HOLOTYPE— Length 39 nun,

wiiltli (tragmiMitaix, missing last third ol body whorl) 23

nun. in back-reef lagoonal facies {Sti/Iopliora bioherm)

of the Suwannee Formation, Terramar Pit. westernmost

Polk Countx-. Florida UF75984.

Etymology:

Florida.

Named for the Suwannee Formation of

Discussion: \'asiiiu siiwnnncensis is the first known

\ase shell to be collected in the Eastern American Oli-

gocene. The new species is most similar to \' humcro-

smu Vaughan. 189fS from the Jackson Eocene Moodys

Branch Formation, but differs in having a lower, flatter

spire, in having more luunerous primary spiral cords on

the body whorl, in having three veiy large and promi-

nent spiral cords on tlii' spire whorls, and in having a

larger, much more prominent, and more elevated snb-

sutural collar. The new species is also similar to V. sith-

capitclhnii Heilprin. 188fS from the ,\<juitanian Miocene

Tampa Member ol the Arcadia Formation, but differs in

having a lower. Hatter spire, in having finer liodv whorl

sculpture', and in having smaller, stumpier, and less de-

veloped shoulder and siphonal spines. As tar as is pres-

ently known. V. siiwiinnceiisis was restricted, ecological-

ly, to the coral bioherms of the Floridian Peninsula.

Family Volutidae

Subfamily Volutinae

Genus Falsilt/iia Pilsbn and Olsson, 1954

Falsili/ria krnclrcwi new species

(figures 37, 38)

Description: Shell of average size for genus, fusiform,

with higli. elevated spire; shoulder, particularly of body

whorl and pt'uultimate whorl, shaipiv -angled, producing

distinctiv sloping spire whorls: whorls ornamented with

8-1(1 strong, shaqilv-angled. evenly-spaced axial ribs;

shoulders, particularly oi body whorl and penultimate

whorls, with 8-10 large, shaqi, spinelike knobs, each cor-

responding to Ml axial rib: bodv whorl smooth, with 2-

(UF 7592S) (iragiiiciitarv. missing pi>rti(in nl

n.sp.. holotxpe (UF 7549S4), length oVJ mm.

l)()(l\ wliiui anil siphonal canal), length 24 nmi. 48,49. Viisiiin Miuciniwi'nsis Petuch,

Pajic 132

THE NAUTILUS, Vol. 110, No. 4

Fij;uri's 50-72. Mitridac, Mariiincllidac. Olividac. C.'Diiidac, Tnriidac, and (Aliclniidac Iroiii the Suwannee liaek-reef lat;oons. 50.

i-'nsimilra coiujuislu iCJonrad, 1S4S), len<j;tli .3.5 mm. 51. Ihjaliim siliiiflin in i ball. 19l(it. length 12 mm. 52,53. I'crsicitlri ilockcn/i

Peliich, n.sp., iioIot>pe (UK 75988), leni;th 14 nnii. 54,55. Pcrsindti inariuili I'etneh. n.s|)., holotNpe lUF 75989), lent;th 11 nmi.

5G,57. Pnmum jcssicac Petucli, n.sp., h()l()t\pe (UK 75992), lenijtli Hi i 58,.59. I'ninnm sandnic relneh, n.sp.. iiolotxpe (V¥

75993), length IS mm. 00,61. I'l-rsiruhi simnnncrn.si.t Petneh, n.sp., holotvpe {UK 75990) length 9 nun. ()2,()3. Dcntiiniin^o diilli

E. J. Petucli. 1997

Paec 1:33

3 lai"m' s[)iral cords aromul aiitciiiii' ciul; spire wlinrls

ornaiiu'iitfd (particularK' ovi last wliorji with siiii^lc, lariat'

spiral cord: aperture wide, Haiiiiu;: colmnclla with S

large, e\enlv-spaced, headed plications: protoconch pro-

poitioualK' ven' sinall, composed ol 2 narrow, smooth

whorls.

Material examined: HOLOTVFE — Length 37 mm.

witlth 21 nun. in hack-reef lagoonal facies (Stt/lophora

hioherni) oi tlie Suwannee Formation, Terramar Pit.

westernmost Polk Countw Florida, UF759.S5: PARA-

TYPE — length 39 mm, same locality as holot\pe, in the

research collection oi the author.

Et\Tnolog\': Named for Mr. Eric Kendrew of \'alrico,

Florida, who collected the holot\]ie at the Terramar Pit.

Discussion: FdlsiUjiia koiihvwi occurs together in the

Ti-rramar fauna with F iiuin.^fiilili (l^all. 1916) (figures

39, 40). hut can he readiK distinguished h'om its hetter-

kuown congener in having a consisteutK broaden more

inflated, and less elongated shell, with prominent spine-

like shoulder knobs, and in biiug a smoother shell that

lacks the spiral cord sculpture on tlu' bod\ whorl. Like-

wise, two Fahihjria species occm' togt'ther in the ri'efal

eu\ironments of the Burdigalian Miocene C^hipola For-

mation. Of these t^\•o, F. uuinsfuhli appears to be an-

cestral to the smooth-shouldered F. anoptos Hoerle and

X'okes, 1978 while F. kcndrewi appears to be ancestral

to tlie knobbed-shouldered F. pijcnopleura (Gardner,

1937). The smooth-shouldered and lu'a\il\-corded F.

inusiciiKi iHeilprin. ISSfi) from the .\(|uitanian Miocene

Tampa Member of the .\rcadia Formation is \erv similar

to F iiuinsfn'hli and is the most probable descendant

species. \o Falsilijria with strong shouldi'r knobs has \et

been tounti in the Tampa fauna. At present, Ftilsili/hii

kciulrcwi is known onl\- from the Terramar Stylophora

biohernis.

FamiK N'olntomitridae

Genus Couoiuitra Conrad. lSrS5

Couoinitra keiuliTwi new species

(figures 44, 45)

with

spire;

Description: Shell small tor gi'uus. biconic

shaqjiv-angled shoulder and proportiouallv low-

anterior t^^■o-tllirds of bod\ wiiorl smooth, unoruameut-

ed; spire whorls and posterior one-third ol bod\ uhoii

iiea\il\' sculptureil with large. e\euK spacetl rib-like pli-

cations, averaging 20 per whorl; aperture narrow; colu-

mell.i with 5 large plications, with the largest at posterior

i']]d. becoming smaller toward anterior end: anterior-

most tip of siphonal iuca ornaTuented with 4-5 im-

pressed spiral thrciids.

Material examined: HOLOT^'PE — Length 14 nun.

width 8 mm, in b;ick-rei'f lagoonal facies of the Suwan-

nee Formation, Teiramar Pit, westernmost Polk (>)unt\,

Florida, UF759Sfi; PAH.AT>TES— lengths 14 nun and

17 nun. same loc'alit\ as holot\'pe, in the research col-

k-ction ol the .iiithoi'.

Et%'niologv: Named lor Mr. Eric Kendrew of \alrico,

Floiitla. who collected the holot\pe at the Terramar Pit.

Discussion: This distinctive new Suwannee Conoiiii-

tni species is most similar to the svmpatrie and wide-

spread \icksburgian C cniudata Docken'. 1984. but dif-

fers in having a nuicli lowfr spire vvitli a sliaqilv -angled

shoulder and cone-shapetl appearance. Couomitra ken-

(Irewi also differs from C. cniudata in having shoulder

and spire plications that arc proportionally larger and

fewer in number. In these characters, the nevy species

is also similar to C. inodesia Docken, 1984 from the

\icksburg-ageil Forest Hill Formation of Mississippi,

but differs in having a much lower spire and much more

shaiplv-angled slionldei'.

Family Margint'llidae

Subfamily Cdabellinae

Genus Dentimargo Cossmann. 1899

Dcntimargo dalli nevy species

(figures 62, 63)

Description: Shell ol .i\er;ige length lor genus, slen-

der, witli high, elevated spire; suture indented, protluc-

ing slightlv stepped spire wiiorls; shoulder of bodv whorl

only slightly angled; aperture narrow at posterior end.

rapidly expanding and lu'coming flaring and opi'u at an-

terior end; columella with 4 large plications; labnun of

adult tliickened ;nid (.iillused. with smooth inner edge.

Material examined: IIOLOTYPE— Length 15 nun,

width 8 mm, in b;ick-reel lagoonal facies (sea grass bed)

of the Suwannei' Formation, Terramar Pit, vyesterumost

Polk Count\, Florida, UF 75987: PARATYPES— 2 spec-

imens, both 14 mm, in length, same localitv as holotvpe.

in the research collection o( the auti

lor

EtMnoIogv: Named in honor ol W illiam Heaiev Dall,

the foremost pioneer Floridi;ui paleonialacologist.

Discussion: Dciitiinargn dalli is tlie oldest known

member of its genus to be found in the Floridian Ce-

nozoic. The new siiecii'S is most similar to D. vlenantida

Petiicli. 11, s[).. liiilotvpe (UF 75987), leiigtli 15 nun: <J4. I'niiimii <rUiif Petucli, 11. sp,, Imliilvpc (UF 75991 ), leiigtii I 1 iiiiii. 65.

Oliia iOinofj^i/iniui) lirook.svillcnsis (Mansfield. 1937). iciigtli 19 iiiiii. 66. Olnclla i irkshurfj,c>tsis I^ockeiy, !9S4, length 10 iiini. 67.

Piciintfii.sia hrooksvillensis (Mansfield, 1937), length 26 iniii. 68. Piciirofnsia doiiliiif:,! Petucli, n.sp., holotvpe (UF 75995), length

26 mm, 69. Conorbis porcellanus (Conrad, 1S4S), length 36 nini. 70. Siiudiiitcscdphii lindac Petucli, n,sp., holotvpe (UF 75996),

lengtli 17 nun (t\pe of the new genus Simaniifscaphfi). 71. Snwaniic^cdplia lindiic Petuch, n.sp., view of spire of iioloKpe; 72.

Conns {A.sjircll(i) kcndrewi Petucli, n.sp,, lioldtvpe (UF 75994). length 35 iniii.

Paw 134

THE NAUTILUS. \ol. 110. No. 4

(Dall, 1890) iioni llic .\(|uitaiiiaii Miocene Tampa Meiii-

l)cr of the Arcadia Foniiation, hut (litU'is in being a larg-

er more slender and elongated .shell with a more sloping

shonlder and in haxing a nuich more piotracted. dis-

tiiK'tK scalariform spire. At present. Dcidiinan^o dalli is

known onK' irom the Snwannee Formation.

Genus Pcrsiculu Scinnnaeher 1S17

Persiaila dockcn/i new species

(figures 52,53)

Description: Shell large for genus, oxoid, inflated,

slightK' Hatteu(xl dorso-ventralK'; posterior extremities

well-developed, greatly projecting, recun-ed strongly to-

ward right side of shell; aperture narrow, widening

slightlv toward anterior end; posterior end of aperture

recuned sharjilv toward right side of shell; spire whorls

not \isible. buried in thick callus; columella with 5 large

plications: lahrum thickened, smooth along inner edge;

shaip-edged callus preseTit along posterior of columella,

forming open canal in sharpK -recuned posterior end of

apertme.

Material examined: HOLOTYPE — Length 14 mm,

width 9 mm, in back-reef lagoonal facies (open sand bot-

tom) of the Suwannee Formation. Terramar Pit, west-

ermuost Polk C'ount\-, Florida, iJV 75988.

Etymology: Named in honor of Dr. David T. Dockery

ill, of the Mississippi State Bureau of Ck-olog)', in rec-

ognition of his extensive and important contributions to

the systematics of the southeastern U.S. Paleogene mol-

lusks.

Discussion: Of the known Paleogene anil earlv Neo-

gene Prrsirula species, P. dochcn/i is Tuost similar to P.

proamvida (Gardner, 1937) from the Buriligalian Mio-

cene Oak Grove Formation of southern Georgia and

Tiorthwestern Florida. The new Oligocenc" species differs

Irom this possible Miocene descendant in biing a larger,

more elongated shell, in ha\ ing a fai' more recun-ed pos-

terior end of the apeitnre, and in having luuch better

developed, more conspiiiions, and more-projecting pos-

terior extremities. At present. I'lisicula ditckcriji is

know^n only from the Suwannee P'ormation.

PcrsicuUi iiifintcili new species

(figures 54. 55)

Description: Shell ol a\er;ige si/e for genus, elonga-

tely o\ate. with lelaliveU str;iight sides; extremities

rouniled. onl\' laintK' produced, not projecting be\ond

spire; aperture narrow throughout. slighlK recuived to

right side of shell at posterior end; columella with 5 pli-

cations; labrnm thickened, with smooth inner edge.

Niil. kite of the U.S. Geological Sune\. in recognition

ol his important eoTitributions to the s\stematics of the

Vicksburg mollusks.

Discussion: Pcrsiculu iiuiciicili is similar in appear-

ance to P. Iwlensis (Dall, 1916) from the contempora-

neous Flint Rixer Formation, but differs in Iwing a larg-

er, more elongated shell with distinctk straighter sides.

The new species is also similar to the \icksburgian P.

virkshtir<^cnsis Docken-, 1984 from the Mint Springs

Formation of Mississippi, but differs in being a umch

larger, more elongated shell with a lower and more

rounded spire.

Persicnlfi .'iiiwanuccnsis new species

(figures 60. 61)

Description: Shell of a\erage size for genus, extreme-

1\ t'longated and cxlindrical. with straight sides; aperture

narrow throughout, recuned to right side of shell at ex-

treme posterior end; spire area flattened: posterior ex-

tremitx on labial side better developed than posterior

extremit)' of columellar side, projecting be\ond plane of

spire; extreme posteriormost edge of columellar area

with thin, sharp-edged blade, producing open posterior

canal with labial projection; columella with 5 plications;

labrum thickened, with \en fine crenulations along in-

ner edge.

Material examined: HOLOTYPE— Length 9 mm,

width 5 miu. in back-reef lagoonal facies (sea grass

beds?) of the Suwaimee Formation, Terramar Pit, west-

ernmost Polk Gounty, Florida, UF75990; PARATYPE—

length 10 nun, same locality as holotype, in the research

collection of the ;iuthor

Etvmolog>': Named for the Suwannee Formation of

Florida.

Discussion: Pcrsicula \iiu(initcciisis is the most slender

;iiid elongated of all known Oligocene and Miocene Pcr-

.\icul(i species. This new Suwannee marginella is simiku'

onl\ to the sxnipatric /' iiuioicili, but diffei's in being

even narrower and more elongated, in liasing the pos-

terior end of the ;iperture being more recuned, and in

having the dislmrlK piiijectiug posterior labial extremit\'.

Material examined: HOLOTYPE— Length llm.

w idth 7 mm, in back-reel lagoonal facies (sea grass bed?

ol the Snwannee Formation, Ternnnar Pit. westennnost

Polk County Florida, UF 75989.

Etymology: N;uned in honor of l)i'. I''. Stearns Mac-

Genus Pniiniiii I leriiuannsen, 1852

Pill III I III criciii new sp(>cies

(figure 641

Descripti«>n: Shell small for genus. eloTig;iteK o\ate,

with high, pvramidal spire; shonlder slightK angled: ap-

erture nairow ;il posterior end. expanding rapidly toward

anterior end. becoming wide, flaring, and opi'ii; colu-

mella distinctk arcuate, with 4 large plications; labrum

) thickeiieil. smontli aioii',; inner edge.

Material examined: IK )LOTVPE— Length If mm,

w idth 6 mm, in b.uk-ieii lagoonal facies (open sand bot-

tom) of the Suwannee l'"oriiuitioii. Terramar Pit. west-

Petuch, 1997

Page 135

tTiiiiiost Pt)Ik Counts. Florida. UF7.5yyi: PAR.Ji-

TYPES — Lengtlis S nun and 9 mm. .same localit\' a.s lio-

lot\pe, in till- resfatc'li LoIk'ctioTi ol tlic autlioi-.

Etymolog^•: Named tor Ms. Erica Kendrew. danghter

ol Mi-. Erie Kendrew. ol N'alrico, Florida.

Discussion: Ol the three Fniiiiiiit species now known

from tlie Suwannee Formation, P. cricac is tlie smallest

and most slender in outline. In general size and shape.

P. ericae is most similar to F iiifccta iDall, 1915) from

the A([uitanian Miocene Tampa Meniher ol the Arcadia

Formation, hut differs in heing a more slender shell with

a w'i(k-r aperture. The new species mav he ancestral to

till' radiation ol small, high-spired Pnuuim species lonnd

in the' Tampa Member and including such distinctixc

tiixa as P. infccta. P. impagina (Dall, 1915), P posti (Dall.

1915), andF inepta (Dall, 1890). The small un-named.

high-spired PniniiDi illustrated bv MacNeil and Docken-

(1984, plate 35, figures 4,5. as "Margiitclla sp.?'") Irom

the Bvram Formation ol the Vicksburg C^roup is similar

to P criaic and ma\' be conspecific.

Pniituiii jcfisicdc new species

(figures 5fi, 57)

Description: Shell of avei'age size tor genus, cN'lindri-

cal in form, with distinctly angled shoulder: spire high,

protracted, slightlv scalaritorni; spire whorls and shoul-

der of b()d\' whorl sloping; aperture narrow, wiilening

onlv slightU" toward anterior end: columella straight, with

4 large plications: labrum (missing on holotspe, but pres-

ent on parat\pe) slightK thickened, smooth along inner

edge.

Material examined: HOLOT^'PE — Length 1(S nnii.

width 9 mm, in back-reef lagoonal iacies (open sand bot-

tom) of the Suwannee Formation, Terramar Pit. west-

ernmost Polk County. Florida. UF75992: PARA-

TYPES — lengths 14 nnn and 11 nnn (fragmentan).

same locality as holotvpe. in the research collection ol

the author.

Etvniologv: Named t(jr Ms. Jessica Kendrew, ilaugh-

ter ot Mr. Eric Kendrew. of Vahico. Florida.

Discussion: With its large size and high scalaritorm

spire. P. jcssicac is the most easilv-recognizable ol the

Suwannee Pninmn species. The new specii's is most

similar to, and is possibly the ancestor of the ,\<|nitaniaii

P silicata (Mansfield. 1937) from the Tampa member ol

the .'\rcadia Fonriation. Pniniimjessicac dilfers Irom the

early Miocene species in having a more inllated shell

with a more-angled shoulder, and a highei-. more pro-

tracted spire.

Pniniim mndrnc new species

(figures 58, 59)

Description: Shell ol average size for genus, inllated.

ovately subcvlindricak slionlder sli.upK -angled: spire

proportionalK low : spire whorls slightK sloping: aperture

narrow, widening slightly toward anterior end: columella

straight, with 4 large plications: labnnn thickened,

smooth along inner edge.

Material examined: HOLOTYPE— Length IS nnn.

width II iimi. in back-reel lagoonal Iacies (open sand

bottom) ol tin' Suwannee Formation. Terramar Pit,

westernmost Polk Countv, Florida. UF75993: PARA-

TYPES — three specimens, lengths 13 mm, 15 nun. and

19 nnn, same locality as holotvpe. in the research col-

lection ol the author.

Etymology: Named lor Mrs. Sandra Kendrew. wile ol

Mr. Eric Kendrew. ol X'ahico, Florida.

Discussion: Pniiuiiii sandrav is very similar to, and is

midoubtedlv the ancestor ol, the A(|nitanian Miocene P.

gregaria (Dall. 1915) from the Tampa Member ol the

Arcadia Formation. Both sliells liave tfie same large size,

lovy spire, and narrow aperture. The nevy species dillers

from its descendant, however, in being consistently a

wider and more inllated shell. Pniniiin sanclrac appears

to be the commonest marginellid in the Suwannee For-

maticjn at Terramar.

Superiamily Conoidea

Family Conidae

Genus C'())i(/s Linnaeus. 1758

Subgenus Aspnlla Schantuss, 1869

Conus (A.sprclla) kciidrcwi new species

(figure 72)

Description: Shell of average size for subgenus, elon-

gated, tapering, with straight sides; spire flattened, with

faintly stepped, slightlv projecting whorls; shoulder

sharply-angled, boundeil bv thick, lidgelike carina; ca-

rina present on spire whorls, producing depressed sca-

lariform appearance; bodv wlmii heavily sculptured with

(on unique holotvpe) 24 large, evenly-spaced, highly-

raised spiral cords: spire whorls ornamented witli 5 fine,

evenlv-spaced spiral thri'ads: aperture straight, narrow.

Material examined: HOLOTYPE — Length 35 mm,

width 21 mm. in b;ick-reef lagoonal facies (Stt/Iopliora

bioherm) ol the Suwannee Formation, Terramar Pit.

westernmost Polk ( loinitv. Florida. UF 75994.

Etymology: Named lor Mr. Eric Kendrew ol X'alrico,

Florida, who collected the holotvpe at the Terramar Pit.

Discussion: ('.(iiiiis kciiiln'ui, with its e.xtremelv heavy

corded bodv sculpture, wide shoulder carina, and de-

pressed scalarilorm spire, is the most distinctive cone

shell known from the Eastern American Oligocene. The

only contemporaneous cone that even remotely resem-

bles this new species is the svnipatric Suwannee and

Flint River C. (Lcptoronus?) cookci Dall. 1916. Conns

kiiiilirui differs from this more wide-ranging species in

having a much coarser and stronger spiral cord sculp-

ture, in having a distinctiv stepped spire, and in having

Pajie 136

THE NAUTILUS. \<.l. 110, No. 4

a nroiiiincnt shoulder faiiua tliat fxteuds Ijcyoiui the

IkkK' wliorl outline.

Famil\ Tiirridae

Suhlainil\- Turriculinae

CJenu.s ricurofiisid de (Jrcsjorio. 1890

Ph'tirofiisia (hncliiif^i new .specie.s

(figure 6S>

Description: Shell of average size for genus, elonga-

telv rusiloriii. with high, protracted, .scalarifonii spirc",

roiiiKled hod\, and tliiek. elongated siplional canal;

shoulder slightK' angled: suture indented; hotK' whorl or-

namented with 9 large, thick spiral cords that lieconie

thinner toward anterior; siplional canal oriiainenteil with

10-12 fine, thin spiral cords; suture houniled with single,

large spiral cord; sntural cord separated from shoulder

spiral cord hv wide, depressed gap, pioducing slightly

canaliculate appearance; spire whorls ornamented with

3 large spiral cords; aperture pniportionalK small. o\al

in shape.

Material examined: HOLOTYPE — Length 26 mm

width, in liac'k-reet lacies of the Suwannee Formation,

Terramar Pit, westernmost Polk Countw Floritla.

UF75995; PARATYPES-lengths 19 mm and 23 nun.

same localitx' as liolot\'pe. in the icseareh collection ol

the authoi'.

Et^■nlologv: Named for .Mr. William Dowiing of Lake

Woith. Morida. who assisted me at the Terramar Pit and

who collected the holot\|)e.

Discussion: Flctirofiisiii (louliuiii occurs together with

the congeneric P. hnxiksiilli'iisis (Mansfield. 1937){fig-

ure 67) at the Terranuu' Pit. hut is nex'cr as connnon.

The new species dilTers from the better-known P.

hrooksvilli'iisis in being a i-onsistentlv more heavilv-

scul[)tured shell with coarser and thicker spiral cords, in

lacking \arical undulations on the spire and bod\' whorl,

and in ha\ing a nmch narrower ga[) beh\'een the sutnral

cord and shoulder cmd,

( )rder ( iephaiaspidea

Snperiamilv ( Aliclmoidea

I'^amiK (!\lichnidae

(ieuus Siiufiiincsc(ijili(i new genus

Diaj^nosis: (Aclichnid Imlible shells ol greater-thau-

axcrage si/.e. wideK inlhited. o\ate. \er\ thin and Iragile;

spire llattened. planar in profile; siitinc extremek de-

nre.s.sed, [)roducing \('i"\ deep, wide, open c h.iunel (see

figure 71); edge o( shoulder along channel de\eloped

into \ei"\ thin, shaq), erect blade; bod\ \\li(irl smooth,

laintK sculptured with extremcK' numerous, \('n fine

s|)iral threads; aperture ven- wide and llaring, o\al in

sha])e; apical perforation absent, re])laced l)\ raised pro-

tocouch; anal notch well-developed, corres|ionding to

raised shoulder blade along spire clianiu>l.

Type species: Siiii iiiiiii sciijiliii I'uhIiic Petuch, new

species, Suwannee I*"oiuiatiou, earl\ Oligoceue ol Flor-

ida (figures 70,71 ), At present, known ouK Irom the Ter-

ramar Pit. Polk Oountx. Florida.

Et>'m<)logy: A combination ol 'Suwannee' and ".s(v/-

phd" (barrel).

Discussion: This unusual cxiichnid most closeK resem-

bles members of tlu' genus Scapliandcr Montlort. 1810.

paiticularK' in shape and size, but is inuuediatcK' sepa-

rable bv possessing the characteristic w idel\ canaliculate

spire. In luuing a flat, channeled spire. Siiutniiicscnphd

somewh;it resembles a giant version of the diminutive

c\iiclniid geims Aclnirinti Gra\. 1847. but differs in be-

ing a nuich more inflated, rotund shell and in having a

deeper ;iud much wider spire channi'l. Morphologically,

Suwiniiicsraplia combines the large size and inflated

shell form of Scaphdndcr and the flat, channeled spire

ol ArtcdfiiKi.

SinL(iiiius(/ij)liii liiuldi new species

(figures 70, 71 )

Description: As for diagnosis of genus.

Material examined: HOLOTYPE— Length 17 nun,

width I 1 nun, in back-reef lagoonal tacies (sea grass

beds?) ol the Suw;muee Formation, Terramar Pit, west-

ernmost Polk Countv, Florida, UF75996; PARA-

TYPES — lengths 19 mm, ami 21 nnn. same localitv as

the holotvpe. in the lesearch collection of the author.

Etvnn>logv: Named for mv wife, Linda |ovce Petuch.

who stoitaliv watched over our familv while I was off

e\[iloring the Suwannee Formation.

ACKNOWLEDCMFNTS

1 wish to thank Mr. Eric Kendrew of X'alrico, Florida,

for his generous donation ol research material collected

;it the Terramar Pit and for his valuable assistance in the

field. Thanks also go to Mr. William Dowiing of Lake

Worth. Floiida, lor Ins assistance in collecting in the Ter-

ramar Pit, Special thanks go to Mrs. |ude Turner. Pho-

tographv Lai) ;it Florida .Atlantic I'niversitv. for devel-

oping and [)rinting the photographs used here and to

Mrs. CAiithia Mischler. I')epartmeut of (n-olog)-, Florida

Atlantic Universitv. for tvping the mamist'ri[)t.

LITFHATUHE CITEH

Ca.sev, '1", !,, 1903. Notes on tlic {.iorn-.iil rollrctioii of \ icks-

liurg fossils. v\illi (Icscriptioiis ol new species. Proceedings

of the Acadeiiiv ol N.ilur.il Scii'iucs of Pliihnliipliia 55:

261-2S3.

(liiniail, '1'. .\, ISIS. Olisrnations ciii the Koci'ue lonnatioii,

.111(1 (l('srriptii)iis of iiiie liiindri'd and five new fossils ol

(ImI pcndif troni llic viciiiilv ol X'icksliurg, Mississippi.

Willi .III .qipiiiiliv |(iiiiii.il 111 llic \ca(lriiiv of Niiliinil Sei-

i-iirrs (il iinkiilclpliia. Seidiid Series lii!!4 I I l.vl,

I), ill. \\ II n)lli \ ( eiiliiliiiticiii to ihi' liivrrleliiate l''aiiiia

ol llic Oli'Mirciii Beds 111 I'linl Kivrr < leort^i;!. Proceed-

E. J. Pctiicli, 1997

I'ai

137

iiii^s 1)1 tile L'liited Stat<-s Xaliiiiial Mumuiii 5ll2lfS2l:

487-524.

MacNeil. F. S. ami D, T, Dockt-n, 1984. Limci ( )lit^i)cene

GastropDila. ScapliopocLi. and ( 'cplialopoila ol the N'icks-

biirg (Iroiip in Missis.'iippi. .Mississijipi Departnu'iit of

Natural Rcsiinrces (Bnreau ol CH-()li)t:;\ '. Bnllctin 124,

415pp

Man.sfiflil. W. C^. 1937. .\Iollnsks oi llic- T.nnpa and .Suwannee

Liniestone.s ol Florida. State ol Florida Department ol

Con.senation. Geologieal Bulletin 15. 334 pji.

Petueli, K. |. 19S2. Geographical Heteroeln'ons: (Contempo-

raneous (>)e\i.stence of Neogene and Reient Mollnsean

Fanna.s in the Americas. Palaeogeograpli\, Falaeoclima-

tolog\, and Paleoecolog)' 37:277-312.

Petncli, E. |. 1988. Neogene Historv of Tropical .\nieriean

Mollusks; Biogeographx' and Flvohitionan Patterns ol

Tropical Western Atlantic Mollusca. The < 'oas(al Educa-

tion anil Research Foimdation. ( lharlottes\ illc. \irginia.

217 pp.

\anglni. T. W. 1900. A Tertiar\ Coral Reel' Near Bauihridge.

Cleorgia. Science (New Series) 12; 873-875.

X'okes, E. H. 1992. Cenozoic Muricidae of the W'esteni .\tlan-

tic Region, Part IX. Ptcn/iiotiis, Ptnricrui. AspcUa. I)< r-

DWiniin'X, Calotrophon. Artintliolahin. mitl Alliliosa: .\d-

ditions and Corrections, Tulane Studies in (;eolog\ anil

Paleontoli)g\- 25(1-31:81-82.

APPENi:)l.\ I

CJa.stropoil.s From The Bai'k-HccI Lagooua! Facies Ol

The Suwannee Foniiatinn in The Terraniar Pit, Polk

Countv', Florida. Y = Widespread X'ick.slinrt^ian Species;

F = Species Also Known From The Flint Hi\i-r For-

mation: S = Species Known ()nl\ Frcuu Tin' Suwannee

Formation.

Trochidae

Calliostoiiiii siliciitiiiii M.uisfield. 1937 S

Turhinidae

AstriDii lAsli'dliiiiiil i>(>lkiii\i.\ I'etueh, n.S[). S

Potamididae

Piinizisimis kciiilrcwi Petuch, n.sp. S

Tvhscojihnn hlnckaiitcn-ims (Mansfield, 1937) S

Tch'sciijiiuiii liciiuniildciisis (Mansfield. 1937) S

Cerithiidae

Bittiiiiii ciisci/i MacNeil, 1984 V

Ccritliuirlinii nitixtilc (Dall, UJlfi) F

('rrilliiiiin iTiiiriiiiiiii} iitsitliitiiiit Dall. UJlfi F

('chlltiiiiii I new genus?) cooA'c; Dall. UJlfi I*"

('■(■^tiiiiictrilliiinu hnxiksvillciisis (Mansfield, 1937 1 S

Ccstiiiiictrilliiiiiii lirmakfHsi.s (Mansfield, 1937) S

Ccstuiun'rilliiinu pascociisis (Mansfield, 1937) S

Cisltiiiucritliiuin la^inatitiu (Dall, 1916) F

PrisDKiiTrithiuiti prisma Petuch, n.sp. S

Scniiiciia<ius imntJuifowtis MaeXeil 19S4 \'

Souivciiat^iis siliciiiiii ( i^all. 191fi) F

Modnlidae

Modulus lircotikciisis Mansfield. 1937 S

Turritellidae

Apiciilii IxnicHiic (Mansfield, 1937) S

Tonulii ciisciii (MacNeil, 1984) S

Tiiiiiild iiiis\is\ij)piciisis (Conrad. 1848) \'

Crepidniidae

C.dhjptnicd coiinidi MacNeil, 1984 \'

Stromhidae

Oiihaiiliix licni/iiidoi'iisis .Mansfield. 1937 F

Fieidae

F/r)(.s niississij)piciisis Com'ad. 1848 \'

Cassidae

Phaliinn rmlntiini ((Comad. 1848) \'

Personiidae

Distmsio ( lilii/sscniii I cnissidi-its C'onrad, 1848 Y

Naticidae

Aiiipiilliiiiipsis llintensis (Mansfield. 1937) F

Xatira cdsi-ip MacNeil, 1984 \'

Pachi/ci'oininiiii.i diilli Petuch, n.sp. F, S

Pdchi/ci'iiiiiinium iiidtisficldi Petuch, n.sp. S

Siiiiiin iuississij)j)ie}isis (Conrad, 1848) V

C\praeidae

Ci/i>rdc<>rliis kciiihrui Petuch, n,sp. (Y?) S

.Muricidae

Cliinirciis sMopits (de Gregorio, 1890) \"

Poiriciid (Ddlliiitiircx) nifintpicohts (Dall, 1916) F

PtiTipuifiis pnipcposti (Mansfield, 1937) S

TdJUijphis iiiississipp'uiisis (Certmann, 1969) \'

Bu.s\coniilae

Spimftdffiir •^iinmiildliun l\-tueh, n.sp. S

Bnccinidae

PdUdcera caseiji (MacNeil, 1984) Y

P(dlacerd i-ickshiir'^cnsis (Com'ad, 1848) \'

Solcnostcini siiuduiu'ciisis Petuch, n.sp. S

Melongenidae

Mi/hsticd crdssicotiuttd (Comad. 1848) \'

Turlnnellidae

Chnolitlics I irkshiii'ff'itsis (CJonrad, 1848) Y

Tiirl)iiiillii suwd}iii('iisis (Mansfield. 1937) S

Vi7.s7/;/( snudiinrciisis l\'tuch. n.sp. S

\'()lllti(Lie

ludsilipid kciidrcui I'etuili. n.sp. S

F(dsilip-i(i iiidiisficlili I Dall. 1916) F

N'olutomitridae

Coitdinili'd crrinildld Docket, 1984 \'

Cononiitrd kciidrcui Petuch, n.sp. S

Coiioiiiitrti stdiiiincd ((Com'ad, 1848) Y

Mitridae (Pleioptxgmatidae?)

Fusiiiiitra ciuujuistd (faini'ad, 1848) \'

Oli\idae

OliidiOiitoi^i/iiiiid I hniiiksiillcnsis (Mansfield, 1937) S

Oliiclld liicDdkciisis Mansfield. 1937 S

Olivclld iickshur;^ciisis iOoeken 1984 \'

Marginellidae

DcntiiiKir^Ht dnlli Petuch. n.sp. S

Hijdiiiid silicijliivia {DaW. 1916) F

Pcrsicidd dtickcn/i Petuch. n.sp. S

Pcrsiciild ludciicili Petuih. n.sp. S

Pcrsiculd suudiiiiccnsis Petuch, n.sp. S

Pniuuiii cricdc Petucli, n.sp. S

Pnniiiiu jcssicfie Petuch, n.sp. S

Pniiiiiiii sdiidrdc Petuch, n.sp. (\') S

Conidae

Coiius {Lcjiliicniius) cddkci ]~)all, 1916 F

Page 13S

THE NAUTILUS, \<)1. 110. No. 4

C.oniis (Asprclla) kcnilrcni I'ctiicli, ii.sp

Tcrebriclae

Tcrehra (TcrcbirltiiKi) diii-'Hini (Joiirad, 1

TurricUif

Coiiorlii.s porccllaniis (Conrad, 1S4S) V

Crassisjiiirlla lyoplciira (MacNcil, 1984

S

>S48 \'

y

.7 S

1.37) S

Euclatluurlla Uvcoakctisis Maiisfii'kl,

Plcurofusia hrooksiillcusis ( Mansfield

Ph'urofiisia (lowliiif^i Petncli. n..sp. .S

Plcurofusia phttonica (Ca.sew 1903) \'

Pli'unifusiii sciTdta (C^onrad, 184S) V

Poh/stini snhsiiHilis (Ca.sev. 1937) V

( A'lic'hnidae

SuiLdinwsraplui liiuluc Pctucii, n..sp. S

APPENDIX 2

.\dditional Suwannee Formation Ga.stropod.s Tliat Were

Reported B\- Mansfield (1937) and E. V'okes (1992) But

Were Not Collected (In Tliis Study) At The Terramar

Pit. V = Widespread N'icksburgian Species: F = Species

Also Known From The Flint River Formation: S = Spe-

cies Ktiowu Onl\' From The Suwannee Formation.

Fissurcllidae

"Dioihini cliipolanii (Dall)" (prohahK = D nnssissip-

picHsis (Conrad, 1848)) X

Cerithiidae

Cciithiuiii iiforiiumnni Lvell and Sowerhw 1S45 F

"i'lTitliiuiii suaaiiurcnsis" Mansfield, 1937

Turritcllidae

"Turriltlld cl. Iiiilriisis Dall" '= Apiculii ci. iKilciisis)

V

Xenophoridae

"XenopJiora roncliiilinjiliiini Horn" l= .V. Iiuiuilis

(Conrad. 1848)) '

Terei)ellidae

Tcrcbclluiu lienuiuclocn.'iis Mansfield, 1937 S

Cassidae

"Cassis sp." (= C. flinfcnsis Mansfield. 1940i F

Muricidae

Takia poiirlli (E. \'okes, 1992) S

Buccinidae (?)

"Latinis sp." (appears to be a Caulhanis sp.) S

V'olutidae

Liiria iiiississippicnsis Conrad, 1848 V

2 3 5 3 i' ; / 0'

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