JANUARY 31, 1983

THE

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ISSN 0028-1344

Vol. 97

No. 1

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The American Museum of Natural History

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THE

NAUTILUS

Volume 97, number 1 — January 31, 1983

ISSN 0028-1344

CONTENTS

Kathe Jensen and Kerry B. Clark

Annotated Checklist of Florida Ascoglossan Opisthobranchia 1

Ralph W. Taylor

A Biographical Sketch of William Irvin Utterback (1872-1949) 13

Emile A. Malek

The South American Hydrobioid Genus Idiopyrgus Pilsbry, 1911 16

Leslie Hubricht, Ronald S. Caldwell and John G. Petranka

Vitrinizonites latissimus (Pulmonata: Zonitidae) and

Vertigo clappi (Pupillidae) from Eastern Kentucky 20

George P. Hoskin and Anders Waren

Peasistilifer edulis, a New Eulimid Prosobranch, Parasitic

on an Indo- Pacific Holothurian 23

Philippe Bouchet and Serge Gofas

"Terebra" cusentini Philippi, 1836, an American Columbellid Species 26

M. G. Harasewych

A New Species of Columbarium (Gastropoda: Muricacea)

from off Eastern Australia 28

W. F. Ponder and S. J. Hall

Pelycidiidae, a New Family of Archaeogastropod Molluscs 30

Edward M. Stern

Depth Distribution and Density of Freshwater Mussels (Unionidae) Collected

with SCUBA from the Lower Wisconsin and St. Croix Rivers 36

Mark E. Gordon

A Pre-European Occurrence of Glebula rotundata (Bivalvia: Unionidae)

in Arkansas 42

Recent Deaths 43 Reviews 44

Freshwater Snails of Africa and their Medical Importance

bv David S. Brown

For the first time:

A comprehensive account of freshwater snails

in Africa and neighbouring islands.

An exhaustive guide to species

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Vol. 97(1)

January 31, 1983

THE NAUTILUS

ANNOTATED CHECKLIST OF FLORIDA

ASCOGLOSSAN OPISTHOBRANCHIA

Kathe Jensen and Kerry B. Clark

Department of Biological Sciences .

Florida Institute of Technology

Melbourne, FL 32901

ABSTRACT

Distribution records are presented for 25 species of Ascoglossa inollusks in

Florida, including 8 previously unknown for Florida. About 35 species of

Ascoglossa occur in Florida, including undescribed species. The Florida fauna

contains significant contributions from the temperate west Atlantic fauna as well

as the Caribbean fauna. Habitats, diet, and collection data and techniques are

described, brief taxonomic summuries presented, and comparisons drawn be-

tween tropical and temperate populations.

The opisthobranch fauna of the Caribbean

province is relatively well-known, due largely to

the works of Ernst and Eveline Marcus and

others (Engel, 1925, 1927; Verrill, 1900, 1901;

Morch, 1863; Edmunds, 1963, 1964). A species

list and summary of records of Western Atlantic

opisthobranchs including some 20 species of

Ascoglossa from Florida has been published by

Marcus (1977). Also, Marcus (1980) published a

summary of the Western Atlantic species of

Elysiidae, including 13 species from Florida.

However, distributions of species within Florida

are poorly known, with the majority of collec-

tions only from south Florida. Further, informa-

tion about the microhabitats of most species,

necessary to effectively collect these species, is

virtually non-existent, and many species are

known only from preserved material. This pau-

city of data is especially pronounced in the Asco-

glossa, a group well-represented in Florida.

During our extensive field collections and lab-

oratory work with Ascoglossa (Clark & Goetz-

fried, 1978; Clark & Busacca, 1978; Clark et at.,

1979; Stirts & Clark, 1980; Jensen, 1980a, b;

Clark & Jensen, 1981; Clark et ai, 1981; Jensen,

1981a, b; Weaver & Clark, 1981; Clark, 1982;

Jensen, 1982, in press a, b; and several papers in

preparation), we have come across several

species previously unrecorded from Florida.

Also, we include new records for species

previously reported from Florida. Collection

data for these species will be presented in the

present study, along with taxonomic

ecological notes on some of the species.

and

MATERIALS AND METHODS

Most species were collected with suction col-

lectors (Clark, 1971) while snorkeling in shallow

water, by vigourously shaking handfuls of algae

to dislodge animals. Others were found by main-

taining clumps of algae in lighted laboratory

aquaria. After 1 or 2 weeks, animals that were

FIG. 1. Location of study sites in Florida,: 1) Haulover

Canal, Banana River; 2) Indian River, Melbourne: 3) Sebas-

tian Inlet: J,i Fort Pierce Inlet: 5) Upper Florida Keys; 6)

Lower Florida Keys; 7) Tampa Bay.

THE NAUTILUS

January 31, 1983

Vol. 97(1)

too small to see at the time of collecting grew to

visible size.

Fig. 1 shows the locations where we collected

the ascoglossans mentioned in this study. Col-

lecting data for individual species will be men-

tioned in the following section.

Drawings of entire animals were made with

camera lucida from live specimens relaxed in 8%

MgCij.GHiO in seawater. Radular mounts were

made as described by Jensen (1981a).

Development patterns are as classified by

Bonar (1978): Type 1 (planktotrophic); Type 2

(lecithotrophic, metamorphically competent at,

or soon after, hatching); and Type 3 (encapsu-

lated metamorphosis).

LIST OF SPECIES'

OXYNOACEA

Ascobullidae

1. AsrnhHlln ulla (Marcus & Marcus, 1970)*

Lobigeridae

2. Lobiger souve7'bii Fischer, 1856*

Oxynoidae

3. Oxynue antillarum Morch, 1863*

4. Oxynoe azuropunctata Jensen, 1980

Juliidae

5. Bertkelinia caribbea Edmunds, 1963*

Elysiacea

Elysiidae

6. Elyaia canguzua Marcus, 1955**

7. Elysia cauze Marcus, 1957*

8. Eiysia chlorotica Gould, 1870*

9. Eiysia evelinae Marcus, 1957*

10. Eiysia ornata (Swainson, 1840)*

11. Eiysia papulosa Verrill, 1901

12. Eiysia patina Marcus, 1980*

13. Eiysia picta Verrill, 1901 ( = E. duis)

14. Eiysia serca Marcus, 1955*

15. Eiysia tuca Marcus, 1967*

16. Tridachia crispata Morch, 1863*

17. Bosellia corinneae Marcus, 1973

18. Bosellia marcusi Marcus, 1972*

19. Bosellia mimetica Tnnche?,e, 1891*

Caliphyllidae

20. Cal I phylla mediterranea Costa, 1867**

21. Cyerce antillensis Engel, 1927*

''new distribution within Fluriiia; "new rei-oni Inr Flnrida.

22. Mourgona germaineae Marcus & Marcus,

1970**

Hermaeidae

23. Costasiella lilianae (Marcus & Marcus,

1969)**

24. Hermaea cruciata Gould, 1870*

25. Apiysiopsis zebra Clark, 1982

Stiligeridae

26. Placida kingstoni Thompson, 1977**

27. Ercolania coerulea Trinchese, 1893*

28. Ercolania funerea (Costa, 1867)*

29. Ercolania fuscata (Gould, 1870)**

30. Stiliger fuscovittatus Lance, 1962**

31. Stiliger vossi Marcus, 1960

Habitat Characteristics

Haulover Canal, Titusville, Brevard Co.: This

region connects the northern Indian River La-

goon with Mosquito Lagoon, but lacks a direct

oceanic connection. Typical annual salinity and

temperature ranges are about 20-30°/oo and

17-31°C (Gilbert & Clark, 1981), but tempera-

tures as low as 6°C occur in some winters. The

substrate is predominantly fine sand, with abun-

dant seagrasses (Syringodium and Halodule)

and a seasonal drift algal community.

Sebastian Inlet, Indian River Co., and Fort

Pierce Inlet, St. Lucie Co.: Collections from

these areas are mostly from artificial rock-jetty

substrates, which are dominated by Caulerpa

raceTnosa. At Sebastian Inlet, this alga is

seasonal, from about April-November, but C.

racemosa occurs nearly all year at Fort Pierce.

Fort Pierce frequently incurs heavy surf, but

the Sebastian Inlet site is somewhat protected

by a parallel jetty. Salinity and temperature at

Fort Pierce are essentially oceanic, but seasonal

runoff can cause rapid and extreme dilution at

Sebastian Inlet, which often causes a temporary

die-back of Caulerpa at the end of summer.

Florida Keys: Substrates vary from calcare-

ous silt to rock, and usually support a rich

assemblage of siphonalean algae and Thalassia.

rarely Halodule or Syringodium. Salinity usual-

ly varies around 35-37°/oo, and we have

measured temperatures as low as 12°C (Feb.) to

35°C (September). Although absolute distance

from the Upper to Lower Keys is not large (270

km), the progressively more oceanic location of

Vol. 97(1)

January 31, 1983

THE NAUTILUS

the Lower Keys makes them somewhat more

tropical. Most habitats in the Keys are well-

sheltered from waves by offshore reefs. Borrow

pits (roadside areas dredged as a source of fill

for highway construction) and tidal canals pro-

vide areas of higher nutrient levels, and often

provide especially rich collecting areas.

1. Ascobulla ulla (Marcus & Marcus, 1970)

Synonyms: Cylindrdbulla ulla Marcus & Mar-

cus[ 1970.

New records: Fort Pierce Inlet; Key Largo.

Previous records: Key Biscayne (Marcus,

1972a); Brazil (Marcus &'Marcus,' 1970).

Habitat: Burrows in sandy substrate sur-

rounding rhizomes of Caulerpa spp. At Fort

Pierce Inlet, it occurred on Caulerpa racernosa.

at Key Largo usually with C. paspaloides. In

aquaria, large animals frequently climb up into

assimilators of Caulerpa. but usually burrow

near rhizoids.

Food: Caulerpa spp. (Jensen, 1981a).

Development: Type 3 (Clark & Jensen, 1981).

Taxonomic remarks: The taxonomic position

of this specie? has been previously discussed

(Jensen, 1981a).

2. Lobiger souverbii Fischer, 1956

Synonyms: L. sagamiensis Baba, 1952 (see

Baba, 1974).

New records: Fort Pierce Inlet; Sebastian In-

let.

Previous records: Brazil (Marcus, 1957);

Curasao; Guadeloupe; Florida (Marcus & Mar-

cus, 1967); Puerto Rico (Warmke & Almodovar,

1972); Barbados (Marcus & Hughes, 1974);

Jamaica (Thompson, 1977); Costa Rica (Hou-

brick, 1968); Baja California, Mexico (Sphon,

1971); Galapagos (Sphon & Mulliner, 1973);

Hawaii (Kay, 1964); Japan (as L. sagamiense,

Baba, 1952) (Baba, 1974).

Habitat: On Caulerpa racernosa on relatively

exposed localities.

Food: Caulerpa racernosa. Feeding could not

be observed directly because the animals always

crawled to the side of the alga facing away from

the light source. However, emptied algal seg-

ments, which could only have been made by L.

souverbii, have been observed on the Caulerpa,

and growth was detected when the animals had

only this alga to feed on. L. souverbii tested the

food algae with the lobular extensions of the

rhinophores as well as with the oral lobes.

A peculiar characteristic of the radula of this

species is that the oldest teeth (in the ascus and

some in the descending limb) have bristles on

their flanges, whereas the younger teeth (some

in the descending and all in the ascending limb)

are smooth (Jensen, 1980a). In a juvenile speci-

men (shell length 1 mm), however, all teeth in

the radula had the bristles. Since both juveniles

and adults feed on C. racernosa. it must be con-

cluded that the bristles must be of help to the

small animals, which probably have less muscu-

lature in the pharynx and esophagus.

Color patches on the upper surface of the

parapodia vary from very pale pink in some

specimens to brown in others.

3. Oxynoe antillarum Morch, 1863

Synonyms: 0. aguayoi Jaume, 1945.

New records: Fort Pierce Inlet; Sebastian In-

let.

Previous records: Key Biscayne (Marcus &

Marcus, 1970); Virgin Islands "(Morch, 1863);

Brazil; Curasao; Atlantic side of Panama (Mar-

cus & Marcus, 1970); Puerto Rico (Warmke &

Almodovar, 1972); Barbados (Marcus & Hughes,

1974); Costa Rica (Houbrick, 1968); Jamaica

(Thompson, 1977); Cuba (as 0. aguayoi. Jaume,

1945).

Habitat: On Caulerpa racernosa at relatively

exposed localities.

Food: Caulerpa racemosea (Jensen, 1981a).

Development: Type 1 (Clark & Goetzfried,

1978; Clark & Jensen, 1981). This distinguishes

0. antillanmi from 0. azuropunctata Jensen,

1980 from the Florida Keys. The latter species

has type 2 development (Jensen, 1980b;. Clark &

Jensen, 1981), and occurs mostly on C. paspa-

loides in relatively sheltered habitats (Jensen,

1980b).

Taxonomic remarks: see Jensen, 1980b,

1981a.

5. Berthelinia caribbea Edmunds, 1963

New records: tidal canals emptying into Card

Sound, Key Largo; Deepwater Cay, Bahamas.

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January 31, 1983

Vol. 97(1)

Previous records: Jamaica (Edmunds, 19()3;

Grahame, 19(i9; Thompson, 1977); Panama

(shells); Brazil (Meeder & Moore, 197ii); Puerto

Rico (Warmke, 1966); Spanish Harbor Key,

Florida (Moore & Miller, 1979).

Habitat: Caulerpa verticillata growing on

mangrove roots or directly in muddy bottom of

tidal canals; uncommon. Most easily found in

late autumn, when reduction of C. verticillata

concentrates animals; not seen in field collec-

tions; found only by aquarium culture of algae.

Food: Caulerpa verticillata (Edmunds, 1963;

Grahame, 1969).

Development: Type 2 (Grahame, 1969; Clark

& Jensen, 1981).

6. Elysia canguzua, Marcus, 1955

New records: Fort Pierce Inlet.

Previous records: Brazil (Marcus, 1955). This

is the first record since the original description.

Habitat: On Bryopsis plumosa on exposed

outer jetty, and on Codimn sp. inside inlet in

strong current.

Food: Bryopsis plumosa and Codium sp. Mar-

cus (1955) sketched how they pierced the tip of a

utricle of Codium and sucked out the contents.

When feeding on Codium sp., it seemed like E.

canguzua tried to swallow the whole tip of a utri-

cle by spreading the oral lobes and buccal tube

around it. In the laboratory they would also feed

on Bryopsis plumosa and Chactomorpha sp. In

fact, small animals seemed to prefer Bryopsis

over Codium.

Development: Type 1.

7. Elysia cauze Marcus, 1957

New records: Fort Pierce Inlet; Sebastian In-

let.

Previous records: Brazil, south Florida (Mar-

cus, 1977).

Habitat: Usually in sheltered areas such as

tidal canals in south Florida, on a variety of

Caulerpa species; especially common on C. ver-

ticillata as juveniles; occasionally collected on

jetties, but only in sheltered areas. We have

recently separated three species which closely

resemble E. cauze; one feeds on C. racemosa in

the same habitats as E. cauze, hence is very dif-

ficult to separate from it, but the other two eat

C prolifera and Penicillu^s dumetosus, respec-

tively. All four species have the distinctive black

margin on the parapodium, which has previously

characterized E. cauze.

Food: Caulerpa spp. (Clark & Busacca, 1978).

Development: Type 3, possibly type 2; the

type 1 development reported by Clark et al.

(1979) appears to belong to the undescribed

species mentioned above, and probably does not

occur in E. cauze.

8. Elysia chlorotica Gould, 1870

New records: Sebastian Inlet; Indian River at

Melbourne and at Haulover Canal, Titusville.

Previous records: Chesapeake Bay (Marcus,

1972b); Noank, Conn. (Clark, 1975); Minas

Basin, Nova Scotia (Bailey & Bleakney, 1967);

New Jersey (Franz, 1968); Cape Cod (Russel,

1964); Texas (Boone, 1982).

Habitat: On Chaetomorpha sp. in sheltered,

estuarine areas.

Food: Chaetomorpha sp. (Clark, 1975; West,

1977); Cladophora sp. (Clark, 1975; Franz,

1968); Vaucheria (Clark, 1975; Russel, 1964;

West, 1977).

Development; Type 1 (Clark, 1975).

9. Elysia (Telinae Marcus, 1957

New records: Key Largo; Fort Pierce Inlet;

Haulover Canal.

Previous records: Miami (Marcus & Marcus,

1967); Brazil (Marcus, 1957).

Habitat: Shaded, sheltered areas ("under

rocks") with dense growths of centric diatoms

(Key Largo), or among epiphytic diatoms on

Bryopsis spp. or Caulerpa spp. also in shaded

areas (Fort Pierce Inlet). We first collected E.

evelinae in a tidal canal on Key Largo. It was

found on mixed algae, mostly Caulerpa verticil-

lata, but only one or two specimens were col-

lected at a time. After discovering that E.

evelinae fed on diatoms (Jensen, 1980a, 1981a),

it became easier to locate the right habitat, and

thus to collect more specimens.

Food: Biddulphia sp. (Jensen, 1981a).

Development: Type 2, occasionally type 3

(Clark & Jensen, 1981). The two types of devel-

opment are often found within the same egg-

mass. Most larvae hatch as large veligers

Vol. 97(1)

January 31, 1983

THE NAUTILUS

(length approx. 210 /^m) with eyes. They swim

little; most of the time they crawl around testing

the substrate. Often a few veligers in each egg-

mass stay behind and complete metamorphosis

within the egg capsule. It is not known what fac-

tors determine how many larvae have the latter

type of development, but it seems likely that the

feeding conditions of the parents are important,

because egg-masses laid by starved parents

usually all would hatch as type 2 veligers.

Development time at room temperature (ap-

prox. 20°C) is 12 days to hatching and 12-14

days till metamorphosis is completed.

10. Elysia omata (Swainson, 1840)

New records: Sebastian Inlet; Fort Pierce In-

let; patch reef off Windley Key.

Previous records: Miami (Marcus, 1972a,

1980); Jamai'^a ^Thompson, 1977); Barbados

(Marcus & Hughes, 1974); Cura^'ao; Tobago; St.

Vincent; Bermuda (Verrill, 1901; Marcus & Mar-

cus, 1970); Hawaii; Vietnam; Australia (Pease,

1860; Risbec, 1956; Thompson, 1973).

Habitat: On Caulerpa racemosa and Bryopsis

plumosa (FPI), and on Bryopsis sp. (Windley

Key).

Food: Bryopsis spp. (Jensen, 1981a).

Development: Type 1. Egg-mass generally has

2 embryos per capsule as in E. maoria (Reid,

1964); some extrazygotic yolk material is pre-

sent.

Taxonomic remarks: The status of this species

has been widely discussed (Marcus & Marcus,

1963; Thompson, 1973, 1977; Marcus, 1977).

The teeth of the present specimens were smooth

and approximately 150 ^^m long, thus resembl-

ing both the Australian material (Thompson,

1973) and Jamaican specimens (Thompson,

1977).

12. Elysia patina Marcus, 1980

New records: Key Largo.

Previous records: "Florida Keys".

Habitat: Common on Udotea spp. in very shal-

low water of tidal canals. This species prefers

eutrophic areas with moderate tidal flow.

Food: Presumably Udotea.

Development: Type 1.

14. Elysia serca Marcus, 1955

Synonyms: E. clena Marcus & Marcus, 1970

(Jensen, in jiress a).

New records: Long Key; Snake Creek Canal

(Windley Key); Point Elizabeth and tidal canal

on Key Largo; Fort Pierce Inlet; Banana River

at Rt. 520; and Indian River at northern end of

Merritt Island (Jensen, in press a).

Previous records: Brazil (Marcus, 1955;

Horsoe, 1956); Curasao (Marcus & Marcus,

1970, asE. clena); Florida (Marcus, 1972a, asE'.

clena); Barbados (Marcus & Hughes, 1974, as E.

clena). Elysia catulus Gould, 1870 has not been

found in Florida, contrary to a statement by

Marcus (1980). The southern limit of E. catulus

probably coincides with that of eelgrass, Zostera

marina L., in the Carolinas.

Habitat: Seagrass beds, preferably containing

Halophila engelm.anni. In the Florida Keys E.

serca is most often found on Thalassia tes-

tudinum.

Food: Seagrasses, i.e. Halophila engelmanni,

Halodule wrightii, and Thalassia testudinum

(Jensen, in press a).

Development: Type 1 (Clark & Jensen, 1981).

Taxonomic remarks: Will be discussed by

Jensen (in press a).

15. Elysia tuca Marcus, 1967

New records: Florida Keys, Fort Pierce Inlet.

Previous records: Curasao; Elliott Key; Big

Pine Key; Brazil (Marcus & Marcus, 1963, 1970);

Puerto Rico (Warmke & Almodovar, 1972); Bar-

bados (Marcus & Hughes, 1974); Jamaica

(Thompson, 1977).

Habitat: Almost always associated with Hali-

meda spp., less frequently on Penicillus durne-

tosus (Lamouroux) Blainville. Occasionally

found crawling on coral rocks, on Caulerpa spp.,

or on Thalassia testudinum (on which the eggs

are often deposited). This is probably the most

common ascoglossan in the Florida Keys. The

depth range of this species is very broad, from

high subtidal to c. 30 m at Deepwater Cay,

Grand Bahama. At Fort Pierce, the species oc-

curs in heavy surf on Halimeda discoidea, but

prefers H. incrassata in still waters of the

Florida Keys.

6

THE NAUTILUS

January 31, 1983

Vol. 97(1)

Food: Halimeda spp., Caulerpa spp., Avrain-

villea nigricans, and Udotea sp. (Clark & Busac-

ca, 1978). Halimeda spp. is the preferred food of

this species (Stirts & Clark, 1980). Its pedal

lobes are very weakly developed, and the ven-

tral side of the head is almost continuous with

the foot. This is probably an adaptation to feed-

ing on the flat segments of Halimeda spp. Like

Lobiger souverbii, this species crawls away from

the light source when placed under the dissec-

tion microscope, and thus feeding could not be

directly observed.

Development: Type 2 (Clark & Goetzfried,

1978; Clark & Jensen, 1981).

16. Tridachia crispata Morch, 1863

Synonyms: Elysia crispata.

New records: Key Largo; Windley Key; Long

Key; Big Pine Key; Geiger Key.

Previous records: Venezuela to Columbia;

Honduras; Florida; Antilles and Barbados (Mar-

cus & Marcus, 1967; Marcus, 1976, 1977);

Jamaica (Thompson, 1977); Curasao; Bonaire;

Haiti (Marcus & Marcus, 1970, Thompson,

1977).

Habitat: Often found straying away from any

algae, on bare sand or coral rock ("under

rocks"); not uncommon at moderate depths (on

coral reefs). Sometimes associated with Cauler-

pa verticillata. The preferred habitat appears to

be highly eutrophic areas (i.e. borrow pits,

marinas) with dense algal growth, as the

densest populations and largest animals occur

there. Reef animals may wander far from visible

algae, but apparently do require periodic feed-

ings to restore chloroplasts. This species ex-

hibits a pronounced daily activity cycle; usually

emerges from crevices in late afternoon, disap-

pearing in mid-morning, depending on light in-

tensity (Weaver & Clark, 1981).

Food: Caulerpa verticillata is the principal

food of juveniles, but adults will also feed on

Halimeda, Bryopsis, Cymopolia. Penicillus. and

Batophora (Clark & Basacca, 1978; Jensen,

1980a).

Development: Type 2, occasionally type 3

(Clark & Jensen, 1981).

Remarks: Juvenile animals closely resemble

Elysia. They are green with black heads.

smooth parapodia and short rhinophores. The

parapodia of juveniles (Fig. 2) do not meet at the

neck, and are smooth-margined, not ruffled.

Separation of certain species of Tridachia based

on these ontogenetically variable characters

may be unsupportable.

18. Bosellia marcttsi Marcus, 1972

New records: Hens and Chickens Reef off

Windley Key; Borrow pit. Grassy Key.

Previous record: Key Biscayne, FL.

Habitat and Food: Halimeda opuntia (L.)

Lamouroux; this diminutive animal (3 mm)

clings tightly to the alga, requiring vigorous

shaking to dislodge it, and is easily overlooked

because of its size. Uncommon.

Development: unknown; the few large follicles

(Marcus, 1973) suggest type 2 or 3.

19. Bosellia mimetica Trinchese, 1891

New records: Fort Pierce Inlet.

Previous records: Mediterranean (Portman,

1958); Florida (?Miami) (Marcus & Marcus,

1970; Marcus, 1973); Barbados (Marcus &

Hughes, 1974); Brazil (Marcus, 1977).

Habitat: On Halimeda discoidea on very ex-

posed rocks in strong surf. Difficult to shake

loose from algae; slugs are nearly invisible in

situ.

Food: Halimeda spp. (Portman, 1958; Jensen,

1981a).

Development: Type 1 (Clark & Jensen, 1981).

20. Caliphylla mediterranea Costa, 1867

New records: Fort Pierce Inlet; Hens and

Chickens Reef, Windley Key.

Previous records: Virgin Islands (Engel,

1927); Brazil (Marcus & Marcus, 1970); Mediter-

ranean (Gascoigne, 1979).

Habitat: On Bryopsis phmiosa on sheltered/

shaded side of rocks on exposed coast.

Food: Bryopsis plumosa (Gascoigne, 1979;

Briiel, 1904).

Development: Type 1 (Clark & Jensen, 1981).

Taxonomic remarks: Based on morphological

comparisons with the description given by

Gascoigne (1979), the present material may

represent a different species from the Mediter-

ranean.

Vol. 97(1)

January 31, 1983

THE NAUTILUS 7

FIG. 2. A, Juvenik Tridaehia crispata; B, Mourgona germainae; C, Placida kingstoni; D, Riyhl

anterior o/P. kingstoni; E, pcro-s o/'P. kingstoni; F, penial stylet o/P. kingstoni. Scale bar in F =

lOfjim; all other scale bars = 1 mm: m- = vaginal ridge; ov = oviducai aperture: p = penis; st = stylet;

d = duct.

8 THE NAUTILUS

January 31, 1983

Vol. 97(1)

21. Cyerce nnfillenfiifi Engel, 1927

New records: Fort Pierce Inlet; Key Largo;

Windley Key.

Previous records: Virginia Key; Elliott Key

(Marcus & Marcus, 1967); Curasao; Puerto Rico;

Tobago (Marcus & Marcus, 1970); Jamaica

(Thompson, 1977).

Habitat: Most often on Penicillus spp.; occa-

sionally on Halimeda discoidea (FPI). Often in

areas with strong currents.

Food: Penicillu.-i spp., Udotea Jlabellum,

Halimeda spp. (Jensen, 1981a).

Development: Type 2 (Clark & Goetzfried,

1978; Clark & Jensen, 1981).

Taxonomic remarks: The teeth of the Florida

specimens are considerably smaller than those

of the Jamaica specimens (Thompson, 1977),

and also the bases of the teeth are relatively

shorter in the Florida specimens (see also Mar-

cus & Marcus, 1963, 1970). The figure of "Cyerce

cristallina (Trinchese)" shown by Marcus &

Marcus (1967) is actually C. antillensis; to our

knowledge, C. antillensis is the only Cyerce

known from Florida.

22. Mourgona germaineae Marcus & Marcus,

1970

New record: Geiger Key, near end of Key

West Naval Air Station Runway.

Previous record: Puerto Rico (Marcus & Mar-

cus, 1970).

Habitat: Exclusively on dense growths of

Cymopolia barbata in very warm, relatively

sheltered, shallow water. We have been unable

to find this species in more northerly popula-

tions of Cymopolia. but these generally regress

completely in winter. Occurrence within the

Cymopolia zone (immediately shoreward of the

Sargassum zone on hard sand/rock bottom) is

sharply demarcated, suggesting water tempera-

ture, and especially the thermocline, may con-

trol recruitment of this species.

Food: Cymopolia barbata (Jensen, 1981a).

Development: Type 1 (Clark & Jensen, 1981).

Eyes are present in newly-hatched larvae, but

the larvae are morphologically type 1 in other

respects, and could not be induced to meta-

morphose on C. barbata.

Taxonomic remarks: This species is probably

synonymous with Cyerce edmundsi Thompson,

1977 from Jamaica. Thompson mentioned that

the cerata of C. edmundsi contained branches of

the digestive gland, and the teeth closely resem-

ble those of M. germaineae. Unfortunately no

mention was made of the presence or absence of

a large pharyngeal crop, the absence of which

characterizes the genus Mourgona (Marcus &

Marcus, 1970), but the presence of digestive

diverticula in the cerata clearly separates it

from the genus Cyerce (See Swennen, 1961).

Our animals varied from whitish to dark

green, depending on the state of the digestive

diverticula. The diverticula appear as lobular

branches in the flat cerata (Fig. 2). The cerata

are highly adhesive and appear to detach defen-

sively; other slugs die when kept in the same

container as this species, suggesting that it ex-

cretes toxins.

23. Costasiella lilianae (Marcus & Marcus,

1969)

Synonyms: Stiliger lilianae.

New records: Key Largo; Long Key; Big Pine

Key; Geiger Key.

Previous records: Brazil; Puerto Rico (Marcus

& Marcus, 1969); Jamaica (Thompson, 1977).

Habitat: Always associated with Avrainvillea

spp., mostly in sheltered, shallow water; occa-

sionally to 5 m depth (largest animals were

found at this depth).

Food: Avrainvillea spp. (Clark & Busacca,

1978; Jensen, 1981a).

Development: Type 3 (Clark & Goetzfried,

1978; Clark & Jensen, 1981).

Taxonomic remarks: Possibly this species is a

synonym of Doto ocellifera Simroth from Ber-

muda (Clark, in prep.). Thompson (1977) cites

Marcus (1976) as authority for Florida distri-

bution, but this citation did not mention C.

lilianae.

24. Hermaea cruciata (lould, 1870

New records: Key Largo; Haulover Canal (In-

dian River).

Previous records: Chesapeake Bay (Vogel,

1971); Massachusetts (Gould, 1870).

Habitat: In dense growth of red algae in

Vol. 97(1)

January 31, 1983

THE NAUTILUS

sheltered areas, usually in reduced salinity, in

winter (a few specimens in spring and autumn).

Food: GriJJithsia sp. The rasping movements

of the buccal mass were very rapid. No buccal

regurgitation was observed, and often a central

string of red cytoplasm was left in the algal cell

after the animal had moved on to another cell.

Development: Type 1 (Clark & Jensen, 1981).

Taxonomic remarks: This species is prob-

ably synonymous with H. coirala Marcus, 1955,

in which case the range extends to Brazil.

26. Placida kingstoni H\\om\)'S(m, 1977

New records: Tampa Bay; Fort Pierce Inlet;

Indian River; Key Largo.

Previous records: Jamaica (Thompson, 1977).

Habitat: Strongly seasonal occurrence; spring

and fall on Bryopsis plumosa. medium exposure;

sometimes extremely common on Cladophora

sp. (gracilis?) on oyster shells in the Indian

River near Sebastian.

Food: i?7'(/opsrsp/MTOOsa (Jensen, 1981a), occa-

sionally Cladophora sp.

Development- Type 1 (Clark & Jensen, 1981).

Taxonomic remarks: This species is morpho-

logically similar to Plax'ida viridis Trinchese,

but P. viridis has paired longitudinal stripes on

the sole of the foot.

Description: Pigmentation is somewhat dif-

ferent than Thompson (1977) noted. The chro-

matophores are typically purplish-grey, and are

concentrated on the pericardial hump, ceratal

tips, and anterior sides; the anus, penis, oviduct,

and vaginal ridge are free of pigment. The geni-

talia are pseudodiaulic. Cerata are in two dense-

ly packed rows; a 9 mm specimen had about 75

cerata. The anus is mid-dorsal, lying slightly

posterior to the beginning of the ceratal rows.

Digestive diverticula are sparingly branched

within the anterior body, and extend into the

lower half of the rhinophores. The cerata are

very long (half of body length), and contain

single unbranched diverticula; each diverticu-

lum is surrounded by a delicate, sparsely anasto-

mosed albumen gland (Fig. 2).

27. Ercolania coerulea Trinchese, 1893

Synonyms: E. costai Pruvot-Fol, 1951

(Schmekel, 1968), Stiliger cricetus Marcus &

Marcus, 1970.

New records: Key Largo; Long Key.

Previous records: Key Largo (Marcus & Mar-

cus, 1960); Curasao (Marcus & Marcus, 1970, as

S. cricetus), Mediterranean (Schmekel, 1968);

Lesser Antilles (Marcus & Marcus, 1963).

Habitat: Often found inside clumps oi Dictyo-

sphaeria cavernosa and Valonia sp., which have

to be broken up to reveal the presence of the

animals. They occasionally occur intertidally in-

side these algae in cooler months. Also, some-

times on Cladophoropsis sp. In relatively

sheltered, very shallow water.

Food: Valonia sp., Dictyosphaerla cavernosa,

and Cladophoropsis sp. (Jensen, 1981a).

Development: Type 1 (Clark & Jensen, 1981).

Taxonomic remarks: Synonymy with E. costai

was discussed by Schmekel (1968). Shape of

radular teeth, number of radular teeth and

general appearance indicates that it is also

synonymous with Stiliger cricetus. The speci-

men described by Marcus & Marcus (1970) was

obviously a juvenile.

28. Ercolania funerea (Costa, 1867)

New records: Haulover Canal; Indian River at

Melbourne; Sebastian Inlet; Key Largo; Geiger

Key.

Previous records: Puerto Rico; Curasao (Mar-

cus & Marcus, 1970); Mediterranean (Schmekel,

1968); Florida (Marcus, 1972a).

Habitat: On dense growths of floating or

sessile algae, mostly Chaetomorpha spp., and on

Cladophoropsis sp. growing in very shallow

water; occasionally on Cladophora spp. The

largest specimens occur on Cladophoropsis and

Chaefom.orpha. From 1972-1975 this species was

extremely common throughout the northern In-

dian River on Chaetomorpha. but this alga has

nearly disappeared from the area in recent

years.

Food: Chaetomorpha spp., Cladophora spp.,

and Cladophoropsis sp. (Jensen, 1981a).

Development: Type 1 (Clark & Jensen, 1981).

29. Ercolania fuscata {Gould, 1870)

Synonyms: Stiliger fuscat us, ?S. evelinae Mar-

cus," 1959 (see Baba & Hamatani, 1970), S.

vanellus Marcus, 1957, ?S. talis Marcus, 1956.

10 THE NAUTILUS

January 31, 1983

Vol. 97(1)

New records: Sebastian Inlet; Key Largo;

Windley Key; North Indian River.

Previous records: Noank, Conn. (Clark, 1975);

Nova Scotia (Bleakney & Meyer, 1979); Baja

California, Mexico (Ferreira & Bertsch, 1975);

Chesapeake Bay (Marcus, 1972b, also as E.

vanellus); Miami (Marcus & Marcus, 1960 as E.

vanelliis); Virginia Key (Marcus & Marcus, 1970,

as E. vanellus); Jamaica (Edmunds, 1966);

Brazil (Marcus, 1957, also &&.E. talis); ?Chile

(Marcus, 1959, as S. evelinae). Not in Australia

as stated by Thompson (1973) (see Gascoigne,

1978), because Thompson's material has pro-

podial tentacles and a broad foot; possibly this

animal is an Alderiopsis.

Habitat: Most often on Cladophora spp.,

either growing attached to mangrove roots or

free floating; occasionally on Chaetomorpha sp.

(see also Clark, 1975).

Food: Cladophora. spp. and Chaetomorpha,

spp. (Clark, 1975).

Development: Type 1 (Clark, 1975; Clark &

Jensen, 1981).

Taxonomic comments: S. vanellus differs

from E. fuaaita only in pigmentation. Popu-

lations of E. fuscata on Cladophora frequently

contain specimens with "vanellus" pigmentation

as well as normal, melanistic/wsra/a specimens;

Chaetomorpha. populations are almost entirely

melanistic. Thus, we regard "vanellus" as an

ecotypic variant oi fuscata.

30. Stiliger fusc&vittatus Lance, 1962

New records: Indian River at Titusville.

Previous records: California (Lance, 1962).

Habitat: Single occurrence, on algae attached

to marker float.

Food: Polysiphonia spp. (Lance, 1962; own

obs.).

Development: Type 1 (small eggs).

DISCUSSION

We estimate that there are at least 35 asco-

glossan species in Florida; this includes the 25

species for which we have presented new distri-

butional data, 6 additional described species, 3

undescribed Elysia species, 2 undescribed Stili-

ger species, and a new I'lacida. This fauna in-

cludes nearly all species reported for the greater

Caribbean fauna, excepting Cyerce cristallina

(known from Bermuda; Thompson, 1977), Costa-

siella nonatoi Marcus (Brazil, Puerto Rico; Mar-

cus, 1977), Volvatella bermudae. Clark, (Ber-

muda; Clark, 1982), and Polybranckia viridis

(Deshayes) (Marcus, 1977), when synonomies

are accounted for. This high diversity - about

15% of all described species (worldwide) - exists

because the Florida fauna includes both truly

tropical components (e.g. Mourgona germainae,

Berthelinia caribbea) and temperate species

(e.g. Her'm.aea cruciata. Ercolania fuscata). We

attribute part of our success in collecting

Florida species to the "island mass effect" -

higher nutrient availability in areas near large

land masses due to runoff- which seems to

favor high recruitment of ascoglossans. Com-

parable collecting activities in drier, smaller

island areas of the Caribbean (e.g. the Bahamas

and Virgin Islands) often yield much smaller

numbers of individuals and species. This effect

is also visible within Florida, as the Lower Keys

provides relatively poor collecting. There are as

yet no published records for ascoglossans from

the northwest coast of Florida (Gulf of Mexico),

but the relatively productive waters of this area

might provide interesting collections.

Of all Ascoglossa reported from Florida, only

the habitat of Stiliger vossi remains unknown;

this species has not been observed since its

original description.

While many of the new records reported here

increase known ranges by only about 360-400

km, this involves transition between essentially

tropical and subtropical marine habitats, so we

regard these new records as significant. Sand-

bottom siphonalean algal communities, in which

most tropical Ascoglossa occur, are rare or ab-

sent in nearshore waters north of Miami. The

area between Fort Pierce and Sebastian Inlet

marks the northern limit for growth of Hali-

rneda, and several Cauler-pa species do not occur

north of Sebastian Inlet (however, C. ashmeadii

and C. prolifera have recently invaded the

northern Indian River between Sebastian Inlet

and Titusville). Although we have not yet col-

lected species which feed on these algae in the

Indian River, we expect that these spec'" (e.g.

Vol. 97(1)

January 31, 1983

THE NAUTILUS 11

Elysia cauze) may extend their ranges into this

coastal lagoon.

Nearly every siphonaceous alga we have ex-

amined in Florida supports at least one asco-

glossan species, though not in all habitats where

such algae occur. Even at the range limits of

such algae, intensive collecting usually yields

animals. Because the Ascoglossa are highly

stenophagous, we feel that the ranges of species

that occur in Florida are limited primarily by the

occurrence of appropriate foods, rather than

directly by temperature. In central Florida,

many such algae appear to be limited by the oc-

currence of suitable habitats (e.g. quiet, shallow

water, sandy bottom, high salinity), rather than

temperature, though Cape Canaveral seems to

represent a significant thermal barrier farther

north (Parr, 1933).

Several species (Elysia ornata, Cyerce ant.il-

leyisis, Caliphylla mediterranen, Bosellia

mimetica, Aplysiopsis zebra) occur in very high

energy habitats at Fort Pierce or Sebastian In-

let. These species do not seem to possess special

adaptations to strong wave action (except B.

miynetica), but rather seem to exploit micro-

habitat conditions (eddies, dense algal growth,

lees of boulders, submerged tidepools) to avoid

wave energy. In the case of Aplysiopsis zebra,

animals appear to crawl below the sand surface

at low tide. Collectors should be aware that ex-

posed coasts do in fact provide suitable habitats

for Ascoglossa.

There are noteworthy differences in habitat

specificity and population density between

temperate and tropical ascoglossans. Northern

species may occur seasonally in extremely dense

populations (Clark, 1975), but tropical species

usually occur in very low densities; we frequent-

ly spend hours to collect a hundred specimens in

the Florida Keys, but a single handful of algae in

New England may contain several hundred ani-

mals during seasonal peaks. Also, temperate

species occur almost ubiquitously, where suit-

able foods are found, but tropical populations

are frequently strongly clumped, and a distance

of as little as 100 m may mean the difference

between success and failure in finding a given

species, despite uniform occurrence of food

algae and apparently similar features of micro-

habitat. We are unable to explain this variability

at present. However, selective predation may be

a partial cause, because aquarium culture of

food algae shows that juveniles are far more

abundant than adults under field conditions.

This clumped nature of tropical populations,

combined with patchy occurrence of algal foods,

makes tropical populations extremely difficult

to quantify. However, our qualitative assess-

ment of most species is that they are ty|3ically

present all year, with egg production extending

over a period of several months; probably most

species have the potential to produce several

generations per year, though these tend to be

clustered during optimal periods of favorable

temperature. Most true tropical species seem to

reach peak activity in late summer and autumn,

when water temperatures reach as high as

32°C.

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Vol. 97(1)

January 31, 1983

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Trap. Oceanogr. Miami 6:3-137.

1969. Euthyneure Meeresschnecken Brasiliens

(2). Beitr. Neotropisrh. Fauna 6:1-1.5.

1970. Opisthcibranclis from Curavao and fauni-

stically related regions. Stud. Fauna Curafiiu 33:1-129.

Marcus, E. and H. Hughes. 1974. Opisthobranch molluscs

from Barbados. Bull. Mar. Sri. 24:498-532.

Meeder, J. F. and D. R. Moore. 1972. The extension of

range of Berthelinia caribhea Edmunds to Brazil and Pan-

ama(Mollusca, Gastropoda). Carih. Jour. Sci. 11:159-161.

Moore, D. R. and M. F. Miller. 1979. Discovery of living

bivalved gastropods in the Florida Keys. The Nautilus

93:106.

Parr, A. E. 1933. A geographic-ecological analysis of the

seasonal changes in temperature conditions in shallow

water along the Atlantic coast of the United States. Bull.

Bingham Oreanogr. Lab. 4:1-90.

Pease, W. H. 1860. Descriptions of new species of Mollusca

from the Sandwich Islands. Pror. Zool. Sur. London

28:18-37.

Portmann, A. 19;'8. BonelUa mnnetira Trinchese, opistho-

branche retrouve en Mediterranee. Vie Milieu 9:74-80.

Reid, J. D. 1964. The reproduction of the sacoglossan

opisthobranch Elysia maoria. Pror. Zool. Sor. London

143:365-393.

Risbec, J. 1956. Nudibranches du Viet-Nam. Arrh. Mux.

Nation. Hist. Nat. Paris 7:1-34.

Russell, H. D. 1964. New England nudibranch notes. The

Nautilus 78:37-42.

Schmekel, L. 1968. Ascoglossa, Notaspidea und Nudibran-

chia im Littoral des Golfes von Neapel. Rev. Suisse Zool.

75:103-1.55.

Sphon, G. G. 1971. New opisthobranch records for the east-

ern Pacific. Velige-r 13:368-369.

Sphon, G. G. and David K. Mulliner. 1973. A preliminary

list of known opisthobranchs from the Galapagos Islands

collected by the Ameripagos F^,\pediti(}n. Veliger

15:147-1.52.

Stirts, H. M. and K. B. Clark. 1980. Effects of temperature

on products of symbiotic chloroplasts in Elysiu turn Mar-

cus ((Jpisthobranchia: Ascoglossa). J. Exp. Mar. Boil.

Erol. 43:39-47.

Swennen, C. 1961. On a collection of Opisthobranchia from

Turkey. Z„ol. Meded. 38:41-75.

Thompson, T. E. 1973. Sacoglossan gastrojjod molluscs from

eastern Australia. Pror. Malar. Sor. Lond. 40:239-251.

1977. Jamaican opisthobranch molluscs I. J.

Moll. Stud. 43:93-140.

Verrill, A. E. 1900. The nudibranchs and naked tectibranchs

of the Bermudas. Trans. Conn. Arad. Sri. 10:545-550.

1901. Additions to the fauna of the Bermudas

from the Yale expedition of 1901 with notes on other

species. Trans. Conn. Arad. Sri. 11:15-62.

Vogel, R. M. 1971. The biology and a redescription of the

opisthobranch mollusk Hennaea rruriata Gould, from

Chesapeake Bay (Maryland). Veliger 14:155-157.

Warmke, G. L. 1966. Two species of the bivalved gastropod

Berthelinia found in Puerto Rico. The Nautilus

79:139-141.

Warmke, G. and L. Almodovar. 1972. Observations on the

life cycle and regeneration in ' xynoe antillarum Mbrch,

an ascoglossan opisthobranch from the Caribbean. Bull.

Mar. Sri. 22:67-74.

Weaver, S. and K. B. Clark. 1981. Light intensity and color

preferences of five ascoglossan ( = sacoglossan) molluscs

(Gastropoda: Opisthobranchia): a comparison of chloro-

plast-symbiotie and aposymbiotic species. Mar. Behav.

Physiol. 7:292-306.

West, Hilary H. 1977. Chloroplast symbiosis and develop-

ment in Elysia rhlorotiea. Am. Zool. 17:968 (abstract).

A BIOGRAPHICAL SKETCH OF WILLIAM IRVIN UTTERBACK (1872-1949)

Ralph W. Taylor

Department of Biological Sciences

Marshall University

Huntington, West Virginia 25701

William Irvin Utterback was born II October

1872 on a farm near Crawfordsville, Mont-

gomery Co., Indiana. His father, John Walton

Utterback, represented the seventh generation

of Utterbacks in the United States. The Utter-

back family tree, in the new world, has its roots

in one Hermann Otterbach, a native of Trup-

bach near Siegen, Germany, who with forty-one

other German immigrants set foot on the

Virginia coast in April, 1741. This small group

14 THE NAUTILUS

January 31, 1983

Vol. 97(1)

FIG. 1. William I. Utterback, about 1925.

of coal miners and iron workers established the

settlement known as Germanna, Virginia. They

were given land and conferred tax-free status in

return for settling in the outlying areas around

Williamsburg and acting as a barrier between

marauding Indians and the civilized townfolk of

British descent. This hardy German stock soon

became well known for qualities such as in-

dustriousness, literacy and religious endeavors.

Early records mention that nearly all members

of the colony could read and write.

Very quickly the new arrivals became angli-

cized in ways, speech and names. Hermann

Otterbach became Herman Utterback. The

name has remained unchanged and the line con-

tinued unbroken to John Walton Utterback

(1850-1897). John Walton married Martha

Hannah Miller (1854-1936) on December 14,

1871, and ten months later she bore a son

christened William Irvin Utterback.

Young Utterback exhibited a propensity for

learning, and at the age of eighteen years ac-

quired teaching certification. In the fall of 1890

he began, in a rural Indiana elementary school,

a teaching career which would span fifty-five

years. In the year 1905, having earned a B.S.

degree from Wabash College in 1901, Utterback

moved to St. Joseph, Missouri, and took a posi-

tion as high school biology teacher, a post he

would hold intermittently until 1917. At the

direction of the Bishop of the Presbyterian

Church of Missouri, Utterback traveled in 1907

to Arkansas. He established, and became the

first president of. The School of the Ozarks

(presently The College of the Ozarks), a Presby-

terian denominational school. The years 1908-

1911 were spent as an instructor of biology at

Westminster College; the following six years

(1911-1917) were important in the development

of Utterback's scientific career. During this

period he returned to his position as biology

teacher at St. Joseph High School, but more im-

portantly he enrolled in graduate school at the

University of Missouri. Utterback was granted

the A.M. degree in 1915, having done a good

portion of his graduate work under the direction

of Dr. George Lefevre. During the summers of

1913-1914 he had the opportunity of working at

the U. S. Fisheries Lab, Fairport, Iowa, where

he learned a great deal about freshwater mussel

biology and also did much of the work that led to

the publication for which he is today most

remembered, "The Naiades of Missouri,"

published in 1916. These middle years of his life

were the most productive scientifically and a

steady flow of papers, all having to do with the

biology of naiades, resulted.

Other significant events in his life occurred

during this period. He married Bessie Ruby

Austin (1877-1938) in 1905, and from this union

came a son William I., Jr. (1907) and a daughter

Anna Martha (1908).

In the fall of 1917 the Utterback family moved

to Spokane, Washington, where William was

again employed as a high school biology teacher

at Lewis and Clark High School. Mrs. Utterback

was apparently adversely affected by the

climate of the Spokane area and Utterback was

forced to again relocate.

On October 24, 1919, Utterback accepted a

Vol. 97(1)

January 31, 1983

THE NAUTILUS 15

position at Marshall Normal School (presently

Marshall University) in Huntington, West

Virginia, for which he was paid the princely sum

of $2,000 for a ten-month academic year. Pro-

fessor Utterback from this time forth adopted

the Huntington area and remained at Marshall

University until his retirement at the age of 73

in 1945.

Professor Utterback was, for many years, the

only biologist on staff at Marshall. Old news-

paper clippings indicate that he was an interest-

ing, efficient and dedicated staff member. His

teaching load included Comparative Anatomy,

Embryology, Zoology, In.ertebrate Zoology

and other courses as student needs dictated. He

also served as advisor to pre-medical students.

His advancing age and heavy course load

seems to have reduced his energies given to out-

side research, ^''"v scientific publications were

produced du ing the Marshall years. Professor

Utterback was a devoutly religious man and was

an elder in the Huntington First Presbyterian

Church. His religious beliefs often showed even

in his scientific works.

"Yet in the last analysis we scientists, who deal

so much with the material and natural law,

may lose sight of the spiritual and supernatural

in the fact that the GREAT JEHOVAH (the

SELF-EXISTENT ONE) is the Alpha and

Omega whether we consider Evolution in the

life history of the individual or even of all

Phyla."

Phylogeny and Ontogeny of Naiaden. 1928.

While it is true that the production of scien-

tific papers waned in the later years, he did not

stop writing. In two books and several lesser

publications he sought to document his ances-

tory (The Utterback Family 1622-1937), and to

bring together his religious beliefs and scientific

thinking. Through the book "The Second Tri-

angle" and a lesser publication "The Great Life

Cycle" Professor Utterback brings together the

concepts of Evolution and Special Creation. His

thesis states that Seth, the third born of Adam

and Eve, carried the God-created line and

blended it, through marriage, with the line of

Anthropoid human beings which had evolved

through time outside the Garden of Eden.

I have in my possession a mimeographed

booklet titled Single Verse Poems produced by

Utterback about 1930. The following four line

verse attests to the conflict which garnered

much of his thinking in later life.

Special Creation and Evolution

Special Creation is mutation

By God's almighty hand

Evolution is Revolution

Through laws at HIS rommand.

Utterback the man

Professor Utterback was deeply involved in

many other aspects of community life. He held

memberships in the A.A.A.S., the West Virginia

Academy of Science, Chi Beta Phi and the

American Institute of Geneology. In addition he

was an active member of the local chapter of the

Sons of the American Revolution, working

through the ranks to become chapter president

in 1942.

Utterback epitomized the early American

naturalist and corresponded regularly with such

distinguished biologists as Drs. F. C. Baker, R.

E. Coker, A. E. Ortmann, V. Sterki and W. H.

Dall, as well as Thaddeus Surber, Bryant

Walker and L. S. Frierson. In the summer of

1913-14 he surveyed, in a row boat, three hun-

FIG. 2. William 1. Utterback, about i;t45.

16 THE NAUTILUS

January 31, 1983

Vol. 97(1)

dred miles of the Osage River of central

Missouri; a feat which seems overwhelming in

light of present day thinking and technology.

The enclosed picture of Utterback in full field

regalia says a great deal about Utterback the

naturalist.

Upon retirement Professor Utterback moved

to Los Angeles, California, where he lived with

his daughter until his death on May 16, 1949.

The professor was buried May 20, 1949 in

Woodmere Cemetery, Huntington, West

Virginia.

Publications of William Irvin Utterback

The Myth of the Manitou. The Wabash (local news-

paper). Crawsfordsville, Indiana. (No date avail-

able.)

1914 Mussel Resources of Missouri. U. S. Bureau of

Fisheries. Econ. Cir. #10. pp. 1-6.

1916 The Naiades of Missouri. Univ. of Notre Dame

Press, Notre Dame, Indiana, pp. 1-200.

1916 Breeding Records of Missouri Mussels. Nautilus

30:13-21.

1916 Parasitism among Missouri Naiades. A^ner. Mid,

Nal. 4:518-.521.

Ca. 1920 The Second Triangle. Gorman Press, Boston.

80 pp.

1927 The Great Life Cycle. Gentry Bros. Printing Co.,

Huntington, WV. One oversized printed page.

1928 Phylogeny and Ontogeny of Naiades. Proc. W. Va.

Acad. Sn. 2:60-67.

1930 A new genus of freshwater mussels (Naiades).

Proc. W. Va. Acad. Sci. 4:66-69.

1931 Sex behavior among Naiades. Proc. W. Va. Acad.

Sci. 5:43-45.

1933 New glochidia. Proc. W. Va. Acad. Sci. 6:32-36.

1937 The Utterback Family 1622-1937. Gentry Bros.

Printing Co., Huntington, W\'. 470 p.

Selected Additional Readings

Johnson, Richard I. 1969. The Unionacea of William I.

Utterback. The Nautilus 82:132-13.5.

Fuller, S. L. H. 1974. Neglected papers on naiades by W. I.

Utterback. The Nautilus 88(3):90.

THE SOUTH AMERICAN HYDROBIOID GENUS IDIOPYRGUS

PILSBRY, 1911

Emile A. Malek

Department of Tropical Medicine

Tulane University

Medical Center

New Orleans, Louisiana 70112

ABSTRACT

The shell and soft parts of the hydrobioid Idiopyrgus and I. souleyetianus

Pilsbry, 1911, are described from material from two sites in the state of Minos

Gerais, adjacent to the state ofEspirito Santo. Brazil, the type locality. Examina-

tion of other specimen's from the states ofBahia and Mato Grosso indicated a wide

geographic distribution in Brazil of species belonging to this gemis. Hydracme

Haas, 1938, and Aquldauama Dwvis, 1979, are considered synonyms, while Onco-

melania brasiliensis Rey, 1959, is invalidated.

There is a great need to stabilize the systema-

tics of some South American hydrobioid fresh-

water snails, which are actual or potential first

intermediate host of the lung flukes, Paragoni-

mus spp. Of particular interest are those hydro-

bioids of Brazil. Pilsbry (1911), created the

genus Idiopyrgus from the state of Espirito

Santo, eastcentral Brazil, with /. souleyetianus

Pilsbry as type species. The description of the

genus and species was based on shell and radu-

Vol. 97(1)

January 31, 1983

THE NAUTILUS 17

lar characteristics only. Other species of

Idiopyrgus were later described: namely /.

pilsbryi by F. Baker (1913), and /. walkeri by

Pilsbry (1924). Haas (1938) erected a new genus,

Hydracme, with H. rudolphi Haas as type, from

the state of Pernambuco in the northeast. Rey

(1959) described Oncomelnnia hrasiliensis from

the Paraguay river drainage in Campo Grande,

the state of Mato Grosso. The general anatomy

of the reproductive systems as well as the radula

were presented. On the basis of a preliminary

examination of snails from the same state in

Brazil, but from the Aquidauana river drainage

Davis (1979) believed that Rey's hydrobioid

snails should belong to a new pomatiopsine

genus Aquidauania Davis, rather than to the

Oriental genus Oncomelania. Davis's account

was very brief, and he reserved further informa-

tion on his genus until more material becomes

available.

Specimens collected by the writer in the states

of Minas Gerais and Bahia, and specimens given

to him from Mato Grosso, made it possible to

revise the systematics of this hydrobioid group

from Brazil. Emphasis is made in this report on

a complete description of Pilsbry's Idiopyrgus

and /. souleyetianus and on synonymizing cer-

tain other genera in the same geographic area.

Genus Idiopyrgus Pilsbry, 1911

Shell dextral, turreted; spire long with convex

whorls and deep sutures. Aperture obliquely

ovate, its basal margin being rounded and ad-

vanced; peristome simple or slightly thickened

within. Operculum horny, paucispiral.

Tentacles slender; foot with pedal crease.

Central tooth of radula with few cusps (5 to 7) on

the upper reflection, and also few cusps, 9 and

16 on marginals; outer marginal scoop-like.

Female oviparous; bursa copulatrix large, with

spermathecal duct arising from its middle por-

tion and extending separately from pallial ovi-

duct with which it usually unites near female

genital opening. Sperm duct arises from sper-

mathecal duct close to bursa copulatrix and joins

visceral oviduct. Verge large, curved and sim-

ple, that is, devoid of any appendages or

papillae. Snails are aquatic in freshwater.

Idiopyrgus souleyetianus Pilsbry. 1911

Specimens examined for the description of/.

snuleyetiantcs are from the Brazilian states of

Minas Gerais, Mato Grosso and Bahia: Sete

Lagoas and Pedro Leopoldo, Minas Gerais;

Ceroula stream in Campo Grande, Mato Grosso;

and the following localities in western Bahia:

reservoir behind dam at Sao Desideiro; side road

ditch in outskirts of Barreiras; pond about 10

miles south of Barreiras; and irrigation canal

near Sao Desideiro.

Redescription of shell - Shell (Fig. 1 B) tur-

reted with 6 to 7V2 rounded, convex and smooth

whorls; sutures deep. Adult shells measuring 4

to 6.5 mm (average 5.5) in height, and averaging

2.1 mm in width. In old specimens spire becomes

eroded and thus shortened. Shell with a small

umbilical chink behind aperture at columellar

margin. Aperture obliquely ovate, basal margin

rounded with slight expansion; outer lip slightly

thickened within, sometimes reflected, but

always retracted upward forming a rounded

sinus just below upper insertion. Operculum cor-

neous, transparent, paucispiral, with the

nucleus near columellar margin.

Deseription of aoft parts - Animal grayish

black; tentacles slender, with their distal two-

thirds black. Eyes each with an "eye brow" of

white-yellowish color. Snout slightly bilobed

anteriorly. Foot elongate, truncate anteriorly

and tapering posteriorly beyond attached oper-

culum. Foot with pedal crease (Fig. 1 D), and

there is a suprapedal fold continuous with oper-

culigerous lobe. Omniphoric groove present be-

tween suprapedal fold and snout. Gill with about

50 lamellae, high near origin on left side close to

mantle collar and tapers posteriorly. Central

tooth of radula with 2 or 3 cusps on each side of

a larger cusp on the upper reflection, and with 2

or 3 basal cusps. Lateral tooth with 7 cuSps and

inner marginal tooth with 9 cusps. Outer mar-

ginal scoop-like, with about 16 cusps.

Female oviparous. Ovary yellowish, having a

few branches embedded in the apical portion of

the digestive gland. Oviduct tubular at its

viscera! portion and convoluted in the region of

kidney, then enlarges to form a saccular and

wide posterior portion of pallial oviduct

(Fig. 1 A). The latter extends, almost of the

18 THE NAUTILUS

January 31, 1983

Vol. 97(1)

OG SF

KIG. 1. Idiopyrgus souleyetianus Ptlsbry, 1911. A, Female reproductiir oryatis. B, Female snail, apertural new. C, Male

reproductive organs. D, Male, snail shovring verge, snout, foot, operculum and gill. E, Radula. APO, anterior pallial oviduct;

BC, bursa copulalrix: F, eye; F, foot; FGO, female genital opening: G, gill; MC, mantle collar; 0, operculum; OG, omniphoric

groove; OV, ovary; P, prostate: PC, pedal crease; PPO, posterior pallial oviduct; S. snout; SD, sperm duct; SF, supra pedal fold;

SPD, spervmthecal duct: SR, seminal recefdavle; T, tentacle; TF. Ie>itis; V. verge: VT). ru-s- deferens: VOL), visceral oviduct.

Vol. 97(1)

January 31, 1983

THE NAUTILUS 19

same diameter, as the anterior portion of pallia!

oviduct to female genital opening on right side

of anal opening at mantle collar. Bursa copula-

trix large, oval near posterior pallial oviduct.

Spermathecal duct arises from middle of bursa

and runs anteriorly along pallial oviduct and

joins the latter immediately before female

genital opening. In a few specimens, however,

spermathecal duct opens separately to outside.

Sperm duct arises from spermathecal duct close

to bursa and opens in the tubular visceral

oviduct, close to opening of short stalk of

seminal receptacle.

In the male, testis embedded in apical portion

of digestive gland. Testis larger than ovary of

female and consists of digitiform diverticula

which end in small ducts that unite to form vas

deferens. The latter, convoluted at its begin-

ning, eventually reaches a kidney-shaped pro-

state gland. After receiving secretions from pro-

state gland vas deferens proceeds anteriad,

along esophagus until it reaches base of verge,

and proceeds inside it running along convex

margin to tip. Base of verge fleshy, almost flat-

tened and curves to form a cylindrical terminal

portion which tapers to tip where it forms a

minute knob-like structure. Verge simple

throughout, devoid of any appendages or

papillae (Fig. 1 C and D).

DISCUSSION

No important morphological differences were

found among the hydrobioid material examined

from the Brazilian states of Minas Gerais, Bahia

and Mato Grosso. It is to be noted that Minas

Gerais is adjacent to the state of Espirito Santo

where the Rio doce is the type locality of Idio-

pyrgus souleyetianus. The Mato Grosso material

was examined previously by Rey (1959), who

designated Oricomelania brasiliensis for the

specimens from the Paraguay river drainage in

Campo Grande county, and by Davis (1979), who

designated Aquidauania bi'asiliensis for the

specimens from Aquidauana river drainage. The

writer agrees with Davis that the specimens

cannot be considered an Oncovielania because

of their aquatic rather than amphibious habitat,

and that the bursa copulatrix complex differs

from that of Oncomelania. Davis (1979),

however, did not study in detail the anatomy of

the Campo Grande material but relied on Key's

(1959) description. Despite this he erected a new

genus Aquidauania. Based on the morphologi-

cal information given in the present paper I con-

sider Pilsbry's genus and species, Idiopyrgus

souleyetianws, which were described on the basis

of the shell and radula only, to extend west,

from Espirito Santo, Minas Gerais and Bahia,

into Mato Grosso. I, therefore, invalidate On-

comelania brasilien.sis Rey, 1959, and place

Aquidauania Davis, 1979 into synonymy with

Idiopyj-gufi.

Another hydrobioid genus which was de-

scribed from the same geographic area is

Hydracme, with H. rudolphi as type, from Sao

Francisco river near Jatoba, state of Pernam-

buco (Haas, 1938). This new name was based on

a series of specimens taken from a fish stomach.

Taylor (1966) correctly synonymized Hydracme

with Idiopyrgus because of the great variation

he encountered when examining a large series

of topotypes. The variation within this series,

and among other lots representing undescribed

species from the states of Paraiba and Pernam-

buco, effaces the supposed distinctions between

Hydracme and Idiopyrgus. However, Taylor,

apparently using shell features alone, placed the

genus Idiopyrgus under his subfamily Lit-

toridininae (family Hydrobiidae), together with

Littoridina, DurangonMa, Pyrgophortis. Mex-

ipyrgus and others. It has been pointed out in

the description of Idiopyrgus in the present

report that the verge is simple, without papillae,

lobes or appendages, thus differing completely

from the latter genera in the Littoridininae.

Two other species of Idiopyrgus have been

described on the basis of the shell alone, namely

/. pilsbryi, from north bank of chief affluent of

Papary lake near its mouth, state of Rio Grande

do Norte by F. Baker (1913), and /. walkeri,

from Rio de Valhas, tributary to Sao Francisco

river at Lassance, state of Minas Gerais by

Pilsbry (1924). Probably these two species

should remain valid until their anatomy is

elucidated.

Certain morphological features of Idiopyrgus,

as presented in this paper, place it in the family

Pomatiopsidae, subfamily Pomatiopsinae,

20 THE NAUTILUS

January 31, 1983

Vol. 97(1)

rather than in the Hydrobiidae, as defined by

Davis (1979). The pomatiopsine hydrobioids

have a spermathecal duct which is separated

from the paiHal oviduct, and is derived not from

the ciliated ventral channel as found in the

Hydrobiidae, but from elongation of a bud from

the bursa copulatrix; males have a simple verge,

that is, with one duct and is devoid of appen-

dages or lamellae; there is a pedal crease; the

central tooth of the radula lacks the pronounced

lateral angles of Hydrobia, and there are two or

more pairs of basal cusps, the largest arises

from the face of the tooth, a condition not seen

in Hydrobia. other pairs arise from the face or

from the lateral angle.

ACKNOWLEDGMENTS

The author is indebted to Dr. W. Lobato

Paraense for supplying specimens from Campo

Grande, Mato Grosso, and from Pedro Leo-

poldo, Minas Gerais. This study was supported

by a grant from the World Health Organization.

LITERATURE CITED

Baker, Fred. 1913. The land and fresh-water mollusks of

the Stanford Expedition to Brazil. Proc. Acad. Nat. Sci.,

Philadelphia 65:618-672.

Davis, G. M. 1979. The origin and evolution of the gastropod

family Pomatiopsidae with emphasis on the Mekong River

Triculinae. Acad. Nat. Sci., Philadelphia Monograph

20, 120 pp.

Haas, F. 1938. Neue Binnen-Mollusken aus Nordost-

Brasilien. Arch. Molluskenk. 70:46-51.

Pilsbry, H. A. 1911. Non-marine Mollusca of Patagonia.

Report Princeton Univ. Expeditions to Patagonia

1896-1899, 3:.513-633.

1924. South American land and fresh-water

mollusks: notes and descriptions. Proc. Acad. Nat. Sci..

Philadelphia 76:49-66.

Rey, L. 1959. Molluscs of the genus Oncomelania, in Brazil,

and their possible epidemiological significance. Rei<. Inst.

Med. Trop. Sao Paulo 1:144-149.

Taylor, D. W. 1966. A remarkable snail fauna from Coahuila,

Mexico. Fe^i^r 9:152-228.

VITRINIZONITES LATISSIMUS (PULMONATA: ZONITIDAE) AND

VERTIGO CLAPPI (PUPILLIDAE) FROM EASTERN KENTUCKY

Leslie Hubricht

4026 35th Street

Meridian, Mississippi 39301

Ronald S. Caldwell

Department of Biology

Te.xas College

Tyler, Texas 75702

John G. Petranka

Division of Biological Sciences

University of Michigan

Ann Arbor, Michigan 48109

The Blue Ridge Snail, Vitrinizonit.es latissi-

mus (Lewis), has been previously reported from

the Blue Ridge Physiographic Province of Ten-

nessee and North Carolina, and from the Ridge

and Valley of Virginia (Hubricht, 1970, 1971,

1973; Pilsbry, 1946). Pilsbry (1946) reported its

occurrence from the Cumberland Plateaus near

Gurley, Alabama, based on four specimens in

the collections of the Academy of Natural

Sciences of Philadelphia. Recent collecting by

the senior author in the vicinity of that locality

has failed to yield V. latissimiis, and its current

status in Alabama is uncertain. According to

Pilsbry (1946) it is generally distributed at eleva-

tions above 2000 feet, being restricted to moist

places with abundant moss cover. Hubricht

(1961) expressed the belief that Vitrinizonites

uvidermis Pilsbry represents gerontic in-

Vol. 97(1)

January 31, 1983

THE NAUTILUS 21

FIGS. 1 and 2. 1, Venti-al (uew o/ Vitrinizonites latissimus

(Lewis) from Pine Mountain. Width: 18.9 mm. 2, Vertigo

clappi Brooks and Htnit from Puie Mountnin. Height:

1.5 mm.

dividuals of V. latissimus which have resorbed

the calcareous layers of the shell. All individuals

examined fi-om Kentucky localities had rigidly

calcified shells.

Vertigo clappi Brooks and Hunt is known

from Marion County, West Virginia, in the

Allegheny Plateaus, and from Jefferson, Green-

brier, Hampshire, and Pendleton counties. West

Virginia, and Loudon County, Virginia in the

Ridge and Valley (Briscoe, 1963; Brooks and

Kutchka, 1938; ' MacMillan, 1949). Little is

known about its habitat preference or other

aspects of its ecology.

We have recently collected specimens of V.

latissimus from Pine and Big Black Mountains,

Harlan County, Kentucky, and Vertigo clappi

from Pine Mountain, Harlan County, Kentucky.

These collections represent significant exten-

sions of the known ranges of the two species and

are the first verifiable reports of either species

in the Cumberland Plateaus.

Pine Mountain is a northwest-facing mono-

clinal mountain extending some 200 km along

the northwest edge of the Cumberland over-

thrust block. Elevation varies from approxi-

mately 650 m in the southwestern half to about

850 m in the northeast. The Pine Mountain site

is located along U. S. Hwy. 421, 0.6 km south-

east of its junction with Ky. Hwy. 221, elevation

about 700 m, lat. 36°53'57"N, long. 83°20'15"W.

The site is a north-facing mountainside of mesic

woods with numerous moss-covered boulders of

Pennington limestone. Specimens of V. latissi-

mus were collected exposed on vegetation when

active in humid weather, and beneath moss mats

and in leaf litter when not active. Vertigo clappi

was found only beneath moss mats and accumu-

lated detritus on top of limestone boulders.

Other land snails collected at this site are listed

in Table 1. Further collecting at four sites of

similar elevation and exposure along Pine

Mountain from U. S. Hwy. 421 to U. S. Hwy.

119 near Whitesburg, Ky. yielded no further

specimens of either species. The distribution of

these two species on Pine Mountain would thus

appear to be very patchy.

Big Black Mountain is a maturely dissected

residual mass of horizontally bedded Pennsyl-

vanian sandstones and shales rising to an eleva-

tion of about 1260 m, some 250 m above the ad-

jacent Cumberland peneplain. Vitrinizonites

latissimus was initially collected at two sites by

members of the Eastern Kentucky University

malacology class (B. Branson, pers. comm.), and

has been subsequently collected by the authors

at two additional sites. Three of the sites are

along Ky. Hwy. 160, and range in elevation

from 1005 m to 1200 m. The fourth site was near

the summit, elevation 1240 m. All sites were of

mesic woodland with varying degrees of ex-

posed sandstones and shales. Vitrinizonites

latissimus was not abundant at any site on Big

Black Mountain, but appears to be rather widely

distributed.

The apparent disjunctions of the Kentucky

populations of these two snail species from the

main part of their ranges, along with their

restriction to higher elevations, suggests that

these populations may represent relicts of a

formerly widespread distribution. Davis and

Barbour (1978) summarized evidence for the

refugial nature of Big Black Mountain with

respect to vascular plants, birds, mammals, and

carabid beetles. Twenty-three species of plants

and animals of northern or Appalachian af-

finities are largely restricted in their Kentucky

distribution to Big Black Mountain. A number of

22 THE NAUTILUS

January 31, 1983

Vol. 97(1)

TABLE 1. Land syiails collected with Vitrinizonites latissi-

mus (Lewis) and Vertigo clappi Brooks and Hunt at Pine

Mountain, Kentucky.

Hendersonia occulta (Say)

Pomatiopsis lapidaria (Say)

Carychium nannodes Clapp

Carychium clappi Hubricht

Cionella morseana Doherty

Columella simplex (Gould)

Gastrocopta pentodon (Say)

Gastrocopta contracta (Say)

Gastrocopta corticaria (Say)

Vertigo gouldi (Binney)

Haplotrema concavum (Say)

Pallifera secreta Cockerel!

Philomycus venuslus Hubricht

Punctum blandianum (Pilsbry)

Discus nigrimontanus (Pilsbry)

Discus patulus (Deshayes)

Anguispira mordax (Shuttleworth)

Succinea ovalis (Say)

Striatum meridionalis (Pilsbry and Ferriss)

Ventridens collisella (Pilsbry)

Gastrodonta interna interna (Say)

Paravitrea capsella (Gould)

Paravitrea subtilis Hubricht

Paravitrea multidentata (Binney)

Mesomphix cupreus (Rafinesque)

Mesomphix perlaevis (Pilsbry)

Mesomphix inomatus (Say)

Glyphyalinia rinmla (Hubricht)

Olyphyalinia cumhrrlandiana (Clapp)

Guppya sterkii (Dall)

Euconulus fulvus (Miiller)

Allogona profunda (Say)

Triodopsis albolabris (Say)

Triodopsis denotata (Ferussac)

Triodopsis tridentata (Say)

Triodopsis vulgata Pilsbry

Mesodon injlectus (Say)

Mesodon appressus (Say)

Mesodtin sayanus (Pilsbry)

Mesodon zaletus (Binney)

Stenotrema stenotrema (Pfeiffer)

Stenotre7na. edvardsi (Bland)

these species of small mammals have been

reported to occur on Pine Mountain as well

(Caldwell, 1980). Many of these populations

presumably represent once more widely distri-

buted Pleistocene relicts.

Branson and Batch (1968) noted that the land

snail faunas of Pine and Big Black Mountains

bear a closer affinity to the faunas of the

Eastern Division of the Cumberland Subregion

of Pilsbry (1900) than to the Western Division.

This view is supported, and our collections pro-

vide further evidence of this region's faunal af-

finities to the Ridge and Valley and Blue Ridge

Physiographic Provinces.

ACKNOWLEDGMENTS

The authors are most grateful to Charles K.

Smith for photographing the Vitrinizonites

latissim.us, and to SEM operator Henry

Southgate for the micrograph of Vertigo clappi.

SEM costs were underwritten by the University

of Kentucky Graduate School.

LITERATURE CITED

Branson, B. A. and D. L. Batch. 1968. Land snails from Pine

and Big Black Mountains, Kentucky. Sterkiana 32:7-17.

Briscoe, M. S. 1963. A survey of land and freshwater snails

in Jefferson County, West Virginia. Sterkiana 9:41-48.

Brooks, S. T. and G. M, Kutchka. 1938. Occurrence of the

family Pupillidae in West Virginia. Ann. Carnegie Mus.

27:63-85.

Caldwell, R. S. 1980. First records of Sorex dispar and Mi-

crosorex thompsoni in Kentucky with distributional notes

on associated species. Tran^. Ky. Acad. Sri. 41(l-2):46-47.

Davis, W. H. and R. W. Barbour. 1978. Kentucky's high

country- a biological treasure. Trans. Ky. Acad. Sd.

39(3-4):138-141.

Hubricht, L. 1961. The status of Vitrinizonites uvidermis

Pilsbry. The Nautilus 74(4): 166.

1970. The land snails of North Carolina.

Sterkiana 39:11-15.

1971. The land snails of Virginia. Sterkiana

42:41-4.3.

1973. The land snails of Tennessee. Sterkiana

49:11-17.

MacMillan, G. K. 1949, The land snails of West Virginia.

Ann. Carnegie Mu.s. 31:89-239.

Pilsbry, H. A. 1900. Mollusca of the Great Smoky Moun-

tains. Proc. Acad. Nat. Sn. Phila. 52:110-150.

1946. Land Mollusca of North America (north of

Mexico). Acad. Nat. Sci. Phila. Monogr. 3. Volume 1,

Part 2.

Vol. 97(1)

January 31, 1983

THE NAUTILUS 23

PEASISTILIFER EDULIS, A NEW EULIMID PROSOBRANCH,

PARASITIC ON AN INDO-PACIFIC HOLOTHURIAN

George P. Hoskin

Division of Microbiology

Food and Drug Administration

Washington, DC 20204

and

Anders Waren

Department of Zoology

University of Goteborg, Box 25059

S-40031, Goteborg, Sweden

ABSTRACT

Peasistilifer edulis new species is described from NE Aicstralia and New Cale-

donia. It closely resembles previotcsly described species of the genus but has a taller

spire and a pr opart ioyiaUy smaller aperture. P. edulis lives as an ectoparasite on

the Indo-Pacific Holothuria (Halodeima) edulis (Lesson) and sucks body fluid from

the lacunae in the body wall of the host.

The family Eulimidae is a large group of meso-

gastropods with highly variable morphology

whose species are ecto- or endoparasites of

echinoderms. A review of the genera is given by

Waren (in press). The new species described

here was found some years ago when the senior

author prepared a study on the biology and

anatomy of Mucronalia nitidula Pease, 1860

(Hoskin and Cheng 1969, 1970). Later, Waren

(1980) described the genus Peasistilifer with M.

nitidula as type species and listed other species

belonging to the genus.

Peasistilifer edulis n. sp.

Description - Shell straight, conical, high and

pointed, rather solid, opaque, very polished and

white. The larval shell consists of about 2.5

whorls (Fig. 3) and its height is 320-340 Mm. The

whorls are slightly convex, perfectly smooth,

more cylindrical than the postlarval whorls, and

differentiated from them by a distinct scar. An

adult female has 10.5 postlarval whorls, the first

of which increases its diameter very rapidly;

later whorls increase slowly and regularly. The

postlarval whorls are unusually convex for the

family. Their largest diameter is about 1/3 the

height of a whorl from the lower suture. Except

for a few indistinct incremental lines, they are

perfectly smooth and polished. There are no

growth scars of the kind characteristic of many

eulimids. The aperture is low and rounded. The

outer lip, seen from the side, protrudes slightly

at the suture, is shallowly sinuated below the

suture, protrudes again at the middle of its

height, and is slightly retracted below this point

(Fig 4.). Some New Caledonia specimens have a

broad and shallow umbilicus behind the low part

of the parietal callus; in others (the type

specimens) the callus forms an expanded pad

covering this area.

Dimensions - Females are 1.5-1.7 times

larger than males but slightly narrower in pro-

portion to their height (Figs. 1 and 2). A large

female (Fig. 2), height 9.69 mm, diameter 4.74

mm; aperture height 2.50 mm, breadth 1.79

mm. Holotype, found to be male, height 5.48

mm, diameter 2.75 mm, height of the aperture

1.05 mm, breadth 0.55 mm.

Soft parts - The tentacles and the area

around the posterior pedal gland are light-

yellow; other soft parts are more or less color-

less. (In P. yiitidula there is one reddish spot in

front of each eye, on the tentacle, a series of red

spots along the rectum, and a reddish line along

the osphradium. Remaining parts of the body

are white.) The tentacles are rather long and

slender with large black eyes at their bases. The

male has a large penis behind and to the right of

the right tentacle. The foot is rather small but

functional and is equipped with anterior and

posterior pedal glands. The pedal gland opens

into a furrow on the underside of the foot and

produces a filament of mucus used for attach-

ment to the host. There is a distinct propodium.

On the sides of the foot are two pedal flaps.

The left one is larger and extends from the

24 THE NAUTILUS

January 31, 1983

Vol. 97(1)

FIGS. 1-3. 1 and 2, Peasistilifer edulis, shell. 1, Holotype,

Heron Island. SE Australia. USNM 787954. Height 5.1,8

mm, male. When, collected the shell was white and highly

polished. Apparent markings due to presentation damage. 2,

Female, Rot Maitre, New Caledonia, 9.69 mm high.

(Photographs not to scale.) 3, Operculum of P. edulis. Height

1.81 mm. Extreme tip broken; dashed line indicates broken

part.

posterior end of the opercular lobe along the

side of the foot, ending between and just to the

right of the proboscis and the left tentacle; its

anterior part is drawn out into a short tongue.

The right flap is smaller and thinner and does

not reach as far posteriorly as the left one. It

also lacks the tongue-shaped anterior end, but

ends in the corresponding position on the right

side. These flaps are highly muscular and prob-

ably cover the base of the shell when the snail is

parasitizing.

The proboscis is long (but proportionately

shorter than in P. nitidula) and may not be com-

pletely retractile; it was partly extended in

preserved P. edulis specimens, whereas in P.

nitidula it was completely retracted. The oper-

culum is large, paucispiral and equipped with a

large bulbous muscular attachment deeply in-

serted into the opercular lobe (Fig. 5). The oper-

culum has radial growth lines and a few spiral

lines, one of which is predominant. A distinctly

thinner zone along the outer edge acts as a pack-

ing to ensure tight fitting of the operculum.

Type locality - On reef flat. Heron Island,

Queensland, Australia; attached to body wall of

Holothuria (Halodeima) edulis (Lesson). Speci-

mens were collected by the senior author,

December 1967.

Holotype - One large male specimen from the

type locality, U.S. National Museum of Natural

History (USNM), Division of Mollusks, reg. no.

786715 (Fig. 1).

Paratypes - Six specimens from the type

locality, USNM, 787954; one large female (Fig.

2) and one male from the body wall of H. edulis,

Ilot Maitre, off Noumea, New Caledonia, April

27, 1978, 1 m deep, on the reef flat, coll. P.

Bouchet, material in Museum National

d'Histoire Naturelle, Paris; a few specimens

from Suva, Fiji, January 1967, coll. G. Hoskin;

specimens destroyed during anatomical study.

Biology - P. edulis lives as an ectoparasite on

FICS. 4-6. 4, Sinuation of outer lip of P. edulis. 5, Apex of P.

edulis. Scale iine 250 \im. 6, Apex of P. nitidula. Scale line 250

Vol. 97(1)

January 31, 1983

THE NAUTILUS 25

its host, H. edulis. It penetrates the tissues of

the holothurian with the proboscis until it

reaches a suitable lacuna from which to suck

body fluid. The dual functions of the proboscis

are attachment and provision of food.

After removal from its host, P. edulis has ob-

vious difficulties reattaching. Four specimens

were removed and isolated for 2 hours. When

replaced on the host, none attached during 2.5

hours of observation. In a similar test, 10 speci-

mens of P. nitidula, isolated for 7 days, reat-

tached after about 30 minutes. In another test,

four specimens of P. edulis and 16 of P. nitidula

were placed in the center of a tank with a speci-

men of H. edulis caged in one end and a

specimen of Holothuria atra caged in the other

end. Ten of 16 P. nitidula reattached to H. atra

after a mean of 32 minutes, whereas only one P.

edulis reattached (after 44 minutes) during the

2.5-hour experiment. All specimens moved to

the vicinity of the normal host species. When

given a choice between H. atra. and H. edulis, P.

nitidula attached itself to H. atra; when given

access only to H. edulis, it parasitized this

species after a few hours' delay.

These tests demonstrated a distinct difference

between P. edulis and P. nitidula in ability to

reattach. It may be that the proboscis of P.

edulis becomes injured when removed from the

host. Such injury has been observed in several

species of Melanella, which also parasitize holo-

thurians.

All specimens of P. edulis were obtained from

a few hundred H. edulis; the junior author ob-

tained 660 specimens of P. nitidula from about

1800 H. atra. which were examined for parasitic

snails in NE Australia, New Caledonia, and the

Loyalty Islands. No snails were found on the

"wrong" host. Apparently there is a high degree

of host specificity in the choice of host in the two

species. The junior author encountered no speci-

mens of P. nitidula or P. edulis during his ex-

amination of about 3000 specimens of other

holothurians. The size and the number of whorls

in the larval shell indicated that P. edulis has

planktotrophic larval development.

Remarks - P. edulis can be recognized among

eulimids parasitic on holothurians by its opercu-

lum which has a peg or bulbous attachment (pre-

sent only in Peasistiiifer), by its distinctly con-

vex whorls with their largest diameter well

below the middle, by the high number of whorls,

and by its tall, conical spire. The larval shell of

P. nitidula, the species of Peasistiiifer that P.

edulis most closely resembles, consists of 3.5

whorls rather than 2.5 (Figs. 3 and 6) and is pro-

portionately broader than that of P. edulis.

Large females of P. edulis resemble species of

Niso in the shape of the shell, but Niso species

usually have flat whorls, a very broad umbilicus,

a keeled shell base, and a more or less distinct

axial sculpture of sharp but indistinct straight or

curved lines. The shells of Niso species are

usually brightly colored and they do not have

opercula with pegs.

In some specimens of P. edulis the umbilicus is

well-developed (Fig. 2); in others it is completely

absent (Fig. 1). However, we believe that this is

an intraspecific variation, not an indication that

two species are involved, because there are also

intermediate specimens. If this assumption is

wrong and there are two species in our material,

the name P. edulis should be used for the form

without umbilicus, because the holotype belongs

to that form.

ACKNOWLEDGMENTS

We thank Dr. P. Bouchet for the sample of P.

edulis from New Caledonia and the staff of

Heron Island Research Station, Queensland,

Australia, for making working facilities avail-

able to us during visits there. The East- West

Center in Hawaii supported the field study of

the senior author and provided financial assis-

tance; the Department of Zoology, University of

Hawaii, provided research facilities during part

of this study.

LITERATURE CITED

Hoskin, G. P. and T. C. Cheng. 1969. A comparison of the

anatomy of two species of Mucronaha (Mollusca; Proso-

branchia): with notes of the distribution of M. nitidula in

the Pacific Basin. Am, Zool. 9:808.

1970. On the ecology and microanatomy of the

parasitic marine prosobranch Mucronaha nitidula Pease,

1860. Proc. Symp. Mollusca. Mar. Biol. Assoc. India

26 THE NAUTILUS

January 31, 1983

Vol. 97(1)

3:780-798.

Waren, A. H. 1980. Description of new taxa of Eulimidae

(Mollusca, Prosobranchia) with notes on some previously

described genera. Zool. Scr. 9:283-306.

. (in press). A generic revision of the family Euli-

midae. J. Molluscan Stud. Suppl.

"TEREBRA" COSENTINI PHILIPPI, 1836,

AN AMERICAN COLUMBELLID SPECIES

Philippe Bouchet and Serge Gofas

Museum National d'Histoire Naturelle

55, Rue de Buffon, 75005 Paris, France

ABSTRACT

Terebra cosentini Philippi, 1836, an alleged Mediterranean species, is shown to

be the correct name for the tropical amphiainerican columbellid generally knawn

as Mazatlania ac\cu\ata (Lamarck, 1822), and should he excluded from Mediterra-

nean faunal lists.

The alleged occurrence of a species of Terebra

in Sicily has long been controversial for Mediter-

ranean conchologists. Although this occurrence

has been denied by a number of malacologists,

the species T. cosentini is still quoted in modern

checklists of the Mediterranean fauna (Paren-

zan 1970; D'Angelo & Gargiullo 1978; Piani 1980

who recognizes its columbellid nature).

De Blainville (1830: pi. 6C, fig. 1) was the first

author to record the presence of this species in

the Mediterranean under the name Buccinum

aciculatum Lamarck, 1822, originally described

from an unknown locality. Blainville's only two

specimens are said to have been received from

Prof. Bonelli from the Provence coast. A few

years later Philippi (1836: pi. 11, fig. 29) in-

troduced the name Terebra cosentini with

Naples, Italy, as the type locality. In volume 2 of

his work, he himself synonymized his T. cosen-

tini with T. cu^iculata (Lamarck). Naples is not

mentioned any longer, but the species "is said to

live in Tarento" Sicily. A year later, the same

species is again described as Buccinum pulchel-

lum by Calcara (1845:41, pi. 4, fig. 23), with

Palermo, Sicily, as type locality.

From the mid- 19th century onwards, we do

not know any original record of this species in

the Mediterranean until modern checklists.

Monterosato, who extensively collected and

FIGS 1 and 2. Mazantlania cosentini (Philippi. 18.36). 1,

Copy of Philippi' s figure 29:17 mm.. 2, Specimen from Las

Caracas, Districto Federal, Venezuela: IJ,.2 rmru

published on the Sicilian malacofauna denied

(1872:59) the Mediterranean origin of the

species. Over the last years, we have visited a

number of Italian private collections, and cor-

responded with a number of collectors in

southern Italy. We could not find any specimens

with accurate Mediterranean locality data.

Shells of this species are rare in collections and

Vol. 97(1)

January 31, 1983

THE NAUTILUS 27

always seem to originate from exchanges and/or

from 19th century collections. They are always

accompanied by such locality data as "Mediter-

ranean" or "Sicily". A number of such samples

are present in MNHN collections. Needlessly to

add, we have never encountered T. cosentini

ourselves in the Mediterranean. With this

evidence we feel certain that the species must be

excluded from the Mediterranean fauna.

The name Terebra anculata (Lamarck) was

first associated with a locality by Hinds (1843):

west coast of central America. Despite Philippi's

synonymizing, both aciculata and cosentini

Vi^ere kept separate in the major 19th century

monographs, due to their widely separate geo-

graphical ranges. A little later, Dunker (1853)

described Terebra nodosoplicata from an

unknown locality. He compared his new species

with Terebra cosentini Philippi and with Buc-

einum clavula Menke, the latter name being ap-

parently a manuscript one. The new taxon was

not figured and has always been regarded as a

synonym of T. aciculata (Lamarck).

The subgeneric name Euryta was introduced

by H. & A. Adams (1858) for a small group of

Terebra comprising aciculata Lam., consent iyii

[sic] V\\\\., fulgurata Phil., granulosa Lam. The

name being preoccupied by Euryta Gistel, 1848,

the new name Mazatlania was proposed by Dall

(1900) without in either case a type species being

designated. Thiele (1929) figured a radula of

Mazatlania aciculata (Lamarck) and transfer-

red the subgenus to the family Columbellidae, as

a subgenus of Pyrene. The designation of

aciculata as type-species oi Mazatlania by Wenz

(1941:1142) appears to be the first valid one.

Examination of samples of M. aciculata from

both sides of central America and comparison

with specimens from historical collections la-

belled as "Mediterranean" T. cosentini leaves no

doubt as to their synonymy.

However, the name Buccinum aciculatum

Lamarck, 1822, is preoccupied by Buccinum,

aciculatum Gmelin, 1791. Thus the correct

name for the amphi-American species generally

known (Radwin 1978:336) as Mazatlania acicu-

lata is M. cosentini (Philippi, 1836). We give

here a copy of the original figure by Philippi and

illustrate a specimen from Venezuela, close to

the type locality as restricted by Radwin.

In conclusion, the synonymy of this columbel-

lid species can be summarized:

Mazatlania cosentini (Philippi, 1836)

Burcinum aciculntum Lamarck, 1822 (mm B. aficulatiim

(Gmelin, 1791)

Terebra cosentini Philippi, 1836

Buccinum pulchetlum Calcara, 1845 (non B. pulcheUum de

Blainville, 1829, nee B. pulcheUum Dujardin, 1837)

Terebra nodosoplicata Dunker, 18.53

Mazatlania hesperia Pilsbry & Lowe, 1932 {Jide Radwin,

1968).

LITERATURE CITED

Adams, H. and A. Adams. 1853. The genera of Recent Mol-

lusca. vol. 1. Van Voorst, London. 484 pp.

Blainville, H. de. 1830. Faiine Frani;aise. Mollusques.

Levrault, Paris. 320 pp.

Calcara, P. 1845. Cenno sui Molluschi viventi efossili delta

Sicilia. Stamperia Reale. Palermo. 65 pp.

Dall, W. H. 1900. Some names which must be discarded.

The Nautilus 14:44-45.

D'angelo, G. and S. Gargiullo. 1978. Guida alle Conchiglie

Mediterranee. ^'abri, Milano. 224 pp.

Dunker, G. 653. Diagnoses Molluscorum Novorum.

Zeitschr. Malak. (1853): 110-117.

Hinds, R. B. 1843. Description of new Shells, collected dur-

ing the voyage of the Sulphur, and in Mr. Cuming's late

visit to the Philippines. Proc. Zool. Soc. Lond. 11:149-168.

Lamarck, J. B. de. 1822. Histoire naturelle des animanx

sans Vertebres, 7 Paris. 678 pp.

Monterosato, A. di. 1872. Notizie intorno alle Conchiglie

Mediterranee. M. Amenta, Palermo. 61 pp.

Parenzan. P. 1970. Carta d'identita delle conchiglie del

Mediten-aneo. vol. 1 (Gasteropodi). Bios Taras, Taranto.

283 pp.

Philippi, R. A. 18.36-1844. Enumeratio Molluscorum

Siciltae. vol. 1 (1836) and vol. 2 (1844). Berlin. 268 +

303 pp.

Piani, P. 1980. Catalogo dei Molluschi conchiferi viventi nel

Mediterraneo. Boll. Malacologico 16:113-224.

Radwin, G. E. 1978. The family Columbellidae in the West-

ern Atlantic Part lib. The Pyreninae (continued). The

Veliger 20:328-344.

Thiele, .J. 1931. Handbuch der systematischeri Weichtier-

kunde, vol. 1. G. Fischer, lena. 778 pp.

Wenz, W. 1941 (in 1938-1944). Handbuch der Paldozoologie.

Gastropoda. 2. G. Borntraeger, Berlin, pp. 949-1639.

Riassunto (Italian Summary)

Terebra cosentini Philippi, 1836, una pretesa

specie mediterranea, e dimostrato essere il cor-

retto nome per il Columbellide tropicale, sia

28 THE NAUTILUS

January 31, 1983

Vol. 97(1)

atlantico che pacifico, generalmente conosciuto dovrebbe essere escluso dalle liste di faune

come Mazatlania aciculata (Lamarck, 1822), e mediterranee.

A NEW SPECIES OF COLUMBARIUM (GASTROPODA: MURICACEA)

FROM OFF EASTERN AUSTRALIA

M. G. Harasewych

College of Marine Studies

University of Delaware

Newark, Delaware 19711

The genus Columbarium Martens 1881 (as

defmec* by Darragh, 1969) is represented in the

Recent fauna by eight species, which inhabit

outer continental shelf and upper continental

slope communities off South Africa, eastern

Australia, New Zealand and Japan. Trawling

operations off the southern coast of Queensland

have brought to light a remarkable new species

of Columbarium, which more closely resembles

several Tertiary species than any in the Recent

fauna. This new species is described herein.

Columbarium harrisae new species

Figs. 1-4

Description - Shell large (to 110 mm), heavy,

fusiform; spire angle 37°-42°; protoconch of 2

whorls, large, bulbous, glassy, with deviated,

disjunct apex; transition to teleoconch indis-

tinct, marked by gradual acquisition of a peri-

pheral keel and axial growth lines; teleoconch

with up to 8 sharply shouldered whorls; suture

adpressed; siphonal canal long, straight, heavy;

spiral scupture of 2-4 cords on body whorl and

20-26 finer threads on siphonal canal; axial

growth lines produce 21-24 short, open spines

per whorl along the shoulder, with correspond-

ing scales on each of the spiral cords and

threads; shell color white with brown spots be-

tween spines and scales; aperture ovate; outer

lip smooth; columella smooth, with a raised peri-

stomal plate that forms a notch below the

suture, and extends along the inner edge of the

siphonal canal; operculum corneous, sharply

ovate, with terminal nucleus; periostracum and

soft parts unknown.

Type material - Holotype - United States Na-

tional Museum, Washington, D.C. (USNM

806997) length 106 mm; Paratype 1 - (USNM

806998) length 70 mm; Paratype 2 - Delaware

Museum of Natural History, Greenville, Dela-

ware (DMNH 153524) length 99 mm; Paratype 3

- The Australian Museum, Sydney, Australia

length 73 mm.

Type locality - East of Lady Musgrave

Island, Queensland, Australia, in 140 fathoms

(256 meters).

Range - Specimens have also been taken off

Lady Elliot Island, Queensland, Australia, in

150 fathoms (275 meters) (Trevor, 1982).

Remarks - The large size, high spire, heavy

shell, stout siphonal canal and characteristic

protoconch readily distinguish this new species

from all other Recent members of the genus.

Columbarium harrisae most closely resembles

C. imlneratum (Finlay and Marwick, 1937) from

the Paleocene of New Zealand, from which it

differs by its larger size and more elongate shell.

Young specimens of C harrisae lack the raised

peristomal plate, and bear a strong resemblance

to C. rugatum (Aldrich, 1886), from the Lower

Eocene of Alabama, but may easily be discerned

on the basis of protoconchs.

The high spire, thick shell and prominent

spiral sculpture are primitive characters within

the Columbariinae, being shared by such groups

as Histricosceptrum, Peristarium and Coluzea.

Vol. 97(1)

January 31, 1983

THE NAUTILUS 29

FIGS. 1-4. Columbarium harrisae rww species. 1, Hulutype. USNM ilUtiinK. dredged east of Lady Musgrave Isiand.

Queensland. Australia, in 256 meters (l.OX). 2, Paratype 1, USNM 806998, same locality and depth (l.OX). 3, Protoconch

of paratype 1 110. OX). 4, Operculum of paratype 1 (.5.0X).

Such features of the Pacific species of Colum-

barium as enlarged protoconch, reduced spiral

ornament, lower spire, and long, thin siphonal

canal are modifications that have arisen since

the closing of the Tethys Sea. Columbarium

harrisae appears to be an offshoot from a

primitive stock and not closely related to the

other Australian members of the genus.

This new taxon honors Valerie Harris of

Caloundra, Queensland, who generously pro-

vided the type material. Thanks are due Richard

M. Kurz, Wauwatosa, Wisconsin, who provided

additional material and information.

LITERATURE CITED

Darragh, T. A. 1969. A Revision of the Family Columbari-

idae (Mollusca: Gastropoda). Proc. Roy. Soc. Vict. 83(1):

63-119.

Trevor. N. 1982. New Species of Columbariidae? Keppel

Bay Tidings 21{l):l.

30 THE NAUTILUS January 31, 1983 Vol. 97(1)

PELYCIDIIDAE, A NEW FAMILY OF ARCHAEOGASTROPOD MOLLUSCS

W. F. Ponder and S. J. Hall

The Australian Museum, Sydney,

N.S.W., Australia 2000

ABSTRACT

A new family is proposed for Pelycidion Fischer ( = AWixia Cossmann and Nan-

noteretispira Habe), a genus of minute gastropods previously associated with the

Rissoacea (Mesogastropoda). The radula is rhipidoglossate and the elongate-

pupiform shell lacks a nacreous layer. The family has a world-wide distribution

in warm temperate to tropical areas and is also known, from the Tertiary of

France. It is provisionally placed in the superfamily Trochaxea.

A generic review of the Rissoidae currently in

progress has shown that many genera included

in that family are wrongly placed. A few cannot

be easily located in any known family and one of

these, Pelycidion Fischer, is the subject of this

review of the group.

Pelycidion vemistuluyn was first named from

Hong Kong and Senegal, West Africa. This

minute, tall-spired species has been included in

the Rissoidae by Thiele (1929), Wenz (1939),

Coan (1964) and Ponder (1967). Cossmann

(1921) was apparently not aware of the ex-

istance oi Pelycidion but included, in the Hydro-

biidae, Allixia, a genus he had previously (1913)

named. This genus name is based on an Eocene

species (Fig. lA, B) from the Paris Basin and is,

in shell characters, identical with Pelycidion.

Allixia recently has been transferred to the

Rissoidae (Gougerot, et al., 1975). Another

genus name, Nannoteretispira Habe, 1961, is

also based on a shell (Fig. IE) virtually identical

to that of the type species of Pelycidion. A

radula and operculum of a single dried animal

loaned by Dr. J. McLean was examined and the

radula was found to be rhipidoglossate. Because

no archaeogastropod family can be used to ac-

commodate the shell seen in Pelycidion, or has

the same radular details, a new family is pro-

posed for it below.

Thiele (1929) and Wenz (1939) list Epigrus

Hedley as a synonym of Pelycidion. The type

species (Rissoina cylindracea T. Woods, 1878)

of that genus, however, is much larger than

species of Pelycidion and has a taenioglossate

radula (personal observation).

Abbreviations

AMS - The Australian Museum. Sydney

IRSB - Institut Royal des Sciences Naturelles de Belgique,

Brussels

LACM - Los Angeles County Museum of Natural History,

Los Angeles

NSMT - National Science Museum, Tokyo

OM - Zoology Museum, Oxford University, Oxford

USNM - National Museum of Natural History, Washington,

D.C.

Family Pelycidiidae family nov.

Diagnosis - Shell minute, elongate pupiform,

imperforate, with smooth or finely spirally

striate teleoconch; protoconch paucispiral or

multispiral, with reticulate or spiral sculpture.

Aperture subcircular, simple, peristome contin-

uous. No inner nacreous layer. Operculum

horny, circular, with central nucleus. Radula

rhipidoglossate, c.l5 -i- 2 -i- 1 -t- 2 + c.l5, central

teeth simple, without lateral thickening, about

V2 size of lateral teeth, central and lateral teeth

multicuspate, marginal teeth small, unicuspid.

Head-foot and anatomy unknown.

Remarks - The new family is distinguished

from other rhipidoglossate families by the com-

bination of characters given in the diagnosis. Its

relationships are obscure although it can be re-

garded as trochacean. A rather heterogeneous

family that appears to show some similarities,

the Skeneidae, has radulae with short central

Vol. 97(1)

January 31, 1983

THE NAUTILUS 31

FIG. 1. Shells o/Pelycidion. A. B, Pelycidion acicularis (Cossmann). Bercheres, Eure-et-Loir, Pans Basin, France (Lutetien..

Eocene): B, detail of protoconch: C, D, Pelycidion \a.ni\\?i\s (Watson), Masthead Islanct. Queensland. Australia, 31-S6 m (AMS);

D.'diHail of protoconch; E, Pelycidion sp. Dahoiney, West Africa, 5.5 m (LACM); F, G, Pelycidion venustnlum Fischer (\n Foiin

& Perier). Bale di Cansado. Cap Blanc. Mauritania, West Africa (IRSB). H. Pelycidion japonicus (Hai<V. holotype. Scales:

shells = 0.1 mm: protoconchs = 0.05 ynm.

32 THE NAUTILUS

January 31, 1983

Vol. 97(1)

teeth with lateral thickenings and, as in the

other trochacean families, the marginal teeth

are multicuspate. Skeneids have 2-4 pairs of

lateral teeth and the shell has no inner nacreous

layer. The shells of the Skeneidae are usually

depressed-trochiform to subplanate and fre-

quently umbilicate. In addition the protoconch is

paucispiral in all known species. The radulae of

Liotiidae, Trochidae and Turbinidae have more

than 3 pairs of lateral teeth. The Phasianellidae

and Turbinidae have calcareous opercula and

differ markedly in radular features. The only

trochacean genus that is similar in general shell

features is Halistylus Dall. This genus has a

markedly different radula from Pelycidion, hav-

ing very long marginal teeth and weakly-

defined, broad-lateral and central teeth each

with only a single, weak cusp (personal observa-

tion).

Genus Pelycidion Fisher

(in Folin & Perier, 1873)

Type species — Pelycidion venustulum Fischer

(in Folin & Perier, 1873, p. 182) by monotypy.

Recent, Senegal, West Africa (here restricted).

Synonymy -

Pelycidion Fischer {in Folin & Perier, 1873): 316: nomen

nudum.

Pelecydium err. auct.

Allixia Cossmann, 1913: 141. Type species: Allixui acicu-

laris Cossman, 1913; original designation. Middle Eocene,

Parish Basin.

Nannoteretispira Habe, 1961: 273. Type species: Nanno-

teretispira japonica Habe, 1961; original designation. Re-

cent, Japan.

Diagnosis - Shell minute, elongate-pupoid,

with subcircular aperture with simple peris-

tome, outer lip prosocline; protoconch of IV2 to

about 3 whorls, weakly spirally sculptured if

paucispiral, complexly sculptured with raised

reticulate threads if multispiral, nucleus smooth

or minutely pitted (Figs. IB, D, 2C, D, 3C, D).

Teleoconch of several (about 5-6) whorls, ap-

parently smooth or with spiral striae (Figs. lA,

C, E-G, 2A, B, 3A, B, E). Head-foot unknown.

Operculum: circular, with central nucleus,

horny, number of whorls not known (Fig. 2E).

Radula - Rhipidoglossate, with relatively

large central teeth 4-1-1-1-4, rather long,

parallel-sided, cusps small and sharp, primary

cusp about twice length of adjacent cusps.

Lateral teeth elongate, with narrow, long, sim-

ple bases and recurved cutting edge bearing

long, sharp cusps; inner lateral teeth with c.6

cusps, outer lateral teeth with wider cutting

edge than inner lateral teeth and with c.9 cusps.

Marginal teeth small, curved, with a single cusp,

C.15 per V2 row (Fig. 21, G).

Distribution and members of the genus Pely-

cidion - Indo-Pacific: (Mucronalia xanthais

Watson, 1886 (Fig. IC, D) ? = iV. japonica Habe,

1961 (Fig. IE)); South Africa: (Modulus afri-

canus Bartsch, 1915 = A^. curiosus Turton, 1932

and A'^. becki Turton, 1932); eastern Pacific:

(Modulus kelseyi Bartsch, 1911); Caribbean:

(Modulus megalomastomus Olsson and McGinty,

1958 (Fig. 3A)); West Africa: (P. venustulum

(Fig. IF, G) and P. sp. (Fig. 3B-D)); Eocene

France: (A. ancularis (Fig. 1 A) (see Gougerot et

al. 1975 for a recent revision of the Tertiary

(Eocene- Pliocene) species and subspecies from

France)).

Material examined - P. venustulum; one lot

so named, Dautzenburg Colin. (IRSB, IG10591)

one specimen (LACM). A'', japonica; holotype

and paratype (NSMT, 39823). A^. africanus;

holotype (USNM, 250422), 4 specimens

(BMNH), 3 specimens (OM). A^. curiosus and A^.

becki Turton, 1932; types (OM). A^. kelseyi; holo-

type (USNM, 111369), a few lots ex LACM

(AMS). A. cwicularis; 1 lot ex J. le Renard

(AMS). M. xanthais; several lots (AMS). M.

megalomastomus; holotype and paratype

(ANSP); two specimens ex D. Moore (AMS).

Remarks - Fischer records his species (P.

venustulum) from Hong Kong and Senegal,

West Africa. No type material has been located

but 4 specimens, only one of which is in

reasonably good condition, are identified as this

species in the Dautzenberg collection (IRSB,

IG10591). These specimens (Fig. IF, G) from the

mission Gruvel (1909-1910) from dredgings in

Bale di Cansado, cap Blanc, south of Port-

Etienne (now Nouadhibou), Mauritania, West

Africa (ex Bavay), agree closely with the original

description of the species and with Mannofereti-

s-pir a japonica Habe (Fig. IE) and with Mucro-

nalia xanthias Watson (Fig. IC, D). Another

specimen, in better condition, from 12 miles E.

Vol. 97(1)

January 31, 1983

THE NAUTILUS 33

«ii55g

FIG. 2. Pelycidion <:/: kelseyi (Bartsch). W. undt' ofEa^-it Idand. San Benito Jdandji. Baja California, Mexico

(LACM). A-E, shelU; B, microaciilpture; C, D, pnitoconch: D, trucrosrulpture of protoconch; E, operculum, in-

ner sidf: F. G, radula. Scaler: A. C. E = 0.1 >nm: B. D = 0.01 mm: F. G = 0.001 mm.

34 THE NAUTILUS

January 31, 1983

Vol. 97(1)

--■*-

B

C I-

^/ m

^t

H

H

D

HG. 3. A, Pelycidion niegalomaslom;i (Ol.sson & MrGinly). shell, 2 m. Glovers Reef Lagoon. N.E. British

Honduras (AMS). B-D, Pelycidion sp.. 55 m, 19 km. E. ofCotonou. Dahomey. WV.s/ Africa (LACM). B, shell: C,

protoconch: D, protoronch microsadpture: E, teleoconch microsculpture. Scales: A. B. C = 0.1 mm: D, E =

0.01 mm.

Vol. 97(1)

January 31, 1983

THE NAUTILUS 35

of Cotonou, Dahomey, West Africa (6°24'N,

2°31'E), 55 m. (Fig. 2E) is smaller and has fewer

whorls. It is possibly a different species (Fig.

3B-D). Specimens of Allixia acicul.a'r':s Coss-

mann (Fig. lA, B) also appear to be congeneric

as do several other species listed above. The

variation in the sculpture and number of whorls

of the protoconch is not considered to be impor-

tant at the generic level in view of the close

similarity of the other shell features.

The interspecific differences in the protoconch

are probably due to the adoption of different life

history strategies. The West American species

and its Caribbean analogue have a paucispiral

protoconch with a large initial whorl (Figs. 2C,

3A) suggesting that direct development occurs

in these species. A multispiral protoconch (Figs.

IB, D, 3C) suggests a planktotrophic larval

stage. This type of protoconch is atypical of the

Archaeogastropoda but is commonly encoun-

tered in the Mesogostropoda, Neogastropoda

and Heterogastropoda.

The original figure of P. venustulum shows a

shell with a markedly convex inner lip. It is here

assumed that this feature is erroneously

depicted. This inaccuracy presumably led Tryon

(1887) to suggest that Hemistomia Crosse, a

genus in the Hydrobiidae, might be a synonym

of Pelycidion.

ACKNOWLEDGMENTS

We would like to thank the curators responsi-

ble for the molluscan collections in the USNM

and OM for facilities being made available to the

senior author to study type material. Specimens

of P. vpniistulum were loaned by Dr. J. Van

Goethem, IRSB and the holotype of A^. japonica

was loaned by Dr. T. Habe, then of NSMT. Pro-

fessor J. le Renard kindly donated specimens of

A. acicidaris to The Australian Museum. Dr. J.

McLean of the Los Angeles County Museum

loaned and donated west American material, in-

cluding the specimen containing a dried animal

and Dr. D. Moore donated two specimens of N.

megalomastomus. Dr. P. Bouchet assisted with

the locality details of the specimens of P.

venustulum. Mr. E. K. Yoo and one of us

(S.J.H.) are responsible for the S.E.M. work

which was carried out in the Electron Micro-

scope Unit, University of Sydney. Miss B.

Duckworth and E. K. Yoo did the drawings. We

thank Dr. C. S. Hickman for critically reading

the manuscript. This work was supported in

large part by an Australian Research Grants

Committee grant to the senior author.

LITERATURE CITED

Coan, E. 1964. A proposed revision of the rissoacean fami-

lies Rissoidae, Rissoinidae, and Cingulopsidae (Mollusca:

Gastropoda). Veliger 6(3):164-171.

Cossmann, M. 1913. Catalogue illustre des coquilles fossiles

de I'Eocene des environs de Paris . . . App. 5 Annates de

la Societe Malacologique dt Belgique 49:19-238, pis. 1-8.

1921. Essais de Paleoconchologie Comparee

12:1-336, pi. 1-11, Paris.

Folin, L. de and L. Perier. 1867-1886. Lesfonds de la nier,

etude internatu lale sue les particuliarites nouvelles des

regions sous-^ >w. 4 volumes, Paris.

Gougerot, L, I'^ekih and J. le Renard. 1975. Le genre

Allixia C' .nann (Gastropoda: Rissoidae) sa position

sysiematique et sa longevite. Cahiers des Naturalistes.

Bulletin des Naturalistes Parisiens 31(2):41-48.

Habe, T. 1961. Three new gastropods from Japan. Venus,

Japanese Journal of Malaeology 21(3):270-274.

Olsson, A. A. and T. L. McGinty. 1958. Recent Marine Mol-

lusks from the Caribbean Coast of Panama with the De-

scription of Some New Genera and Species. Bull. Amer.

Paleontology 39:1-58, 5 pis.

Ponder, W. F. 1967. The classification of the Rissoidae and

Orbitestellidae with descriptions of some new taxa. Trans-

actio7ts of the Royal Society of New Zealand, Zoology

9(17):193-224.

Ponder, W. F. and E. K. Yoo. 1976. A revision of the Aus-

tralian and Tropical Indo-Pacific Tertiary and Recent

species oi Pisinna I =Estea) (Mollusca: Gastropoda: Rissoi-

dae) Record.^ of the Australian Museum 30(10):150-247.

Thiele, .J. 1929(-1935). Handbuch der systematischen Weich-

tierkunde 1. Jena, 376 pp. (1929).

Tryon, G. W., 1887. Manual ofConchology 9.

Wenz, W., 1938-1944. Gastropoda. Handbuch der Paldo-

zoologie 6(1), Lief 1-7. 1639 pp.

36 THE NAUTILUS

January 31, 1983

Vol. 97(1)

DEPTH DISTRIBUTION AND DENSITY OF FRESHWATER MUSSELS

(UNIONIDAE) COLLECTED WITH SCUBA FROM THE

LOWER WISCONSIN AND ST. CROIX RIVERS

Edward M. Stern

Department of Biology

University of Wisconsin-Stevens Point

Stevens Point, WI 54481

ABSTRACT

Using SCUBA diving, the depth distribution ajid density of freshwater mussels

(Unionidae) in the Wisconsin and St. Croix rivers were examined. Depth is a fac-

tor in determining the distribution of mussels only because it is a reflection of cur-

rent velocity and substrate type. The distribution of freshwater mussels in lotic

habitats was most closely correlated with composition of the substrate. The

greatest species diversity as well as both the highest mean (3. 7/m^) and maximum

(601 m^) densities included substrates with a mixture of particle sizes from mud. to

boulder. Only species o/Anodonta and Lampsilis were characteristic inhabitants

of mud and! or stable sand substrates. The mean density was significantly lower

in a mud-sand bottom, with a value of 0.9 mu^sselslm^. Mussels were absent in a

shifting sand substrate. Baker (1928) reported a total of 28 species from the

Wisconsin River and 15 species from, the St.. Croix River, whereas 25 and 11).

species, respectively, were collected during this study.

Few studies treating the freshwater mussels

(Unionidae) of Wisconsin have been conducted

since Baker (1928) published his comprehensive

monograph. Of the nearly 60 forms reported

(approximately 45 species using present con-

cepts of a species), most formerly occurred or

currently occur in the Mississippi River and/or

in two of its major tributaries, the Wisconsin

and St. Croix rivers. Because an excellent sum-

mary of the unionids of the Mississippi River in

the vicinity of Prairie due Chien, Wisconsin, was

recently published by Havlik and Stansbery

(1977), the emphasis of this study is on the latter

two rivers.

SCUBA diving has been used successfully to

study lacustrine molluscs (Cvancara, 1972; Pace

et aL, 1975, 1979; Ghent et a/., 1978). The use of

SCUBA provides quantitative data on the depth

distribution and density of mussels as well as in

situ habitat observations. However, because

poor visibility and strong currents usually limit

its use in lotic habitats, comparative data are

few.

Methods and Materials

SCUBA diving was conducted during low

water periods in August and September 1978 at

five sites along the Wisconsin and St. Croix

rivers (Fig. 1). Because of the paucity of the

molluscan fauna in the heavily impounded mid-

dle one-third of the Wisconsin River, only locali-

ties well below the last dam were examined.

Fourteen transects, each measuring 2 m by 20

m, were examined during a total of 26 hours of

diving time. The transect lines were constructed

of nylon rope 6 mm in diameter and weighted at

5 m intervals with 2 kg lead weights. Plastic

milk cartons were attached at 10 m intervals to

serve as buoys and the entire transect was an-

chored in place with stakes. At each site,

transects were established at several depths

with maximum depths of 2.5 m and 3.5 m in the

Wisconsin and St. Croix rivers, respectively.

Because seasonal fluctuations in water level in

the Wisconsin and St. Croix rivers can be sub-

stantial, depth determinations are not absolute.

All collection depths represent low water levels.

Vol. 97(1)

January 31, 1983

THE NAUTILUS 37

FIG. 1. Collection localitic.-! on the Wisconsin (1-i) and St.

Croix 15) rivers. Localities are identified in the text.

and high water levels may be an additional 1 to

2 m. All mussels collected within each 40 m^

transect were placed in bags and transported

back to the laboratory for identification. Den-

sities were determined at random within each

transect using a 0.25 m^ wire frame. A number

of physico-chemical parameters were measured

at each site using Hach colorimetric and

titrametric procedures. A determination was

made of the type of substrate at each transect

using the following modification of the Went-

worth scale for particle size: mud < 0.06 mm;

sand 0.06-2.0 mm; gravel 2.0-64.0 mm; and

boulder > 64.0 mm.

Mussels were also qualitatively collected by

hand at several additional sites. Although these

data are not quantitative in relation to surface

area sampled and collection methods varied,

they are used because they include species not

encountered while diving.

Voucher specimens have been deposited in the

Museum of Natural History, University of

Wisconsin-Stevens Point.

Collecting Stations

The following localities, identified in Fig. 1,

were examined for mussels:

1. Wisconsin River at County Park in Dekorra.

(Columbia Co.)

2. Wisconsin River at intersection of Wisconsin

Highways 60 and 00. (Richland Co.)

3. Wisconsin River at intersection of Wisconsin

Highways 60 and E at public landing. (Rich-

land Co.)

4. Wisconsin River at intersection of Wisconsin

Highways 60 and TX at wayside. (Richland

Co.)

5. St. Croix River, T. 35 N, R. 19 W, Sec. 9.

(Polk Co.)

Results and Discussion

Physico-chemical determinations at each

locality are presented in Table 1. Because these

data represent only a single test at each locality,

water quality data compiled by the U.S. Geologi-

cal Survey (1978) for the water year 1977 are

also included for the purpose of valid compari-

son. When compared with the most current data

for those physico-chemical parameters that af-

fect the physiological ecology of freshwater

mussels, all of the parameters measured were

within currently accepted levels for the suc-

cessful propagation and growth of freshwater

mussels as summarized by Fuller (1974). The

current at all sites was moderately swift with an

average surface velocity of 0.6m/sec.

The depth distribution for 21 species in dif-

ferent types of substrates for all transects is

summarized in Table 2. Seven species are not in-

cluded, because too few individuals were col-

lected, and are discussed separately below.

Mussels were not located uniformly across the

river bed. In a lake study, Harman (1972) quan-

titatively demonstrated the relationship be-

tween mollusc distribution and substrate pat-

terns as well as the correlation between species

diversity and substrate diversity. Thus depth is

a factor in governing the horizontal distribution

of mussels primarily because it is a reflection of

38 THE NAUTILUS

January 31, 1983

Vol. 97(1)

TABLE 1. Physico-chemical data for localities on the Wisconsin (l-U) and St. Croix (5)

rivers. Localities 1-5 are identified in Fig. 1 Data for localities 6 (Wisconsin Rii>er at

Muscoda) and 7 (St. Croix River at St. Croix Falls) are from the U.S. Geological Survey

(1978) and are mean values representing measurements for the water year October 1976 to

September 1977.

gradations in current velocity and substrate

type.

In my study, a greater species diversity and

mean and maximum density of mussels were

associated with those transects that included a

bottom with a mixture of particle sizes ranging

from sand to boulder (Table 2). Only seven of the

28 species collected were found in a mud-sand

substrate. Based upon the percent of the in-

dividuals of each species collected in each

substrate type (Table 2), of these seven species a

mud-sand substrate was the typical habitat for

only Anodonta grandis, Lampsilis anodon-

toides, and L. radiata. The remaining four

species were more abundant in a mixed sand-

gravel-boulder substrate (Table 2). The mean

density for each substrate type at each depth

also illustrates these conclusions (Table 3). The

maximum density recorded was 60 mussels/m^

at a transect depth of 1.7 m in a sand-gravel-

boulder substrate.

Cvancara (1972) and Ghent et al. (1978) com-

pared the adaptive morphology of several

species of unionids from diverse habitats by

measuring shell width and angle of the ventral

margin. The greater obesity and ventral angle

exhibited by A. grandis and L. radiata are

believed to be morphological adaptations per-

mitting them to present a wide surface to a soft

substrate in ponds, sloughs, and pools (Ghent et

«i., 1978). Conversely, a narrow ventral angle is

ideally suited to a gravel bottom in running

water. The results of my SCUBA study further

substantiate this interpretation because the

same two species were the most abundant in and

characteristic of predominately mud and sand

substrates (Table 2). (A discussion of these data

is the subject of a future paper.) The lowest

mean density/m^ occurred in a mud-sand bottom

(Table 3).

Only six of the 28 species collected during this

study (A. plicata, E. dilatatuii, F.Jlava, A. gran-

dis, A. carinata, and L. radiata) exhibited a

distribution that included all substrate types ex-

cept shifting sand. Mussels were absent entirely

in a shifting sand bottom. Therefore, depth is

only a factor in determining the density and dis-

tribution of freshwater mussels as a reflection of

current velocity and substrate type.

Many of the distributional records in Baker's

Vol. 97(1)

January 31, 1983

THE NAUTILUS 39

TABLE 2. Depth distribution of freshwater mussels fur

each substrate type: number of individuals, with % of the

species population for each depth-sufjstrate combination ni

parentheses. Data are composite totals for each species at all

transects. *No transects were established at depths between

2.0 and 2.7 to. Genera are identified in Table i. ini = mud:

s = sand; g = gravel: b = boulder.)

T.'XBLE 3. Mean density (f freshwater m.u.'isels for eaeh

depth-substrate combination. Data include all transects.

lM=niud: s = s(uid: g = graret: l) = houlder.)

B-S-g

•-g-b

0.9/»

l.V"

3.1/.

3.6/.^ 3.7/.^

3.1/.'

(1928) survey of Wisconsin were based upon

localities along the Wisconsin antJ St. Croix

rivers. Baker (1928) reported 28 species from

the Wisconsin River and 15 species from the St.

Croix River, while during this study 25 and 14

species, respectively, were collected. A sum-

mary of these records is presented in Table 4.

Although the Baker (1928) records are aifficult

to compare because some species were listed

simply as being statewide or as occurring in the

Mississippi River system, these data do reveal

some changes in species distribution and abun-

dance, especially with regard to several of the

species not included in Table 2.

Although Barnes (1823) cited the Wisconsin

River as the type locality for C. verrucosa

( = tuberculata), this species was not collected by

Baker (1928) nor by me. Cydonnias tuberculata

has also been extirpated from the Mississippi

River at Prairie du Chien (Havlik and Stans-

bery, 1977), where it was last collected by Ellis

in 1930 (van der Schalie, 1950). Today, a tuber-

culata may be restricted to the St. Croix River

and populations here may represent the last

significant ones within the State.

Baker (1928) noted that F. ebeyia was rare in

both the Mississippi and Wisconsin rivers.

Williams (1978) recently collected two in-

dividuals from the St. Croix River near Hudson,

40 THE NAUTILUS

January 31, 1983

Vol. 97(1)

TABLE 4. The freshwater mivisels recorded from the

Wisconsin and St. Croix rivers l/y Baker I192S) and Stern

(this study). Asterisk (*) denotes those species listed .•iimply

(US statewide by Baker (1928). Classification follows that of

Ortmatin (1910) and the species are arranyed alphabetically

under each subfamily.

reported it from the Mississippi River at Prairie

du Chien. No sliells were found while diving.

Plethobasus cyphyus has been extirpated from

the Mississippi River at Prairie du Chien (Havlik

and Stansbery, 1977), where it was last collected

by Shimek (1921). Baker (1928) reported it from

Lake Pepin to the north and indicated that it

was common in the Wisconsin River. Only two

live specimens were collected in the Wisconsin

River while diving, both from a sand-gravel-

boulder substrate in water 1.3-1.7 m deep.

Because a greater number of subfossil shells

were also collected, the species is probably

disappearing from Wisconsin waters.

Baker (1928) reported several "forms" ( = eco-

phenotypes) of P. coccineum from the Wisconsin

River. Havlik and Stansbery (1977) found only

one shell at Prairie du Chien. While diving, a

total of only three live individuals were col-

lected. Where collected live, P. coccineum was

found in water 1.3-1.7 m deep in a mixed mud to

boulder bottom.

Baker (1928) stated that Q. fragosa ( = quad-

rula) in Wisconsin was restricted to the Wiscon-

sin River drainage. Although he also recorded

two other closely related forms from the State,

they were also confined to single drainages.

Baker (1928) noted that Q. quadrula s.l. (sensu

lato-in the broad sense) was neither widely dis-

tributed nor abundant. Only one live specimen

was found while diving in the Wisconsin River

during this study. It was collected from a sand-

gravel-boulder substrate in L5 m of water.

Havlik and Stansbery (1977) found no Q. fragosa

at Prairie du Chien and concluded that this

ecoform is now reduced or estirpated through-

out much of its range.

Like A. grandis, A. imbecillis is a typical in-

habitant of quiet pools in a mud or stable sand

bottom. Baker (1928) noted that it was widely

scattered throughout the State and rare when

found. A single individual of A. imbecillis was

collected in shallow water (1 m) in a mud-sand

bottom from the St. Croix River.

Baker (1928) reported Arcidens confragosus

only from the Mississippi River, where it still oc-

curs (Havlik and Stansbery, 1977). One live and

two subfossil specimens were collected during

this study from the Wisconsin River. Through-

out most of its range, it is most abundant in a

Vol. 97(1)

January 31, 1983

THE NAUTILUS 41

mud bottom in sluggish water at shallow depths

(<1 m), but in the Wisconsin River it was col-

lected from a sand-gravel bottom in a moderate

current at a depth of 1.7 m. An emphasis on col-

lecting in the former habitat might reveal A.

confragosus in larger numbers.

Baker (1928) collected several shells, but no

living specimens, of S. ambigiia on a gravel bar

in 0.3 m of water in the Wisconsin River. Only a

single specimen was collected by Ellis in 1930 in

the Mississippi River at Prairie du Chien (van

der Schalie, 1950). Simpsoniconcha ambigua

was probably never common, and it was not en-

countered while diving. It has a unique glochi-

dial host, the mudpuppy Nectunis maculosus.

An examination of those sites at which its am-

phibian host is known to occur might help to

clarify the distribution of this unionid. Stans-

bery (1970, 1971) has noted thatS. ambigua may

be endangered throughout its entire range. Its

distribution is sporadic and, when encountered,

it is seldom abundant.

The absence of L. anodontoides in the Wiscon-

sin River puzzled Baker (1928) because the river

seemed "ecologically well suited for the species."

Lampsilis anodontoides was collected during

this study while wading in shallow water (0.3 m)

in a stable sand bottom. Despite its apparent

ability to adjust to a variety of habitats from

mud to sand to gravel bottoms, in either a swift

or slow current, and at varying depths (Baker,

1928; Murray and Leonard, 1962; Parmalee,

1967), no live individuals were collected while

diving.

Most of the faunal changes over the last 50

years, as discussed above, are attributable to

man's activities, including changes in water

quality, elimination of host fishes, commercial

overexploitation, and/or the creation of large

impoundments that profoundly and permanent-

ly alter habitats. The construction of over one

dozen dams along a 125-mile stretch of the mid-

dle one-third of the Wisconsin River illustrates

the latter factor.

In a recently completed survey, Mathiak

(1979) reported five species (Quadrula nodulata,

Anodontoides fertissacianus, Lasmigona com-

planata, L. compressa, and Lampsilis higginsi)

from the Wisconsin River that were neither

reported by Baker (1928) nor collected while div-

ing. Most of these were found just upstream

from the confluence of the Wisconsin and

Mississippi rivers and thus represent recent

range extensions.

It is apparent that several species are now

rare and in danger of being extirpated from

Wisconsin waters. However, it is encouraging

that not only is the same basic fauna still

represented some 50 years after Baker's (1928)

study, but that there has been the establishment

of additional species as well.

ACKNOWLEDGMENTS

I wish to thank the following for their

assistance in the field: Dr. William LeGrande,

Pam Gomez, Dave Timm, and especially Rick

Rothman, who helped me with the diving. I also

want to thank Dr. Ruth L. Hine, the Wisconsin

Department of Natural Resources, and the En-

dangered Species Project (E-1) for financial sup-

port provided. Finally, I want to thank

Catherine Stanly for typing this manuscript.

LITERATURE CITED

Baker, F. C. 1928. ne freshwater MoUusca of Wisconsin.

Part II. Pelecypoda. Bull. Wis. Geol. & Nat. Hist. Surv.

70:1-495.

Barnes, D. W. 1823. On the genera Unio and Ala.smodonta:

with introductory remarks. Am. J. Sci. 6:107-127;

258-280.

Cvancara, A. M. 1972. Lake mussel distribution as deter-

mined with SCUBA. Ecol. 53:154-157.

Fuller, S. L. H. 1974. Clams and mussels (Mollusca: Bival-

via). In Pollution Ecology of Freshwater Invertebrates,

C. W. Hart. Jr. and S. L. H. Fuller, eds. Academic Press,

New York. pp. 215-273.

Ghent, A. H., R. Singer and L. J. Singer. 1978. Depth dis-

tributions determined with SCUBA, and associated stu-

dies of the freshwater unionid clams EUiptio c(miplanata

and Anodonta grandis in Lake Bernard, Ontario. Can. J.

Zool. 56:1654-1663.

Harman, W. N. 1972. Benthic substrates: their effect on

freshwater Mollusca. Ecol. 53:271-277.

Havlik, M. E. and D. H. Stansbery. 1977. The naiad mollusks

of the Mississippi River in the vicinity of Prairie du Chien,

Wisconsin. Bull. Am. Malatol. Union 1977:9-12.

Mathiak, H. A. 1979. A river survey of the unionid musseh

of Wisconsin 1973-1977. Sand Shell Press, Horicon, WI.

75 pp.

Murray, H. D. and A. B. Leonard. 1962. Handbook of

unionid mussels in Kansas. Univ. Kansas Mus. Nat. Hist.,

Misc. Pub. 28:1-184.

42 THE NAUTILUS

January 31, 1983

Vol. 97(1)

Ortmann, A. E. 1910. A new system of the Unionidae. The

A'oMh/Mt.- 23:11-4-120.

Pace. G. L.. E. J. Szuch and R. \V. Dapson. 1975. SCUBA

assisted studies of freshwater snails. Bull. Am. Malncol.

Union 7.975:68.

1979. Depth distribution of three gastro[)ods in

New Mission Bay, Lake Michigan. The Nautilus 93:S1-'M.

Parmalee, P. W. 1967. The fresh-water mussels of Illinois.

Illinois State Mus., Sci. Ser. 8:1-108.

Shimek, B. 1921. Mollusks of the McGregor. Iowa region.

Iowa Conserv. 5:1.

Stansbery, D. H. 1970. Eastern freshwater mollusks. The

Mississippi and St. Lawrence River systems. In Papers on

the rare and endangered mollusks of North America,

A. H. Clarke, ed. Malacologia 10:9-21.

1971. Rare and endangered freshwater mollusks

in eastern United States. In Proc. of a symposium on rare

and endangered mollusks (naiads) of the United States,

S. D. Jorgensen and R. W. Sharp, eds. U.S. Dept. Interior,

Fish and Wildlife Serv., Bur. Sport Fish, and Wildlife.

Twin Cities, Minnesota, pp. 5-18.

U.S. Geological Survey. 1978. Water resources data for

Wisconsin Water Year 1977. U.S. Geol. Surv., Water Res.

Div., Madison, Wisconsin.

van der Schalie, H. and A. 1950. The mussels of the Missis-

sippi River. Am. Midi. Natur. 44:448-466.

Williams, D. D. 1978. Aspidogaster conchicola in St. Croix

River, Wisconsin clams. Proc. Helminthological Soc.

Wa.sh. 45:257-258.

A PRE-EUROPEAN OCCURRENCE OF GLEBULA ROTUNDATA

(BIVALVIA: UNIONIDAE) IN ARKANSAS

Mark E. Gordon

Department of Zoology

University of Arkansas

Fayetteville, Arkansas 72701

Glebula rotunda t a (Lamarck) is reported for

the first time from Arkansas. It is distributed

from eastern Texas to the Apalachicola River in

Florida (Clench and Turner, 1956) and has been

found primarily within about 200 km of the Gulf

of Mexico (Parker, personal communication); al-

though, Branson (1969) reported a specimen

from the Neosho River system in Oklahoma.

Speculation on the presence of G. rotundata in

Arkansas had been made by Call (1895) and Gor-

don, et al. (1980).

A specimen of Glebula rotundata (University

of Colorado Museum no. 30468) was identified

from a group of about 6000 specimens collected

from the Tillar Farms locality, Drew County, of

the Arkansas Archaeological Survey. The site

represents a silted-in oxbow of Bayou Barthole-

mew and dates from between 1400 and 1600

A.D. All specimens represent a naturally occur-

ring assemblage and did not represent an Indian

midden. The site is within the region of the state

in which Call (1895) expected G. rotundata to be

found. Living specimens have not been found, as

yet, in Bayou Bartholemew.

I would like to thank Dr. Neal Trubowitz,

Arkansas Archaeological Survey, and Dr. Mar-

vin Jeter, University of Arkansas-Monticello,

for information concerning the Tillar Farm

locality and Mr. Robert S. Parker, Freeport

Sulphur Co., Belle Chasse, La. for distributional

information.

LITERATURE CITED

Branson, B. A. 1969. Glebula in Oklahoma. Sterkiana 36:22.

Call, R. E. 1895. A study of the Unionidae of Arkansas, with

incidental reference to their distribution in the Mississippi

Valley. Trans. Acad. Sri. St. Louis 7:1-65.

Clench, W. J. and R. D. Turner. 1956. Freshwater mollusks

of Alabama, Georgia, and Florida from the Escambia

River to the Suwannee River. Bull. Fla. State Mus.

1:99-239.

(iordon, M. E., L. R. Kraemcr and A. V. Brown. 1980.

Unionacea of .Arkansas: historical review, checklist, and

observations on distributional patterns. Bull. Ani. Malac.

Union 1979:31-37.

Vol. 97(1)

January 31, 1983

THE NAUTILUS 43

RECENT DEATHS

Harry S(tephen) Ladd, retired paleontolo-

gist, died Noveml)er 30, 1982, at the age of 82,

in Bethesda, Maryland. Although primarily an

expert on Pacific Island coral reefs and the Ter-

tiary mollusks of the Southwest Pacific, Dr.

Ladd was well-known to malacologists, and was

always helpful to the many students that visited

the U.S. National Museum. He was born Jan. 1,

1899, in St. Louis, Missouri, and obtained his

Ph.D. at the University of Iowa. He spent two

years mapping the geology of the Lau Group in

Fiji. He began his distinguished career at the

U.S. Geological Survey in 1940, and continued

research at the U.S. National Museum until

1978. He received the Distinguished Service

Award of the Interior Department in 1965 and

the Paleontological Society Medal in 1981. He

published over 60 articles, mainly on fossil

mollusks, and was a contributor to The

Nautilus. Additional information in Ajner'ican

Malacologists, p. 334, and the Washington Post.

p. B18of Dec. 8, 1982.

Charles B. Wurtz, consulting biologist, en-

vironment ecologist, and former Associate

Editor of The Nautilus (1958-74), died October

21, 1982, at the age of 65, in Philadelphia, PA.

Son of a physician, he was born Dec. 6, 1916, in

Philadelphia. He was a student of land mollusks

under Drs. H. A. Pilsbry and H. B. Baker, and

received his Ph.D. in 1955 at the University of

Pennsylvania. He led many limnological surveys

for the Academy of Natural Sciences of Phila-

delphia from 1948 to 1954, and later began his

own environmental consulting service. He

taught biology at La Salle College from 1963 to

1971. Dr. Wurtz published about 70 articles,

many of them appearing in The Nautilus.

"Chuck" as he was known to his many friends

was always helpful to fellow biologists. His in-

fectious laugh, good cheer and keen mind will be

missed by many. He is survived by his wife, Elsa

M. Hofheinz Wurtz. Additional information in

American Malacologists, p. 489 and La Salle

(Quarterly College Magazine), Fall 1966, vol. 10,

pp. 8-12."

William E(rwood) Old, Jr., malacologist,

died of a heart attack, on December 31, 1982, at

age 54, in New York City. He was born April 14,

1928, in Norfolk, Virginia, and attended the Col-

lege of William and Mary. He served in the

Army during the Korean War. Bill Old joined

the American Museum of Natural History in

1960, and became a specialist in marine mol-

lusks, pul)lishing many papers, some in conjunc-

tion with W. K. Emerson. Bill was extremely

devoted to his science, and was well-known as a

Charles B. Wurtz (191ti-19S2)

Williiim K. Old. Jr. (192»-i98-J)

44 THE NAUTILUS

January 31, 1983

Vol. 97(1)

shell show judge in many parts of the United

States. He was constantly assisting amateur, as

well as professional, malacologists. He went on

several expeditions, including ones to the Gala-

pagos and Mexico, and helped build the Ameri-

can Museum's mollusk collection into a leading

research resource. Friends of Bill Old may make

tribute to his memory by contributing to the

William E. Old Malacology Fund, c/o Dr. W. K.

Emerson, American Museum of Natural His-

tory, Central Park West, NY, NY 10024.

-R.T.A.

BOOK REVIEWS

Sea Shells of Southern Africa by Richard

Kilburn and (illus. by) Elizabeth Rippey. 249

pp., 46 pis. of colored paintings, text draw-

ings. Macmillan South Africa (in the U.S.:

International scholarly Book Services, P.O.

Box 1632, Beaverton, OR 97075). $49.95.

This is by far the best of the recent guides to

the shelled mollusks of this region. About 600

species are illustrated, described, and are with

comparative and habitudinal remarks. Authors

and dates accompany the scientific names. Ex-

cellent diagnostic drawings of bivalve hinges

make identifications easier. Dr. Kilburn has

added professional taxonomic information

throughout the book. The introductory chapters

have an excellent historical account and a good

coverage of collecting and curatorial methods.

The colored paintings are adequate for iden-

tification purposes. This book joins the ranks of

other fine faunal guides, such as those of Keen,

Kay and Powell. - R. Tucker Abbott,

American Malacologists, Inc.

Seashells of Oman by Donald and Eloise Bosch

(edited by Kathleen Smythe). 206 pp., numer-

ous color photos. Longman Group, Ltd., Lon-

don and New York. $35.00.

This is a beautifully illustrated book on 258

species of gastropods and 96 bivalves found in

the southeastern Arabian Peninsula. Authors,

dates and correct scientific names, together

wath habitudinal information make this a recom-

mended book for this area. - R.T.A.

Seashells of the Arabian Gulf by Kathleen

Smythe. i23 pp., 20 pis. (8 in color). Allen and

Unwin, Inc., Winchester, MA. $25.00.

This small, simple guide covers only a few

species and is poorly illustrated. The price is for-

bidding. -R.T.A.

The Freshwater Molluscs of Canada by Arthur

H. Clarke. 446 pp., 179 pis. (plus 50 in color).

University of Chicago Press, 5801 S. Ellis

Ave., Chicago, IL 60637, $39.95. In Canada:

National Museum of Natural Sciences,

Ottawa, Canada KIA 0M8. $39.95.

At long last Canada has a complete and well-

illustrated handbook for the identification of its

179 species of freshwater mollusks. Dr. Clarke

has considerably expanded the coverage of his

1973 monograph of the 103 species found in the

more northerly Canadian Interior Basin. This

new book is for the layman and biologist not

familiar with mollusks. Excellent photographs,

including SEM views of minute species, ecologi-

cal notes and easily interpreted distributional

maps accompany each species. A two-page

spread is devoted to each species. The colored

paintings of 50 Unionidae mussels are superior

to anything previously published on the subject.

The onerous synonymies and detailed locality

records of a monograph are omitted.

Perhaps a little more attention could have

been given to generic differentiation, particular-

ly in the Lymnaeidae, and perhaps in some

future edition the details of live gastropods and

bivalve larvae will be given. This is an excellent

popular guide, and its price is in keeping with to-

day's publishing costs. - R.T.A.

James Graham Cooper - Pioneer Western Nat-

uralist. By Eugene Coan. 1982. 255 pp., 23

figures, map. A Northwest Naturalist Book,

University Press of Idaho, University Station,

Box 3368, Moscow, Idaho 82843. $11.95.

It is fortunate that natural historians often

have a need to know details concerning early

workers in their fields of endeavor. Dr. Eugene

Coan is one such scientist who, dismayed by the

lack of information on James G. Cooper, an im-

portant contributor to knowledge about our

Vol. 97(1)

January 31, 1983

THE NAUTILUS 45

country's natural history during the 19th Cen-

tury, set about to fill that hiatus by collecting

the data which grew into the volume cited

above. The book was meticulously researched in

museums, libraries, and archives throughout the

country resulting in a much more complete pic-

ture than had previously existed of Cooper's life,

work, and travels from 1830 to 1902. The bio-

graphy is replete with direct quotations from

Cooper's writings and those of others, inter-

spersed with the author's comments which

maintains the continuity. Each chapter is

documented with extensive notes.

Cooper, a physician, is perhaps best known for

his participation in the Pacific Railroad Survey,

1853-1855, but he also took part in a number of

other major and minor expeditions to various

portions of the United States. From an early

age he was fascinated with collecting and study-

ing natural objects, and apparently also inspired

others to collect specimens for him to study. He

lived during what has been called the "Golden

Age" of biological observation when many new

species were being described from world-wide

explorations. He knew and corresponded with

important naturalists of his day such as Baird,

Dall, Carpenter, and others who encouraged his

surveys. Much of Cooper's malacological work

was devoted to Recent land and freshwater

moUusks, although he frequently described

fossils and marine species.

Assembled in the book are extensive lists of

the zoological taxa Cooper described carefully

annotated with interpretive notes, localities,

and citations of repositories for the type

specimens of 3 brachiopods, over 100 mollusks,

1 insect, 13 fish, 2 reptiles, 4 birds, and 1 mam-

mal that he discovered. The bibliography con-

tains all of Cooper's works, those of his father,

William Cooper, a naturalist in his own right,

together with literature cited by Coan in the

text. This book gives a most interesting view of

life over 100 years ago in the United States,

besides providing what is essentially an

obligatory reference for those investigating the

natural history of this country.

Jcweph Rosewater

Division nf Mollusl<s

National Museum of Natural History

Washintrton, D.C. 20.560

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46 THE NAUTILUS January 31, 1983

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THE AMERICAN MALACOLOGICAL BULLETIN

- Volume 1 - 1983 -

Volume 1 of The Arney-irnn Malacolagical Bulletin will be available in

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THE

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Volume 97, number 2 - April 6, 1983

ISSN 0028-1344

CONTENTS

William Miller, III

Distribution of Pyramidellid Gastropods in Late Pleistocene Nearshore Sedimentary

Environments, Dare County, North Carolina 49

Robert F. McMahon and Colette O'Byrne McMahon

Leaping and Swimming as Predator Escape Responses in the Jackknife Clam,

Ensis yyiinor Dal! (Bivalvia: Pharellidae) 55

Branley A. Branson, James B. Sickel and Bruce M. Bauer

Notes on Rare and Endangered or Threateneil Pleurocerid Snails from the

Cumberland River, Kentucky 58

Robert Robertson

Extraordinarily Rapid Postlarval Growth of a Tropical Wentletrap (Ep/^j/r/wm alhiduvi) 60

Paul D. Hartfield and Charles M. Cooper

Distribution of Corbieula fluininea. the Asiatic Clam, in Mississippi 66

Fred G. Thompson

The Planorl)id Snail Micronienetus dilatatus aims (Pilsbry) in the West Indies

and Central America 68

Artie L. Metealf

A New Humboldtiana (Pulmonata: Helminthoglyptidae) from Northwestern

Coahuila, Mexico 69

William L. Pratt

hWmg Promenetus exacuosus (Pulmonata: Planorbidae) from North Central Texas 73

Derek S. Davis

The Freshwater Snail, Menetus dilatatus, (Planorbidae) in Nova Scotia 74

Eva Pip

Variability in North American Lymnaea stagnalis L. (Gastropoda: Lymnaeidae) 77

Frederick C. Hill

Unexplained Occurrence of the Mactrid Bivalve, Rangia cuneata. from the Arrowhead

Farms Indian Site Near Louisville, Kentucky 79

Robert S. Prezant

Auto-drilling in the Oyster Drill Thais Haernastoma (Muricidae) 81

Richard E. Petit

Katherine V. W. Palmer - An Obituary, 1895-1982 85

Recent Death 84

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Vol. 97(2)

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THE NAUTILUS 49

DISTRIBUTION OF PYRAMIDELLID GASTROPODS IN LATE

PLEISTOCENE NEARSHORE SEDIMENTARY ENVIRONMENTS,

DARE COUNTY, NORTH CAROLINA

William Miller, III

Earth Sciences Department

Tulane University

New Orleans, LA 70118

ABSTRACT

Examination offossiliferous sediments at the Stetson borrow pit indicates that

the late Pleistocene, molluscan fauna of Dare County. North Carolina, included

seven species of ectoparasitic pyramidellid snails that were distributed in recur-

rent taxonomic associations within three major nearshore environ.m.ents: 1) baxk-

barrier estuarin£ (characterized by Odostomia impressa (Say) and Turbonilla

powhatani Henderso7i and Bartsch; also containing Odostomia virginica Hender-

son and Bartsch), 2) transitional (with only 0. impressaA and 3) inner shelf

marine (characterized by Turbonilla interrupta (Totten): with rare Sayella

chesapeakea Morrison, Odostomia conoidea acutidens Dall, Odostomia dux Dall

and Bartsch, 0. impressa, arid 0. virginica).

The Pyramidellidae comprises a large, taxo-

nomically confused group of minute opistho-

branch snails that have been studied largely by

conchologists. The shells of the organisms are

commonly found in Cenozoic fossil deposits of

marine and estuarine origin, indicating that

pyramidellids were just as important as com-

ponents of nearshore benthic communities in

the past as they are in modern benthic com-

munities (see Bartsch, 1955; Wells, 1961).

The traditional approach to the study of the

Pyramidellidae by both conchologists and paleo-

conchologists has involved extensive, regional

taxonomic studies of shells that include general

notes on biogeographic and bathymetric distri-

butions (e.g., Bartsch, 1909, 1955; Dall and

Bartsch, 1909). A number of significant studies

of anatomy, feeding habits, and reproductive

behavior has appeared in the past few decades

greatly extending knowledge of the biological

properties of the most common species of

modern Pyramidellidae (e.g., Robertson, 1978;

Scheltema, 1965; Wells, 1959; Allen, 1958; Fret-

ter and Graham, 1949). However, the phyloge-

netic relationships, paleocommunity contexts,

and paleoenvironmental distributions of pyra-

midellid gastropods remain as essentially unex-

plored areas of research in molluscan paleon-

tology.

The purpose of my paper is to describe the

pyramidellid associations (i.e., groups of pyra-

midellid species derived from a particular kind

of community; these are "taxonomic associa-

tions" of Kauffman and Scott, 1976, p. 21)

preserved in upper Pleistocene estuarine and

marine deposits in Dare County, and to recon-

struct the paleoenvironmental distributions of

these associations in order to test pyramidellids

as possible indicators of ancient nearshore en-

vironments. In addition, several species previ-

ously known only from recent material are

described for the first time as fossils.

Because the stratigraphic framework of Late

Cenozoic deposits beneath the Atlantic Coastal

Plain is being slowly unravelled mainly through

the use of various methods of subsurface sam-

pling (see Oaks and DuBar, 1974), it is clearly

advantageous to be able to recognize fossil taxa

that are both small enough to be recovered more

or less undamaged by drilling and that are use-

ful in delimiting, together with stratigraphic

and sedimentologic data, ancient sedimentary

environments. Benthic foraminiferids, ostra-

codes, and fragments of larger moUusks have

50 THE NAUTILUS

April 6, 1983

Vol. 97(2)

been used in this capacity traditionally. I pro-

pose that pyramidellids could be employed as

paleoenvironmental "guides" with considerable

ease because: 1) no special techniques other than

screening sediment are required to isolate speci-

mens, 2) entire shells can be recovered from

auger stems or split-spoon cores, and 3) most

species can be determined (at least tentatively)

using a lOX handlens. It remains to be demon-

strated that pyramidellids have recognizable

distributional patterns, recurrent in space and

time, which allow them to be used in this role.

Locality, Stratigraphy, and Methods

Specimens used in this study were collected

from three-dimensional exposures of back-

barrier deposits in the walls of the Stetson bor-

row pit, and from a single power-auger boring in

marine deposits below the floor of the pit. The

borrow pit is located in northcentral peninsular

Dare County, North Carolina, 15 km west of

Croatan Sound and 10.5 km east of Alligator

River, on the north side of U. S. Highway 64

(Fig. 1).

The Stetson pit is an important Pleistocene

fossil locality in northeastern North Carolina,

because the low, flat, featureless topography of

the Outer Coastal Plain precludes natural sur-

face exposure of the underlying Late Cenozoic

formations, and because in 1978 the borrow pit

was the only artificial exposure of fossiliferous

Pleistocene on the Dare County peninsula. The

walls of the pit contain a 4-meter vertical ex-

posure of shelly, sandy lagoonal sediments over-

lain by about 2 meters of unfossiliferous, muddy

sand of river-estuarine origin. A careful study of

the sedimentary structures, stratigraphic rela-

tionships, and fossil shells in the lagoonal

deposits indicates that these beds were depo-

sited in a patchwork of backbarrier sedimentary

environments including shallow subtidal muddy-

bottom areas that surrounded shallow subtidal

to intertidal sand shoals, oyster banks, tidal

channels, and lagoon-margin bays and beaches

(see Miller, 1978, for a detailed environmental

reconstruction). A power-auger boring in the

floor of the pit revealed an additional 2 meters

of backbarrier muddy sand underlain by about 1

meter of muddy sand containing a mixture of

marine and estuarine shells. This transitional

unit grades downward into a thick sequence (10

meters +) of marine deposits. The marine

sediments show signs of having been deposited

in subtidal, inner shelf sedimentary environ-

ments that probably included lower shoreface,

sand shoal, and intershoal trough areas (Miller,

1978). A generalized stratigraphic column of the

borrow pit is shown in Fig. 2.

Fifteen bulk samples, ranging from 0.5 to 4

liters, were collected from surface exposures

and the bore hole. Over 26,000 fossil shells were

separated from the samples by wet screening,

then were dried and identified to species if possi-

ble. A total of 350 pyramidellids, or 1.3% of the

fossil shells recovered, were later separated

from the collection and re-examined with a bino-

iPf^Pi^'*

FIG. 1. Indux map ^lu>wing lucahuiL uj Stetson borrow pit m

northern mainland Dare County, North Carolina.

FIG. 2. Stratigraphic column of upper Pleistoceyie deposits

at the Stetson borrow pit. (Numbers in brackets indicate

number of samples collected.)

Vol. 97(2)

April 6, 1983

THE NAUTILUS 51

cular microscope to double-check determina-

tions and to note the range of morphologic vari-

ability of each species. A total of seven species

was identified (Table 1; Figs. 3 through 9).

On the Use of Pyramidellid Names

in Paleontology

Anyone beginning a study involving the Pyra-

midellidae is at once faced with the enormous

taxonomic difficulties that characterize this

group. Even the compilation of simple species

lists from a series of samples is hampered by the

many look-alike "species" which appear to have

been named by authors who applied a strictly

typological, rather than a biological species con-

cept in establishing new taxa. Either the myriad

of pyramidellid species is the result of unpre-

cedented oversplitting, or rampant adaptive

radiation in the Late Cenozoic, partly obscured

by morphologic convergence, has visited near-

unmanagable diversity upon the taxonomist of

pyramidellid mollusks. A combination of these

two extreme interpretations is probably nearer

the truth. Yet there may be new hope of un-

scrambling the systematics of this group in the

form of taxonomic studies that utilize soft part

anatomy and reproductive behavior (e.g.,

Robertson, 1978). After this approach has been

more widely employed, many pyramidellid

|i

mm

FIGS. 3-8. Fossil pyramidellid snails from, the Stetson pit:

3. Sayella chesapeakea Morrison. 1939; 4, Odostomia im-

pressa (Say. lS-21); 5. Odostomia conoidea acutidens Dall,

ISSi: 6, Odostomia dux Dall and Bartsch, 1906: 7, Turbonilla

powliatani Henderson and Bartsch. 1911,: 8, Turbonilla inter-

rupta(ro«e«. 1835).

"species" will probably be recognized as morpho-

types and the list of true species will be con-

siderably shortened (Abbott, 1974). (I have

found that it is actually possible to identify a

great deal of polymorphic variation within cer-

tain species using only conchological character-

istics (see Fig. 10). This method can be used by

paleontologists to control the number of species

included in lists that are generated from collec-

TABLE 1. Associations of fossil pyramidellid snails.

Inner shelf

marine (3 samples)

Odostomia impressa

Odostomia dux

Odostomia virginica

Sayella chesapeakea

Turbonilla interrupta

52 THE NAUTILUS

April 6, 1983

Vol. 97(2)

E

E

CM

FIG. 9. Odostomia virginica Henderson and Bartsch, 1914.

impressa trifida bedequensis trifida

E

E

lacking

CORD FUSION

partial

complete

FIG. 10. Variation in spiral ornamentation on whorl swr-

face above the aperture o/ Odostomia impressa (Say). Varia-

tion was obsenied in every sample containing the species.

(From a biological point of view, the variation could be the

expression in shell morphology of an tmderlying develop-

mental polymorphism, possibly expressed functionally in the

soft parts of living 0. impressa f.see Gilbert, 1980). Develop-

mental polyynorphism in certain species of Pyramidellidae

may have resulted in morphotypes that have been regarded

previously as distinct species. For comparison, three of

Bartsch's (1909) species and subspecies categories, applied to

apparent morphotypes of 0. impressa, are shown along the

top of the drawing next to approximately equivalent morphs).

tions of fossil shells and to approximate the

biological species concept when using fossil

material.) Unfortunately for the paleontologist,

spermatophore morphology, location of the

female pore, and reproductive behavior patterns

have no chance of being preserved with the dead

shells of pyramidellids, and the conchological

literature remains as the indispensible source of

taxonomic information for the identification of

fossil specimens. Published descriptions and il-

lustrations of species consulted in this study in-

clude: Abbott, 1974; Morris, 1973; Wells "and

Wells, 1961; Bartsch, 1955, 1909; and Morrison,

1939.

Pyramidellid Associations

Three recurrent taxonomic associations of

pyramidellid snails were found in the Stetson

borrow pit samples: 1) backbarrier estuarine as-

sociations, 2) a transitional association, and 3)

inner shelf marine associations (Table 1). These

associations were components of high-domi-

nance, "physically accomodated" benthic com-

munities in which organism-environment inter-

actions were generally more significant than

biological interactions, and in which community

organization was rather loosely structured

(Johnson, 1972). Although pyramidellid distribu-

tion was no doubt partly controlled by the occur-

rence of preferred hosts and other organisms

that could be easily parasitized, environmental

distribution outlined below was probably also

controlled by ambient salinity ranges (Larsen,

1976; Orth, 1976; Wells, 1961, p. 256). Together,

trophic resource availability and salinity appear

to be the most important parameters controlling

the successful post-larval establishment of local

pyramidellid populations in temperate near-

shore waters.

Communities living in the physico-chemical

mosaic of backbarrier environments contained

three species of pyramidellids, two of which

have not been previously reported as fossils

(Tables 1, 2): Odostomia impressa (Fig. 4),

Odostomia virginica (Fig. 9), and Turbonilla

powhatani (Fig. 7). 0. impressa. the numerically

dominant species, is a biologically well-known

ectoparasite of oysters and sometimes other

mollusks (Robertson, 1978; Wells, 1959; Allen,

1958). It is no coincidence that samples contain-

ing 0. impressa also contained large numbers of

the valves oi Crassostrea virginica (Gmelin) (see

Wells, 1961). 0. virginica is biologically un-

known. T. powhatani occurs in moderate num-

bers in samples and appears to have been re-

stricted to backbarrier deposits. The possibility

that this species is an ecophenotype of Tur-

bonilla interrupta cannot be excluded. Host

preferences of both 0. virginica and T.

powhatani are, to my knowledge, undeter-

mined. Co-occurring with the backbarrier estua-

rine associations are the following species of

mollusks, in addition to oysters: Gemma gemma,

(Totten), Mya arenaria Linne, Ensis directus

Vol. 97(2)

April 6, 1983

THE NAUTILUS 53

TABLE 2. Fossil occurrences of pyrnmidellid species.

Odostomia impressa (Pliocene)

Odostomia concoidea (Miocene)

Turbonilla interrupta (Miocene)

Odostomia dux

Odostomia virginica

Sayella chesapeakea

Turbonilla powhatani

= Clark, 1906; Richards, 1962; Shimer and Shrock. 1965; DuBar, Solliday, and

Howard, 1974; DuBar, Johnson, Thorn, and Hatchell, 1974; Bailey, 1977.

Conrad, Petricola pholadifonnis (Lamarck),

Mulinia lateralis (Say), Macoma balthica

(Linne), Anadara transversa (Say), Mitrella

lunata (Say), Nassarius ohsoletus (Say), Crepi-

dula convexa Say, and Lunatia heros (Say).

In deposits beneath the backbarrier beds at

the Stetson pit, a mixed assemblage of estuarine

and marine mollusks also contains abundant

shells of 0. impressa. These transitional

deposits record the partitioning of a coastal

compartment in the Dare County area in late

Pleistocene time, possibly by an accreting bar-

rier shoal or island, and the local initiation of

estuarine conditions (see Miller, 1978). The

eurytopic nature of 0. impressa is shown by the

occurrence of this species in a sedimentary en-

vironment that apparently was too rigorous for

other pyramidellids. It even has a limited occur-

rence in the subjacent marine deposits (Table 1).

Could polymorphism in 0. impressa be related

to its apparent eurytopic character (see Fig. 10)?

The most species-rich associations in samples

came from marine beds below the transitional

deposits, and contained the following species: 0.

impressa, 0. virginica, Odostomia dux (Fig. 6),

Odostomia conoidea acutidens (Fig. 5), Sayella

chesapeakea (Fig. 3), and Turbonilla interrupta-

(Fig. 8). 0. dux and S. chesapeakea are described

for the first time as fossils (Table 2). In this asso-

ciation, T. interrupta is by far the most abun-

dant species (Table 1). It is a well-known, shal-

low water species that parasitizes a variety of

bivalve mollusks (Morton, 1967, p. 185; Abbott,

1974). The presence of rare 0. impressa and 0.

virginica indicates that small populations of

these species apparently were able to live in

marine areas as well as adjacent estuarine envi-

ronments. 0. c. acutidens is unknown biologi-

cally. 0. duj: may have fed upon tubicolous

polychaetes, such as Sabellaria, and probably

preferred near-normal marine salinities (Wells

and Wells, 1961). S. chesapeakea may be another

eurytopic pyramidellid that occasionally ranged

into open marine waters from its normal mud-

FIG. 11. Schematic representation of the distribution of

pyramidellid associations in nearshore environments:

1-ba^kbarrier estuarine associations: 2-transitional associa-

tion: 3-inner shelf marine associations.

54 THE NAUTILUS

April 6, 1983

Vol. 97(2)

flat habitat (see Morrison, 1939). Co-occurring

with the inner shelf marine associations were

the following mollusk species: Mulinia lateralis,

Pleuromeris tridentata (Say), Crassinella lunu-

lata (Conrad), Mercenaria mercenaria (Linne),

Donax variabilis Say, Spisula solidissima (Dill-

wyn), Retiisa canaliculata (Say), Terehr-a dislo-

cata (Say), Polinices duplicatus (Say), Mitrella

lunata, Nassariiis acutus (Say), and Olivella

mutica (Say).

The distribution of pyramidellid associations

in late Pleistocene nearshore environments of

the northern Dare County area is summarized in

Fig. 11.

Conclusions

The stratigraphy and paleontology of the Stet-

son borrow pit section show that the late Pleis-

tocene history of the Dare County area included

an important change in the configuration of

coastal environments involving a shift from

marine to estuarine conditions. This replace-

ment of environments is reflected in the vertical

succession of pryamidellid associations at the

borrow pit from a species-rich, inner shelf asso-

ciation, to a transitional zone containing a single

species, and finally to a moderately species-rich

backbarrier association. The change is also

reflected in a shift through time in the relative

abundance and rank of eurytopic species (e.g.,

0. im.'pressa), and in the appearance and dis-

appearance of apparently stenotopic species,

such as T. powhatani (an estuarine species) and

0. dux (a marine species) in the stratigraphic

column.

In conclusion, pyramidellid gastropods appear

to be useful as indicators of nearshore sedimen-

tary environments when considered in their

paleosynecological and stratigraphic contexts.

Although taxonomic complexities have made

the Pyramidellidae a less than popular group of

mollusks with paleontologists, I believe this

family of gastropods holds great promise as a

source of raw material for studies of polymor-

phism and its role in macroevolution, in paleo-

ecological studies of deployment and function of

morphotypes of variable species in ancient ben-

thic communities, and in evolutionary studies of

the adaptive radiation of parasites that have a

good fossil record.

Acknowledgments

I extend my thanks to Dr. Hubert C. Skinner

(Tulane University) for suggesting manuscript

improvements and for his general encourage-

ment. Duke University and the North Carolina

Department of Natural and Economic Re-

sources provided grant support that made the

field work possible.

LITERATURE CITED

Abbott, R. T. 1974. Amertcaji Seashells (2nd edition). Van

Nostrand Reinhold, New York, 663 pp.

Allen, J. F. 1958. Feeding habits of two species of Odos-

tomia. The Nautilus 72:11-15.

Bailey, R. H. 1977. Neogene molluscan assemblages along

the Chowan River. North Carolina. Southeastern Geol.

18:173-189.

Bartsch, P. 1909. Pyramidellidae of New England and the

adjacent region. Pror. Boston Soc. Nat. His., 34(4):67-113.

1955. The pyramidellid mollusks of the Pliocene

deposits of North St. Petersburg, Florida. Smithson.

Misc. Collect. 125(2); 102 pp.

Clark, W. B. 1906. Mollusca. In. Shattuck, G. B. The Plio-

cene and Pleistocene Deposits of Maryland. Maryland

Geol. Survey, Baltimore, p. 176-210.

Dall, W. H. and P. Bartsch. 1909. Monograph of west

American pyramidellid mollusks. U. S. National Mus.

Bull. 68, 258 pp.

DuBar, J. R., J. R. Solliday and J. F. Howard. 1974. Strati-

graphy and morphology of Neogene deposits, Neuse

River Estuary, North Carolina. In. Oaks. R. Q. and J. R.

DuBar (eds.), Post-Miocene Stratigraphy: Central and

Southern Atlantic Coastal Plain. Utah State Univ. Press,

Logan, p. 102-122.

DuBar, .1. R., H. S. Johnson, B. Thorn and W. 0. Hatchell.

1974. Neogene stratigraphy and morphology, south flank

of the Cape Fear Arch, North and South Carolina. In,

Oaks, R. Q. and J. R. DuBar (eds.), Post-Miocene Strati-

graphy: Central and Southern Atlantic Coastal Plain.

Utah State Univ. Press. Logan, p. 139-173.

Fretter, V. and A. Graham. 1949. The structure and mode

of life of the Pyramidellidae, parasitic opisthobranchs.

Jour. Marine Biol. Assoc. U. K. 28:493-532.

Gilbert, J. J. 1980. Developmental polymorphism in the

rotifer Asplanchna sieboldi. Am.. Sci. 68:636-646.

Johnson, R. G. 1972. Conceptual models of benthic marine

communities. In, T. J. M. Schopf (ed.), Models in Paleo-

biology. Freeman, Cooper and Co., San Francisco,

p. 148-159.

Kauffman, E. G. and R. W. Scott. 1976. Basic concepts of

community ecology and paleoecology. In, R. W. Scott and

R. R. West (eds.), Structure and Classification of Paleo-

Vol. 97(2)

April 6, 1983

THE NAUTILUS 55

communities. Dowden, Hutchinson and Ross. Strouds-

burg, Penn., p. 1-28.

Larsen, P. F. 1976. Patterns of distribution of estuarine

organisms and their response to a catastrophic decrease

in salinity. In. The Chesapeake Research Consortium,

Inc., The Ejjects of Tropical Storm Agnes on the Chesa-

peake Bay Estuarine System. Johns Hopkins Univ. Press,

Baltimore, p. 55.5-565.

Miller, W., HI. 1978. Ecological units, paleocommunity

structure, and ecological history of late Pleistocene de-

posits in Dare County, North Carolina. Unpubl. M. S.

thesis, Duke Univ.. 135 pp.

Morris, P. A. 1973. A Field Guide to Shells. Houghton

Mifflin, Boston, 330 pp.

Morrison, J. P. E. 1939. Two new species of Sayella with

notes on the genus. The Nautilus 53:43-45.

Morton, J. E. 1967. Molluscs (4th edition). Hutchinson,

London, 244 pp.

Oaks, R. Q. and J. R. DuBar. 1974. Introduction. In. R. Q.

Oaks and J. R. DuBar (eds.), Post-Miocene Stratigraphy:

Central and Southern Atlantic Coastal Plain. Utah State

Univ. Press, Logan, p. 2-8.

Orth, R. J. 1976. The effect of tropical storm Agnes on the

benthic fauna of eelgrass, Zostera marina, in the lower

Chesapeake Bay. In. The Chesapeake Research Consor-

tium, Inc., The EJfects of Tropical Storm Agnes on the

Chesapeake Bay Estuarine System. Johns Hopkins Univ.

Press, Baltimore, p. 566-583.

Richards, H. G. 1962. Studies on the marine Pleistocene.

Part II. The marine Pleistocene mollusks of eastern

North America. Trans. Am. Philosoph. Soc, New Series-

52(3):42-141.

Robertson, R. 1978. Spermatophores of si.\ eastern North

American pyramidellid gastropods and their systematic

significance (with the new genus Boonea). Biol. Biill.

155:360-382.

Scheltema, A. H. 1965. Two gastropod hosts of the pyra-

midellid gastropod Odostomia bisuturalis. The Nautilus

79:7-10.

Shinier, H. W. and R. R. Shrock. 1965. Index Fossils of

North America (8th edition). Mass. Instit. Tech. Press,

Cambridge, 837 pp.

Wells, H. W. 1959. Notes on Odostomia impressa (Say).

The Nautilus 72:140-144.

1961. The fauna of oyster beds, with special

reference to the salinity factor. Ecol. Mun. 31:239-266.

Wells, H. W. and M. J. Wells. 1961. Three species of Odosto-

mia from North Carolina, with description of new species.

The Nautilus 74:149-157.

LEAPING AND SWIMMING AS PREDATOR ESCAPE RESPONSES IN THE

JACKKNIFE CLAM, ENSIS MINOR DALL (BIVALVIA: PHARELLIDAE)

Robert F. McMahon and Colette O'Byrne McMahon

Department of Biology, Box 19498

The University of Texas at Arlington

Arlington, Texas 76019

ABSTRACT

Two distinct behaviors, "leaping" and "swimming", were observed to follow dis-

lodgement from the substratum in the Western Atlantic jackknife clam. Ensis

minor Dall, from, the Laguna Madre, Texas. Leaping involves extension of the tip

of the foot into the substratum followed by an anteriorly directed thrust of the foot

fram the pedal gape. During swimming the foot remains flax- id and maximally ex-

tended from the pedal gape while rapidly repeated shell valve adduction draws

water into the mantle cavity ventrally and forces it as a jet of water from the

anterior pedal gape propelling the clam posteriorly. Both behaviors are im-

mediately succeeded by burrowing arid are presumed to be highly adaptive

predator escape responses in this species.

Swimming behavior is best known and de-

scribed in the superfamily Pectinacea whose

members swim perpendicularly to the hinge axis

with the commissure plane approximately 45° of

horizontal and in the family Limidae which swim

with the commissure plane oriented vertically

56 THE NAUTILUS

April 6, 1983

Vol. 97(2)

(Morton, 1964; Stanley, 1970). Also reported to

swim occassionally are some species of the

families Solemyacidae (Solemya velum Say)

(Morse, 1913), and the Cardiidae {Laevicardiurn

laevigatum (Linne)) (Stanley, 1970). In the

superfamily Solenacea (Razor-shells) swimming

has been observed in the families Solenidae

{Solen marginatiLS Pulteney) (Poli, 1791-1795,

Deshayes 1844-1848) and Pharellidae (Ensis

directus Conrad) (Drew, 1906; Stanley, 1970).

This paper discusses both "leaping" and "swim-

ming" behavior as predator escape responses in

a second species of the family Pharellidae, the

jackknife clam, Ensis minor Dall, that were

observed in individuals collected on March 21,

1981, in 5-15 cm of water on a sandy shore of

low slope on Bird Island Beach, Padre Island

National Seashore, on the Laguna Madre,

Texas.

In this area the E. minor population is very

dense (300-500 individuals/m^). Individuals

were dislodged from their burrows by gently

fanning water over the sediment surface to re-

move the sand covering them. Ensis minor with

a maximum shell length of 76 mm (Andrews,

1977) is a much smaller bivalve than the West-

ern Atlantic species, Ensis directus, which can

reach a length of 20 cm (Stanley, 1970; Abbott,

1974). Specimens of E. minor collected at Bird

Island Beach had a mean shell length of 28.9 mm

(s.d.= ±2.66, s.e.= ±0.30, range = 22.3-42.4,

n = 79) and a mean shell height of 4.1 mm

(s.d.= ±0.36, s.e.= ±0.041, range = 3.4-5.9,

n = 79).

Many specimens of E. minor dislodged from

the substratum showed one or the other of two

distinct escape behaviors. The first involved a

"leaping" response in which individuals moved

rapidly across the sand surface before initiating

burrowing behavior. The second involved a dis-

tinct "swimming" response in which individuals

left the sand surface and moved both vertically

and horizontally through the water column

before settling and re-entering the substratum.

Both leaping and swimming were carefully

observed in the field and in specimens removed

from the field into 3.8 liter glass vessels.

Leaping in E. minor is initiated by an individ-

ual lying on its right or left valve by extending

the tip of the foot into the substratum followed

by an anteriorly directed thrust of the foot, ex-

tending for a distance nearly that of the shell

length, from the anterior pedal gape which

forces the clam to move posteriorly across the

sediment surface. Up to five to ten consecutive

foot thrusts may occur within a few seconds,

moving the clam distances of up to 10-15 cm

from the site of dislodgement. Leaping behavior

is then immediately followed by burrowing into

the substratum as described for Ensis by

Trueman (1967).

Leaping behavior has also been described for

the large Western Atlantic species E. directum

but instead of anteriorly directed thrusts as oc-

curs with E. minor it involves extending the

foot and bending it dorsally beneath the shell,

followed by a sudden straightening of the foot to

the anterior projecting the clam posteriorly.

These leaping movements are repeated several

times (Drew, 1906). Less commonly, E. directus

may leap as does E. minor by repeated thrusts

of the foot anteriorly (Drew, 1906).

Specimens of E. minor also displayed a unique

swimming behavior when dislodged from the

sediment which involved entering the water col-

umn for extended periods of time and traveling

over relatively large horizontal distances. Swim-

ming was accomplished by extending the foot to

its maximal length (nearly equivalent to shell

length) from the anterior pedal gape. In this ex-

tended condition the foot is somewhat flacid and

its diameter appears to be less than that of the

pedal gape at the anterior margins of the shell.

Foot extension greatly increases the volume of

water in the pallial cavity and apparently allows

water to be drawn diffusely into the cavity be-

tween the unfused portion of the ventral mantle

edges as the shell valves open. Thereafter, the

valves are rapidly adducted. During adduction

the free mantle edges appear to seal the ventral

margin of the pallial cavity, forcing a jet of

water to be expelled from the relatively con-

stricted anterior padal gape around the ex-

tended foot, driving the individual quickly

through the water in a posterior direction with

the extended foot trailing behind. During swim-

ming, valve adduction occurs at least several

times per second propelling the individual rapid-

Vol. 97(2)

April 6, 1983

THE NAUTILUS 57

ly from the sediment surface and away from the

site of dislodgement. This swimming behavior

was maintained in some individuals for at least

15 to 20 seconds, permitting horizontal move-

ments of well over a meter from the dislodge-

ment site. On cessation of swimming, individ-

uals initiate burrowing immediately on settling

to the sediment surface.

Swimming has been previously described for

adult specimens of E. diredus in which it is

associated with a repeated rapid extension of

the foot from the anterior pedal opening, the

shell valves being adducted each time the foot is

retracted into the pallial cavity. Simultaneous

foot retraction and shell adduction forces a jet of

water out the anterior pedal opening propelling

the individual posteriorly across the sediment

surface without entry into the water column

(Drew, 1906). This description is very similar to

that reported for the related but smaller species

Solen marginatus (Deshayes, 1844-1848). This

swimming behavior of these two larger species,

E. directus and S. marginatus, is quite different

from that of the smaller E. minor, (which is ac-

complished by a rapid valve adduction alone)

and appears to be a variant of "leaping" activity

in which movement is accomplished by a succes-

sion of rapid anteriorly directed extensions of

the foot, a behavior not much modified from

that associated with burrowing. In contrast,

swimming behavior in E. minor seems to be

distinctly different from that associated with

either leaping or burrowing.

Stanley (1970) also described swimming in E.

directus as "an anterior swimming movement

accomplished as in Solemya by retracting the ex-

tended, plug-like foot and adducting the valves

to expel a jet of water posteriorly". This sort of

swimming behavior was never observed in E.

minor and in light of Drew's (1906) report of

posteriorly directed swimming in E. directus

and of the anatomy of the genus, Ensis

(Trueman, 1967), it is probable that such

"anterior swimming movement" does not occur

in this genus.

Drew (1906) speculated that leaping and

swimming behavior in E. directus was not pri-

marily for escape, as this large bivalve is one of

the fastest and strongest of burrowing species

(Trueman, 1967; Stanley, 1970), apparently

much better able to avoid predators by burrow-

ing deeply. Instead, swimming in E. directus

was hypothesized to be a method for rapidly

changing position after settlement while leaping

was presumed to allow dislodgement of individ-

uals from highly confining or otherwise unsuit-

able substrata and possibly for escape if a

specimen was somehow dislodged from the sub-

stratum (Drew, 1906).

Ensis minor (SL<76 mm) is of a much smaller

adult size than either E. directus or S. margina-

tus. Because of its short siphons individuals

must lie close to the sediment surface where

they could be easily dislodged by predators as

brachyuran decapods and shore birds that probe

the substratum for their prey. Once dislodged,

specimens of E. minor with their thin fragile

shells and open pedal and siphonal gapes would

be readily subject to predation. Therefore, leap-'

ing and swimming behavior after dislodgement

would be of the greatest adaptive advantage to

E. minor allowing dislodged specimens to move

rapidly away from the point of attack before re-

entering the substratum. If burrowing behavior

was initiated immediately after dislodgement a

predator could easily relocate and attack a dis-

lodged individual within the 20-100 seconds it

requires for Ensis to completely re-enter the

substratum (Drew, 1906; Trueman, 1967; Stan-

ley, 1970). However, if burrowing is preceeded

by leaping or particularly by swimming away

from the point of original dislodgement an in-

dividual may gain enough time to completely re-

enter the substratum before it is once again

detected by the predator. It now seems probable

that small species or juvenile specimens of Ensis

are capable of a much more efficient and pro-

longed swimming behavior than has previously

been suspected (perhaps being generally of the

type described herein specifically for E. minor)

as small specimens of E. directus has been taken

in tow nets at the sea surface (Drew, 1906). Such

efficient swimming behavior may be lost in the

adults of large species as E. directum and 5.

marginatus that primarily depend on efficient

burrowing to avoid predators but appears to be

retained in adults of small species as E. minor

which are far more susceptible to dislodgement

April 6, 1983

Vol. 97(2)

from the substratum by predators than are

larger species.

Acknowledgments

We wish to express our gratitude to the Uni-

versity of Texas Marine Science Institute, Port

Aransas Marine Laboratory, Port Aransas,

Texas, 78373 for providing its laboratory and

collection facilities to us during the course of

this study, and to the editor of The Nautilus for

literature and nomenclatorial assistance.

LITERATURE CITED

Abbott, R. T. 1974. Anwrica-n Seashells, Second edition.

Van Nostrand Reinhold Co., New York. 663 pp.

Andrews, J. 1977. Shells and Shores of Texas. University of

Texas Press, Austin, Texas. 365 pp.

Deshayes, M. G. P. 1844-1848. Histuire Naturelle des

Mollusques (Exploration Scientifque de I'Algerie). Paris.

(As cited by Forbes, E.. and S. Hanley. 1853. Pp. 245 in

A History of British Mollusca and thtir Shells, Vol. I.

John Van Voorst, London, 486 pp.).

Drew, G. A. 1906. The habits and movements of the razor

shell clam, Ensis directus, Con. Biol. Bull. 12:127-138.

Morse, E. S. 1913. Observations on Wving Solemnnya (velum

and borealis). Biol. Bull. 25:261-281.

Morton, J. E. 1964. Locomotion. In; Physiology of Mollusca,

Vol. I., K. M. Wilbur and C. M. Yonge, eds. Academic

Press, New York, pp. 383-423.

Poli. 1791-1795. Testacea Utriusque Siciliae. (As cited by

Tryon, G. W., Jr. 1884. Pp. 130 in Structural and Sys-

tematic Conchology. Vol. III. Academy of Natural Sciences

of Philadelphia, Philadelphia, 453 pp.).

Stanley, S. M. 1970. Relation of Shell Form to Life Habits

of the Bivalvia (Mollusca). Memoir 125, The Geological

Society of America, Inc., Boulder, Colorado. 296 pp.

Trueman. E. R. 1967. The dynamics of burrowing in Ensis

(Bivalvia). Proc. Roy. Soc. Lond. 166:459-476.

NOTES ON RARE AND ENDANGERED OR THREATENED PLEUROCERID

SNAILS FROM THE CUMBERLAND RIVER, KENTUCKY

Branlev A. Branson, James B. Sickel' and Bruce M. Bauer^

ABSTRACT

Data for Lithasia armigera, L. geniculata and Pleurocera alveare, three species

listed as Rare and Endangered or of Special Concern in Kentucky and elsewhere,

from the Cumberland River are presented. Ancillary records for Helisoma

anceps, Physa Integra, and Ferrissia rivularis are included.

Recently, a task force (Branson et al. 1981)

was assigned the job of generating a list of rare

and endangered plants and animals of Ken-

tucky. Included in that list are 65 molluscan

species, 49 unionid clams and 16 snails, 12 of

which are aquatic. During the writing process, it

became obvious that we were hampered by a

paucity of published information on the aquatic

gastropod fauna or by the lack of recent surveys

'Department of Biology, Murray State University, Murray.

Kentucky 42071

'Soil Systems, Inc., 525 Webb Industrial Drive, Marietta,

Georgia 30062

of the fauna. This stimulated a tlurry of activi-

ties (Branson and Batch 1981, 1982a, b, c) in at-

tempt to fill in some of the voids in distributional

knowledge and to ascertain the status of the

species considered rare and endangered.

Thus, the first co-author has been conducting

some extensive surveys and biological investiga-

tions of the unionid clams of western Kentucky,

particularly in the Cumberland River drainage

above and below the Lake Barkley Dam. Coinci-

dentally, he and his assistants made collections

of some important pleurocerid snails at various

sites below the dam, one of the few stretches of

the Lower Cumberland River still flowing free-

Vol. 97(2)

April 6, 1983

THE NAUTILUS 59

ly. Because this section of that river was former-

ly rich in pleurocerid snails it was feared that

the giant impoundment and the cold water

released below the dam had either extirpated or

greatly decimated the operculates. However,

this shallow-water reservoir has little tempera-

ture stratification, so temperature problems

have not developed. The daily water tluctuations

and consequent water-quality changes are of

more importance with respect to the molluscan

populations.

One of those species is Lithasia armigera

(Say) 1821. This species is listed as Rare and En-

dangered in Kentucky (Branson et al. 1981) and

elsewhere (Federal Register 1980 -candidate

for listing). The species is truly Rare and En-

dangered throughout much of its range, particu-

larly in the Upper Cumberland River (Branson

and Batch 1982c) where it is heavily impacted by

strip-mining pollutants. However, from the re-

sults of Sickel's observations in the Cumberland

River below the dam there appears to be a thriv-

ing and healthy population in that stretch of the

river, i.e., from the dam to the mouth.

Sickel's collecting sites were: Cumberland

River, River Mile 16, at Pinkneyville, Livings-

ton County, Kentucky, 8 November 1981, 31

specimens (EKU 11939); River Mile 17.3, Crit-

tenden County, 20 September 1981, 30 speci-

mens (EKU 11942); River Mile 26.9, Lyon Coun-

ty, 7 October 1981, 40 specimens (EKU 11941);

River Mile 27.1, Lyon County, 8 October 1981,

21 specimens (EKU 11940). In addition, Bauer

secured 11 specimens (EKU 11937) from a

massive population in the Cumberland River at

Dycusburg on the Livingston-Crittenden county

line. River Mile 20, 5 October 1981. All im'-

mature shells (12 mm or smaller) are banded,

and 65% of the adult shells bear 1 to 5 bands

within the aperture.

The presence of an abundance of young shells

indicates that the population below Barkley

Dam is self-sustaining. Notwithstanding any ad-

ditional environmental changes in the lower

stretches of the Cumberland River, these sites

may be considered as a refugium for this and the

next species below.

Lithasia geniculata Haldeman 1840 is listed

as Endangered in Kentucky (Branson et al.

1981) and is currently being considered for

federal listing (Federal Register 1980). It is tru-

ly rare throughout its range, thus Sickel's

discovery of a small population at Cumberland

River Mile 17.3, Crittenden County, Kentucky,

20 September 1981, is of considerable interest.

He secured three specimens only, all banded

(EKU 11943). No immature shells were col-

lected, and we have no indication of the popula-

tion's size, although some estimates may be

forthcoming after SCUBA-gear observations.

Pleurocera alveare (Conrad) 1834 is con-

sidered as of Special Concern in Kentucky

(Branson et al. 1981) since Cumberland River

specimens have not been reported in many

years. Goodrich (1934) found the species abun-

dantly in the Cumberland River above Burnside,

a site now inundated by the impounded waters

of Lake Cumberland, and recent collecting

(Branson and Batch 1982c) in stretches of the

river above the lake and elsewhere failed to

disclose specimens. Thus, Bauer's discovery of a

good-sized population in the Poor Fork of the

Cumberland at Gatum, Harlan County, Ken-

tucky, 30 September 1981 (12 specimens: EKU

11938), is noteworthy. He also made collections

from the Martins Fork below Martins Fork

Dam, Harlan County, without securing speci-

mens of this species, although he did find a

thriving community of Helisoyna anceps (Menke)

1830 (EKU 11935) and Physa integra Haldeman

1841 (EKU 11936). The environmental condi-

tions below the dam are vastly changed, in-

cluding the presence of silt and lowered

temperatures.

Ferrissia rivularis (Say) 1817. Because of the

very few published records from Kentucky

waters, the single specimen removed from a

Lithasia shell taken from Cumberland River

Mile 17.1 by Sickel is herewith reported (EKU

11944).

LITERATURE CITED

Branson, Branley A. and Donald L. Batch. 1982a. The gas-

tropoda and sphaeriacean clams of Red River, Kentucky.

The Veliger 24:200-204.

(1982b). Distributional records for gastropods

and sphaeriacean clams of the Kentucky and Licking

rivers and Tygarts Creek drainages, Kentucky. Brim-

leyana 7:137-144.

60 THE NAUTILUS

April 6, 1983

Vol. 97(2)

1982c. Molluscan distribution records from the

Cumberiand River, Kentucky. The Veliger 24(4):351-354.

1981. The gastropods and sphaeriacean clams of

the Dix River system, Kentucky, Trans. Ky. Acad. Sri.

42:54-61.

Branson, Branley A., Donald F. Harker, .Jr., .Jerry M. Bas-

kin. Max E. Medley, Donald L. Batch, Melvin L. Warren,

Jr., Wayne H. Davis, Wayne C. Houtcooper, Burt Monroe,

Jr., Loy R. Phillippe and Paul Cupp. 1981. Endangered,

Threatened, and Rare animals and plants of Kentucky.

Trans Ky. Acad. Sci. 42:77-89.

Federal Register. 1980. Part IV. Department of the Interior

Fish and Wildlife Service. Republication of lists of endan-

gered and threatened species and corrections of technical

errors in final rules. 45(99):33768-33779.

Goodrich, C. 1934. Studies of the gastropod family Pleuro-

ceridae-III. Occ. Pap. Mus. Zool. Univ. Mich. 300:1-59.

EXTRAORDINARILY RAPID POSTLARVAL GROWTH OF A

TROPICAL WENTLETRAP (EPITONIUM ALBIDUM)

Robert Robertson

Academy of Natural Sciences of Philadelphia

Nineteenth and the Parkway

Philadelphia, PA 19103

ABSTRACT

Observations and experiments were conducted on the West Indian wentletrap

Epitonium albidum (Orbigny, 18^2) to see how it grows, how fast it grows, and to

determine whether one varix is grown per 2U hours. Combining aquarium with

field data, and data on both sexes (epitoniids are protandric). growth data were

obtained from 23 specimens. The best data set comes from, 8 Barbados males that

greivfor longer than 5 days, in which the mean daily teleoconch length increment

was 0.15 + O.OJt mm, and the mean number of ribs added per day was 1.2 + 0.5.

Three females grew 0.33 to 0.50 mmJday in teleoconch length. The data show that

E. albidum can grow extremely fast, but that there is no consistency about 1.0 rib

being grown per day. Even thick-shelled tropical wentletraps may be among the

fastest-growing gastropods.

Observations are reported on the way Epitonium teleoconchs grow, and an. ex-

planation is given on why the shells are rarely seen with the outer lip between

varices. Further observations show that hunger can cause Epitonium ribs to

become abnorm.ally closely spaced, yet rib-spacings (counts) are considered to be

major taxonomic characters in epitoniids.

According to Bosch (1965), the Hawaiian wen-

tletrap Epitonium ulu Pilsbry, 1921, begins to

lay egg masses within the remarkably short time

of three weeks after the planktotrophic veliger

settles and metamorphoses near or on its scler-

actinian coral host Fungia scutaria Lamarck,

1801 (Robertson, 1970: 45-46). According to

Guinther (1970) "sexual maturity" can even be

attained in "perhaps [a] little more than two

weeks," but he may have been referring to male

maturity (epitoniids are probably all protandric:

Robertson, 1981b). Taylor (1977:258, Fig. 7) has

published data on the early postlarval growth

rate of E. ulu (she did not try to count varices):

when with the actiniarian sea anemone Aiptosta

sp., not its normal host, shell length (of one

specimen only?) increased from 0.6 mm to 3.5

mm in 15 days (0.19 mm/day). "Field growth ex-

periments" conducted by Guinther (1970) on "E.

ulu indicate that [the] growth rate is greatest as

the juvenile snail approaches sexual maturity,

after which [the] growth rate decreases nearly

geometrically." Bosch (1965) reported a max-

imum shell length in E. ulu of 16.5 mm.

Vol. 97(2)

April 6, 1983

THE NAUTILUS 6i

Epitonium ulu is an unusual wentletrap in

having a fairly thin shell with only traces of

varices. The Hawaiian holotype (Acad. Nat. Sci.

Philadelphia [ANSP] no. 127818) is 14.2 mm

long and has a total of about 135 faint varices on

the 9.7 remaining whorls of its teleoconch (about

21 varices on the last whorl). These shell

features and the fast early postlarval growth

rate caused me to wonder whether wentletrap

varices are grown one per 24 hours (possibly

each night).

Such an occurrence is not without precedent

among gastropods: Berry (1962, 1963) reported

that Malayan Opisthostoma spp. and Diplom-

matina spp. (operculate land snails of the family

Diplommatinidae) grow one interspace and one

thin axial rib per day under optimal conditions

(ribs are grown somewhat more rapidly in con-

tinuous darkness, and irregularly when the

snails are too dry). According to Berry (1963),

the same things are true of Malayan Oophana

sp. and Sinoennea sp. (Pulmonata: Streptaxi-

dae), even though they are not closely related to

diplommatinids. Tillier (1981:182) stated that in

diplommatinids sexual maturity coincides with

definitive peristome growth (i.e. growth is

determinate). Tillier suggested that the New

Caledonian diplommatinid Palaina mareana

Tillier, 1981, begins to grow its peristome and

matures in as few as 80 days after hatching, i.e.

after about 80 thin axial ribs are grown.

Epitoniids do not cease growing after becom-

ing female (i.e. their growth is indeterminate),

but if one varix is grown per day the implication

would be that large species such as Epitonium

scalare (Linn., 1758) (up to about 63 mm long)

with big, widely spaced varices (8-15 on the last

whorl) are short-lived, the postlarvae growing

for only about 78 days (count of the total number

of varices on the largest nearly intact specimen

at ANSP), and that E. millecostatum (Pease,

1860-1861), a much smaller species (up to 9.7

mm long) with many very closely-spaced varices

(Robertson, 1981a: 4, Fig. 2) -as many as about

100 on the last whorl only -is relatively long-

lived.

Ankel (1936:149) published a statement bear-

ing on the growth and life cycle of epitoniids: he

claimed that the sex of the European species

Epitonium. dathr'us (Linn., 1758) [as Scala

clathrus] changes with each breeding season.

Multiple sex change seems unlikely (Robertson,

1981b), and Ankel (1936) published no data to

show that there is a discrete breeding season. In

his paper on the longevity of mollusks, Comfort

(1957) had no information about epitoniids.

The main objects of this study were to obtain

some growth rate data on a shallow water,

tropical Epitonium of average size and shell

thickness, and with varices of average size and

spacing, and to ascertain whether one varix is

grown per 24 hours.

Materials and Methods

The species studied was Epitonium albidum

(Orbigny, 1842) (Figs. 1-3), which in the Bahama

Islands (Robertson, 1963), at Virgin Gorda

(British Virgin Islands) and at Barbados lives

with and feeds on the shallowly subtidal ac-

tiniarian sea anemone Stoichactis' helianthu^

(Ellis, 1768). See Robertson (1983).

Epitonium albidum. attains a shell length of

2b + mm (ANSP no. 299697, a sub- fossil shell

from Nassau, New Providence, Bahama Islands;

R. Robertson, leg. 1954). The second largest

specimen collected by me (the largest live-col-

lected) is 15.9 mm long (Figs. 1-3: Virgin Gorda).

The largest Barbados specimen (growth com-

pleted in aquarium) was 14 mm long. The larg-

est Barbados specimen that I collected in 1982

(growth also completed in aquarium) was only

12.0 mm long. Males turn into females between

lengths of 5 and 8 mm (Robertson, 1981b).

Specimens were studied from January to

March 1972 at Virgin Gorda, and from March to

April 1980 and from January to February 1982

at the Bellairs Research Institute of McGill

University, St. James, Barbados. Specimens

were breeding at these times, but whether they

breed year-round is unknown.

Soon after capture of a specimen, the flared or

thickened outer lip was marked with a graphite

pencil. If the outer lip was broken, the two pre-

ceding varices were marked instead. In this way

a replacement rib could be distinguished from a

'Since going to press, I have learned that the genus should

be Stichodactyla (see Dunn, 1981:78-82).

62 THE NAUTILUS

April 6, 1983

Vol. 97(2)

FIGS. 1-3. Shell o/Epitonium albidum 15.9 mm long and 6.6

mm wide (th£ largest specimen collected during this study).

Virgin Gorda, British Virgin Islands. Animal in the female

phase. Growth can be extremely fast: varices are usually

grown faster than 1.0 per day. 1, Apical view. 2, Apertural

view. 3, Basal view.

completely new rib. Once marked, specimens

were 1) kept away from Stoichactis in plastic

petri dishes about half full of sea water changed

daily, or 2) were put with Stoichactis in an

aquarium with running sea water (temperature

about 26° C), or 3) were returned to a Stoichac-

tis in the field and retrieved (if possible) about

two weeks later. Both in aquaria and in the field,

the wentletraps tend to wander from one ane-

mone to another. Thus in the field individuals

are difficult to retrieve.

Stoichactis remained apparently healthy for

up to 12 days in the aquaria, although pedal disc

movement and detachment were frequent. The

fast-growing 7.6-11.5 mm-long female listed in

Table 1 was seen twice (three days apart) to feed

on single Stoichactis tentacles (it may have fed

more frequently than this).

On February 4, 1972, near the southern end of

Copper Mine Bay, on the southeastern coast of

Virgin Gorda, a small group of marked wentle-

traps was released next to an easily recognized

anemone on a near vertical rock face (not one of

the anemones with which the individuals were

originally found). One of these marked animals

was retrieved from the same anemone on

February 19, 1972, 14.8 days later.

A more extensive mark and recapture experi-

ment was conducted at the Bellairs Research In-

stitute, just north of Holetown, western Bar-

bados, off which there is an enormous Stoichac-

tis helianthus colony. On January 28, 1982, all

12 wentletraps from one anemone, and all 39

from another, were marked, measured, and

replaced the same day with the same individual

host anemones. The first of these was approx.

25 cm from the perimeter of the colony where

the anemones were edge to edge. The second

was approx. 15 cm from the first, and approx.

35 cm from the edge of the massed colony. Each

of these anemones had sand around the base

(the wentletraps prefer anemones with sand, in

which they hide and with which they agglutinate

their egg capsules). The numbers of Epitonium

albidum per Stoichactis were the maxima

observed in the three study areas, and it was

hoped that the sand and peripheral position of

the anemones would minimize wentletrap

wandering. On February 11, 1982, 13.8 days

later, an attempt was made to retrieve as many

as possible of the 51 marked wentletraps from

the two anemones. Of the 16 wentletraps col-

lected, 5 had been marked. On February 13,

1982, 15.8 days later, a sixth marked specimen

was found with a Stoichactis about 1 m away.

Judging by their initial and final shell lengths

(between 2.2 and 6.1 mm), all the retrieved

specimens were either immature or male

(Robertson, 1981b).

Observations

Observations were made on the way Epito-

nium teleoconchs grow. A new intervarix (the

space between two varices) and varix grow

simultaneously, thinly at first, then both sub-

sequently thicken. Thus the intervarix and new

rib either are thin and easily broken back to the

preceding rib, or they both are thick and the

outer lip coincides with the last rib. Some indi-

viduals had a thin intervarix and a thin outer lip

Vol. 97(2

April 6, 1983

THE NAUTILUS 63

rib, while others, collected and marked at the

same time of day, had already thickened these

shell parts. Three out of four specimens found

with one Sfoichactis at Virgin Gorda had abnor-

mally thickened penultimate axial ribs.

Specimens found with Stoickactis but kept

separate from the anemone host usually ceased

shell growth in the laboratory, although a thin

outer lip was usually thickened or a replacement

rib was sometimes grown if the lip needed

repair. If new (but not replaced) shell was

grown, the intervarix was usually narrowed

relative to the immediately adjacent inter-

varices and grew within a day or so (Fig. 4). On

one occasion two new ribs were grown, the sec-

ond also with a narrow intervarix (Fig. 5). These

new ribs were noteworthy in being slanted rela-

tive to the preceding rib.

Experimental Results

Growth rate data from the specimens in

aquaria are given in Table 1; field data are in

Table 2. Twenty-three specimens were studied

in all, 14 of them in the immature or male phase,

and 9 of them in the female phase (field data on

females and laboratory data on large females -

FIGS. 4 and 5. 4, Lateral view of th£ anterior end of an

Epitonium aibidum shell showing normally spaced varices, a

narrow intervarix and a new outer lip rib grown the first

night in the laboratory (kept away from Stichodactyla^. 5,

Same, shoimng two nefw narrow intervarices and two new

outer lip ribs groum during two successive nights in the

laboratory (kept away from Stichodactyla).

13-25 mm long -unfortunately are lacking).

The data are difficult to interpret because

days elapsed between measurements are not the

same, because there are few individuals in each

size class, and because data are not normally

distributed when some size classes and days

elapsed between measurements are standard-

ized. Additionally, females must be considered

separately from males.

Several points are nevertheless clear: 1. Non-

normally distributed data and unacceptably high

variance result when animals were measured

less than 5 days apart. 2. Three egg-laying

females gave anomalously high measurements:

mean teleoconch length increments of 0.33 to

0.50 mm per day. 3. One class of measurements

yields sufficient data to make a clearcut and

reliable statement about growth, i.e. males

maintained for 7 or more days (N = 8) grew

0.15 ±0.04 mm in mean teleoconch length per

day (Barbados animals only). The sizes of the

animals do not appear to affect this rate over a

range of 2.2 to 6.2 mm in initial shell length. 4.

The same eight specimens grew a mean number

of 1.2 ±0.5 ribs per day.

Discussion and Conclusions

The observations on teleoconch growth ex-

plain why Epitonium shells are rarely seen with

the outer lip between varices (only when it has

been partially broken back). Similar conclusions

were reached by Linsley and Javidpour (1980)

on much larger, distantly related varix-bearing

prosobranchs such as Cassis and Murex.

The observation that not all individuals were

at the same stage of growth at the same time of

day does not favor the 24-hour cycle hypothesis.

The observation that three specimens collected

with one Stoickactis at Virgin Gorda all had ab-

normally thickened penultimate axial ribs does

suggest, however, that some extrinsic factor

was causative.

When Epiforiinm aibidum is parted from its

host and if growth does not cease promptly, the

spacing between the one or two newly grown

ribs usually narrows (Figs. 4-5). It is note-

worthy that such changes in spacing should so

readily occur, the number of ribs on the last

whorl being such a sacrosanct taxonomic char-

64 THE NAUTILUS

April 6, 1983

Vol. 97(2)

TABLE 1. Data on shell growth in Epitonium albidum, based on Barbados specimens maintained in

aquaria in which Stichodactyla helianthus and sand were available (early February).

'Specimen unable to right itself, and thus did not feed.

TABLE 2. Mai-k and recapture data on shell growth in Epitonium albidum during 13.8 - 15.8 day periods in the field with

Stichodactyla helianthus at Virgin Gorda and Barbados. All observations were made in late January and February. Sex: all

immature or male.

Virgin Gorda (VG) Estimated Mean shell No. ribs grown

or initial shell Days Final shell

Barbados (Bl length I in in) elapsed length I nun)

Mean shell

length increments/

day I mm)

RcpldceinenI New

Mean ribs/

day

acter in epitoniids. The ribs usually are lined up

and joined together from one whorl to the next

(Figs. 4-5), and thus the normal spacing is

established early in postlarval life. Even so,

would the spacing (and "species") change if a

wentletrap moved from a preferred host to a

less "desirable" host?

The data available do not enable me to deter-

mine a growth curve for Epitonium albidum.

Nor do they make possible statistical compari-

sons between the laboratory and field data, or

between individuals in the male and female

phases (do females grow faster than males?).

How well do the data accord with a sigmoid

curve? It is not possible to establish the sizes at

which the first lag phase ends and the second

begins, or the point on the curve where sex

change occurs (is there a deflection?). The best

data set analyzed in the Experimental Results

section (from the 8 Barbados males) may have

Vol. 97(2)

April 6, 1983

THE NAUTILUS 65

come from animals still in the initial lag phase

when the first measurements were made and

from animals probably in the log phase when the

final measurements were made. In other words,

data from these specimens probably do not in-

dicate the greatest mean rate of growth that can

be attained in the log phase. This may help to ex-

plain part of the increased rate of growth of the

three small- to medium-sized females, but does

not explain the inordinately high rates and why

the data are so erratic. Presumably they were at

the middle or high end of the log phase. This

raises questions: Under what conditions does

Epitonium albidum grow large enough to reach

the final lag phase? Is the upper end of the

growth curve usually truncated? What causes

death?

According to Guinther (1970), growth of

Epitonium ulu slows soon after "maturity" is

reached, but this study of E. (tlhiduni indicates

that the second lag phase begins well after the

beginning of female maturity.

It is puzzling to me why the data from fewer

than 5 days growth are so erratic. Two of the

three fastest-growing Epitonium alhidum

females grew for only 2.0 days, but the one that

grew 0.33 mm/day did so for a more sustained

time - 12.0 days (Table 1). In five instances the

growth rate data for & albidum exceed the rate

recorded in young postlarval E. ulu (>0.19 mm/

day) (Taylor, 1977) (Tables 1-2). The percent

daily shell length increase of one male was as

high as 7.5 (2.8 to 5.7 mm in 13.8 days; Table 2).

Other isolated data sets also indicate fast

growth m Epitonium albidum, specifically those

relating to rib growth. In three out of seven in-

stances in the field, 23 or 24 ribs were grown in

13.8 days (1.7/day) (Table 2). There are between

10 and 13 ribs on the last whorl of mature E.

albiduyn (mean of 42 counts: 11.5), so about two

whole new whorls were grown during this time.

In two instances in aquaria, females grew 2.0

ribs/day, but only for 2.0 days (Table 1).

Episodic growth in the much larger proso-

branchs such as Cassis and Murex discussed by

Linsley and Javidpour (1980) must involve tre-

mendously fast spurts of growth. After each

spurt, after each outer lip is thickened, there is a

long period of quiescence insofar as shell growth

is concerned. Although Epitonium has growth

spurts, these are much smaller in scale. Periods

of quiescence involve only hours or a day or so at

most, not months.

What could account for the rhythmic growth

of Epitonium varices? A 24-hour cycle? A tidal

cycle? Rhythmic feeding? An endogenous

rhythm? As for the first and second possibilities:

there is no one-to-one relationship between rib

growth and either of these physical factors.

However, the situation may be more compli-

cated. The day-night and tidal rhythms were

obscured in the laboratory. Considering only the

Barbados field data (Table 2), the mean number

of ribs grown per day is 1.2 ±0.5. It seems

unlikely that a 24-hour cycle would result in

more than 1.0 rib per day. However, the tidal cy-

cle at Barbados is semidiurnal (1.9 high or low

tides per day). Allowing for some lag and

behavioral variance, E. albidum growth spurts

possibly are affected or regulated by the tides.

The data are, however, so variable that the lat-

ter hyjiothesis requires rigorous testing. As for

the third and fourth possibilities: no data are

available.

The data do show that Epitonium albidum

definitely does not consistently grow 1.0 rib per

day, and thus shells cannot be aged simply by

counting total numbers of varices.

Breyer's (unpublished) data show that the

California species Epitonium tinctum (Carpen-

ter, 1864) has an annual life cycle; growth is

slow. Relative to its size and shell thickness, E.

albidum must be one of the fastest-growing

gastropods. The data reported here suggest that

it has a short sub-annual life cycle.

Acknowledgments

I am indebted to Dr. Finn Sander, Director of

the Bellairs Research Institute of McGill Univer-

sity, Barbados, for the continued provision of

excellent facilities. Harriet H. Robertson helped

in many ways in the field, the laboratory, and at

home. Dr. George M. Davis's help with organiza-

tion, analysis, and presentation was invaluable.

Various drafts of the manuscript were also kind-

ly read and criticized by: Dr. Arthur E. Bogan;

Amy Breyer; Helen DuShane; Dr. Kenneth L.

Heck, Jr.; Robert Hershler; Dr. E. Alison Kay;

66 THE NAUTILUS

April 6, 1983

Vol. 97(2)

Virginia Orr Maes; Dr. Finn Sander; Dr.

Charles W. Thayer, and Dr. Joseph Vagvolgyi.

LITERATURE CITED

Ankel, W. E. 1936. Prosobranchia. In: G. Grimpe and E.

Wagler (eds.), Die Tierwelt der Nord- und Ostsee. IX. 1),,

Lief. 29:1-240. Leipzig, Al<ad. Verlagsgesellsch.

Berry, A. J. 1962. The growth of Opinthostoma (Plecto-

stoma) retrirvertens Tomlin, a minute cyclophorid from

a IVIalayan limestone hill. Pi-oc. Malac. Soc. London

35(l):46-49, 1 fig.

1963. Growth and variation of the shell in certain

Malayan limestone hill snails. Proc. Malac. Soc. London

35(5)':203-206, 1 fig.

Bosch, H. F. 1965. A gastropod parasite of solitary corals in

Hawaii. Pacific Science 19:267-268, 1 fig.

Breyer, A. Unpublished (1982). Observations on the repro-

duction, feeding and ecology of the wevtletrap Epitonium

tinctum (Gastropoda: Mesogastropoda). M.A. thesis,

Sonoma State University, ix + 50 pp., 8 figs.

Comfort, A. 1957. The duration of life in molluscs. Proc.

Malac. Soc. London 32(6):219-241, 2 figs.

Dunn, D. F. 1981. The clownfish sea anemones: Stichodac-

tylidae (Coelenterata: Actiniaria) and other sea anemones

symbiotic with pomacentrid fishes. Trans. American

Philos. Soc. 71(1):1-115, 60 figs.

Guinther, E. |B]. 1970. Biology of some Hawaiian Epito-

niidae. In: The Biology of Molluscs. A collection of

abstracts from the National Science Foundation Graduate

Research Training Program June-September 1968. Univ.

Hawaii. Hawaii /(i.s<. Mar. Biol. Techn. Kept. 18:10-11.

Linsley, R. M. and M. Javidpour. 1980. Episodic growth in

Gastropoda. Malacologia 20(1):153-160, 7 figs.

Robertson, R. 1963. Wentletraps (Epitoniidae) feeding on

sea anemones and corals. Proc. Malar. Soc. London

35(2-3):51-63, pis. 5-7.

1970. Review of the predators and parasites of

stony corals, with special reference to symbiotic proso-

branch gastropods. Pacific Science 24(l):43-54.

1981a. ["1980"). Epitonium millecostatum and

Coralliophila clathrata: two prosobranch gastropods

symbiotic with Indo-Pacific Palythoa (Coelenterata:

Zoanthidae). Pacific Science 34(1):1-17, 13 figs.

1981b. Protandry with only one sex change in an

Epitonium (Ptenoglossa). Navtibis 95(4):184-186, 1 fig.

1983. Observations on the life history of the

wentletrap Epitonium albidum in the West Indies.

American Malac. Bull. 1:

Taylor, J. B. 1977. Growth rates in juvenile carnivorous

prosobranchs (Mollusca: Gastropoda) of Kaneohe Bay,

Oahu (Hawaii). Proc. Third Intematl. Coral ReefSymp..

Univ. Miami, pp. 253-259, 8 figs.

nilier, S. 1981. Clines, convergence and character displace-

ment in New Caledonian diplommatinids (land proso-

branchs). Malatotogia 21(l-2):177-208, 35 figs.

DISTRIBUTION OF CORBICULA FLUMINEA THE

ASIATIC CLAM, IN MISSISSIPPI'

Paul D. Hartfield and Charles M. Cooper

ABSTRACT

Corbicula fluminea (Miiller, 177 U) is recorded from 10 of the 12 major drainage

basins of Mississippi. The absence of this clam is associated with shallow, sandy

7-ivern nnd short coastal drainages influenced by tides.

In Mississippi, the Asiatic Clam was first dis-

covered in the Yazoo River in 1963 (Hubricht,

1963). It has since been found in the Pearl, Leaf,

and Coldwater Rivers (Heard, 1965); the Pasca-

goula and Chickasawhay Rivers, and the Ten-

nessee River tributaries in northeastern Missis-

'Contribution of the Mississippi Museum of Natural Science.

Jackson, MS, 39202 and the Sedimentation Laboratory,

ARS, U.S. Department of Agriculture, Oxford, MS 38655.

sippi (Grantham, 1967); the Tombigbee River

(Grantham, 1969); the Tangipahoa and Amite

Rivers (Stern, 1976); the Yalobusha River and

Grenada Lake (Cooper and Johnson, 1980); and

the Big Black River (Hartfield and Rummel,

1981).

Mississippi (Fig. 1) is divided into 12 major

drainage basins. We made 51 collections of

Asiatic Clams in ten of these systems with the

following observations:

Vol. 97(2)

April 6, 1983

THE NAUTILUS 67

FIG. 1. Drainage systems in Mississippi (After Cook, 1959).

Black dots (*) indicate collections by the authors.

1. Yazoo-Sunflower Basin. Dispersed through-

out Bear Creek, the Sunflower, Yazoo, Talla-

hatchie, Yocona, Yalobusha and Coldwater

rivers, and in Lakes Bolivar, Ferguson and

Washington.

2. Big Black River. Locally abundant in the

lower half of the Big Black River and is occa-

sionally found north of Madison County.

3. Southwest Mississippi. Absent from Bayou

Pierre and the Homochitto and Buffalo Rivers.

4. Amite River. Common, but we have not

found any dense populations.

5. Tangipahoa River. Locally common.

6. Pearl River. Abundant throughout the

drainage including the Yockanookanay and

Strong Rivers, and Ross Barnett Reservoir.

7. Coastal Terrace. Absent.

8. Leaf River. Locally abundant throughout

the drainage including Okatoma Creek. Dense

populations are found in the lower part of the

river between Hattiesburg and the mouth.

9. Chickasawhay River. Locally abundant in-

cluding Chunky River.

10. Pascagoula River. Abundant above tidal

influence but not found in the Black and Red

Creeks.

11. Tombigbee River. Locally abundant popu-

lations throughout the drainage including the

Buttahatchee River and the Tibbee, Luxapalila

and Bull Mountain Creeks.

12. Tennessee River. Very abundant in Bear

Creek. Animal middens on the creek bank are

almost exclusively Corbicula.

There are two general regions of Mississippi

where Asiatic Clams have not been collected.

The first consists of the tributaries which drain

into the Mississippi River below Vicksburg, Mis-

sissippi and includes Bayou Pierre, Homochitto

River and Buffalo River. The absence of Corbi-

eula from these streams does not appear to be

due to inaccessibility, since the clam is well-

established to the north in the Big Black River,

to the east in the Pearl River, to the south in the

Amite River and to the west in the Mississippi

River. Other bivalves are also quite rare in these

three streams. Only five species of unionids

were found in Bayou Pierre: Potamilus pur-

puratus. Lajmpsilis ovata ventricosa, L. stra-

minea daibomensis, Leptodea fragilis, and

Tritogonia verrucosa. Only two species, Lamp-

silis radiata luteola and Villosa lienosa have

been collected in the main channel of the Homo-

chitto River, but four additonal species (Toxo-

lasrna texasensis. Fusconaia flava. Uniomerus

decliinis. Anodonta imbeciUis) have been found

in a small tributary. No bivalves were collected

from the Buffalo River.

Physical characteristics shared by the three

streams explain their limited bivalve fauna. Sub-

strates are primarily sand or gravel mixed with

sand. Quicksand varying from 15 to 60 cm deep

is found throughout the drainages. In general

the sand can be considered a very unstable and

shifting substrate on which few bivalve species,

including Corbicula, can survive. The streams

have a low flow channel which meanders within

wide, sandy flood channels. In the Homochitto

River the flood channel may be 400 m wide while

the low flow channel is less than 30 m. The low

flow channel also meanders extensively as a

result of seasonal flooding. One collecting site in

the main channel of the Homochitto in 1981 was

68 THE NAUTILUS

April 6, 1983

Vol. 97(2)

dry sand and over 25 m from the low flow chan-

nel in 1982. In the headwaters where the ratio of

the flood channel is much less, long stretches of

sand and gravel are exposed during low flow.

These sand and gravel bars act as solar collec-

tors during the summer months and water tem-

peratures reach or exceed 35° C. It appears like-

ly that shifting sandy substrates and high sum-

mer water temperatures are limiting factors to

all bivalve fauna including Corbicula in this

drainage.

The second region where Corbicula could not

be found consists of the streams that drain into

the Mississippi Sound between the Pearl and

Pascagoula Rivers, i.e. the Jordan, Wolf, Biloxi,

and Tchoutachabouffa Rivers. All of these

streams have comparatively short and narrow

drainages, are bayou-like at their mouths with

tidal influences extending upstream for several

km. Their headwaters above tidal influence are

shallow and often have sandy substrates, char-

acteristics common with the streams in the

lower Mississippi drainage. Naiad molluscs are

also rare in these coastal drainages. Grantham

(1969) reported three species from the Wolf

River (L. claibomensis, Villosa vibex, Rangia

cuneata). two from the Jordan (Villosa lien.osa

lienosa, Rangia cuneata), four from the Biloxi

(Lampsilis claibomesis. V. lienosa, V. vibex, R.

cutieata) and one from the Tchoutachabouffa (R.

cuneata). During our survey, we also found

Fusconaia cerina and Villosa lienosa in the Wolf

River, but were unable to add to the faunal lists

of the other streams.

Acknowledgments

We wish to thank M. Pierson, C. A. Schultz,

R. Rummel, and D. Heins for providing addi-

tional collections of Corbicula, and Dr. Dave

Stansbery for assistance in identifying naiads.

Specimens and locality data from this study are

on file in the Mollusca Collection of the Missis-

sippi Museum of Natural Science, Jackson, MS.

LITERATURE CITED

Cooper. C. M. and V. W. Johnson. 1980. Bivalve Mollusca

of Yalobusha River, Mississippi. The Nautilus 9i{'l):22-24.

Grantham, B. J. 1967. The Asiatic clam in Mississippi. Proc.

Miss. Water Res. Conf., pp. 81-85.

1969. The freshwater pelecypod fauna of Missis-

sippi. Dissertaiion. Univ. of Southern Miss.

Hartfield, P. D. and R. G. Rummel. 1981. Mussels of the Big

Black River. Miss. Acad. ofSci. Abstracts 26:128.

Heard. W. H. 1966. Further records oi Corhicida fluminea

(Miiller) in the Southern United States. The Nautilus

79:142-143.

Hubricht, L. 1963. Corbieula Jlmninea at Vicksburg, MS.

The Nautilus 77:143.

Sickel, J. B. 1973. A new record of Corhiculn manileti^is

(Philippi) in the Southern Atlantic Slope Region of Geor-

gia. The Nautilus 87:11-12.

Sinclair, R. M. and B. G. Isom. 1963. Further studies on the

introduced Asiatic clam (Corbicula) in Tennessee. Tenn.

Stream Poll. Cont. Board, Tenn. Dept. of Public Health,

pp. 1-51.

Sinclair, R. M. 1971. Annotated bibliography on the exotic

bivalve Corbicula in North America. 1900-1971.

Sterkiana 43:11-18.

Stern, E. M. 1976. The freshwater mussels (Unionidae) of

the Lake Maurepas-Pontchartrain-Borgne drainage sys-

tem, Louisiana and Mississippi. Dissertation, La. St. Univ.

THE PLANORBID SNAIL MICROMENETUS DILATATUS AVUS (PILSBRY)

IN THE WEST INDIES AND CENTRAL AMERICA

Fred G. Thompson

Florida State Museum

University of Florida

Gainesville, Florida 32611

During recent years I collected several sam-

ples of a minute planorbid snail in Haiti and

Jamaica. The specimens are indistinguishable

from a form described as a Pliocene fossil from

Florida. This is also the same as a form de-

scribed from Panama. The snail is properly

Vol. 97(2)

April 6, 1983

THE NAUTILUS 69

classified as follows: Walter (1970, Ann Rept.

Amer. Malac. Union: 47-51) establishes the use

of the generic name Micromenetiis.

Micromenetus dilatatus aims (Pilsbry)

Plariorbis alabamensin avua Pilsbry, 1905; The Nautilua

19:34. (Type locality: Pliocene, Caloosahatchee Forma-

tion, near Clewiston, Florida).

Pronienetus (Microvienetus) alabamensis aims (Pilsbry),

Baker, 1945; Molluscan Family Planorbidae, Univ. Illinois

Press: 190; pi. 121, figs. 40, '41; pi. 12,3, fig. 1; pi. 140,

fig. 25.

Promenetus minutus Taylor, 1954; Rerista Soc. Malac.

"Carlos de La Torre" 9:37-38. (Type locality: Allee Creek,

Barro Colorado Island, Panama).

M. d. avios is generally distributed throughout

the Florida peninsula and intergrades with M. d.

dilatatus (Gould) in northern Florida. It is fig-

ured and discussed in greater detail in a forth-

coming manual on Florida freshwater gastro-

pods. M. alabamensis is a different species.

West Indian records for M. d. aims are: HAITI:

Dept. du Sud, creek 14 km N, Cavaillon, 350 m.

alt. (UF 32419), irrigation ditch, Les Cayes (UF

34995). JAMAICA: St. Elizabeth Parrish,

marshy stream 1.6 mi. SE Snipe (UF 34995).

Undoubtedly the species is more widely distri-

buted in the Greater Antilles. It is seldom col-

lected because of its small size (less than 2 mm)

and its secretive habit of living under stones and

vegetative debris.

A NEW HUMBOLDTIANA (PULMONATA: HELMINTHOGLYPTIDAE)

FROM NORTHWESTERN COAHUILA, MEXICO

Artie L. Metcalf

Department of Biological Sciences

University of Texas at El Paso

El Paso, Texas 79968

ABSTRACT

A new species of helminthoglyptid land snail, Humboldtiana malenae, is de-

scribed from mountains of northwestern Coahuila, Mexico. Its relationships with

other members of the genus in the area are discussed.

Southward from the eastern side of Big Bend

National Park, in Texas, extends a complex but

generally linear series of mountain ranges into

Coahuila, Mexico. These include, north to south,

the Sierra del Carmen, the Sierra Jardin and the

Sierra Maderas del Carmen. Between this latter

range and the Sierra de la Encantada, which ex-

tends on to the south and southeast is the lower

Mesa de los Fresnos. At the southern end of this

mesa is a pass across the mountains called La

Cuesta de Malena or La Cuesta de Plomo.

Standing at the pass, one can observe the arid

basins and ranges of the Chihuahuan Desert to

the west and, to the east, the more mesic

mesquite-grassland of Coahuila, where copious

rains had fallen in the days before a visit made

here in May 1981. The rains had drenched the

area of the pass as well and stimulated activity

in the Humboldtiana reported here. Vegetation

comprised grasses and a shrubby matorral.

Snails were associated with accumulations of

igneous rocks.

I am grateful to Mr. David H. Riskind and Mr.

Robert Burleson, who organized and conducted

the field excursion during which we visited La

Cuesta de Malena, and especially to Mr.

Burleson, who provided transportation. Mr.

William Murray helped in making collections.

70 THE NAUTILUS

April 6, 1983

Vol. 97(2)

Humboldtiana malenae new species

Figs. 1-4

Diagnosis: A relatively small Humboldtiana

with narrow brown bands on shell, the lower-

most band poorly developed; younger whorls

granulose; most of surface of the first whorl

smooth; and with mucus gland ring situated

closely above the dart sacs.

Description of Holotype: Shell subglobose,

24.9 mm in diameter and 21.1 mm high; spire

rising steeply, forming angle of 105°; 4.2

whorls, with body whorl rounded peripherally

and descending; aperture sub-rounded, 13.8 mm

wide and 15.0 mm high, inclined at angle of 25°

to vertical; columellar peristome reflected and

covering most of umbilicus, leaving only a slit;

outer lip thin; most of first whorl smooth but

with low, fine, slightly sinuous growth lines ap-

pearing at 0.9 whorl, these bearing minute

granules, widely separated; both growth lines

and granules gradually becoming stronger on

younger whorls, the former developing into

light yellow growth ridges that alternate ir-

regularly with lower, brownish areas and with

granules becoming larger, light yellow in color,

and more elongate on dorsal surface, occurring

both on and between ridges; granules less

developed ventrally; two pale brown bands arise

at 1.6-1.8 whorls, one centrally and one

peripherally located on the upper surface of the

whorl, these becoming darker in color and bet-

ter defined on younger whorls with both located

above the periphery of the body whorl; in some

places traces of a third brown band below the

periphery of the body whorl are barely discerni-

ble. There are several irregularities on the shell

surface caused by injury and shell regeneration.

Variatioyi in shells of paratypes: The sub-

peripheral brown band, which is almost indis-

cernible on the shell of the holotype, is moder-

ately well developed on some paratypes. The

relationship of width to height of shell is vari-

FIGS. 1-3. 1 and 2, Apcrtural and dorftal vieum of holotype n/'Humhokitiana malenae new

species (2i.9 mm diaineler). 3, Genitalia of a paratype ofH. malenae (dimevsions listed

first, of four, in text under heading "Genitalia").

Vol. 97(2)

April 6, 1983

THE NAUTILUS 71

mm

FIG. 4. Lower region of genitalia of same specimen o/Hum-

boldtiana malenae new species as in Fig. 3. A-atrium,

DS-dart sac. E-epiphallus. EF-epiphaUic flagellum. FO-Jree

oviduct, MG-munis gland ring, P-penis, PR-penial retractor

muscle. PS-prostatic part of spermaviduct, SD-spermathecal

duct, US-uierine part of spermoviduct, V-unaltered part of

vagina, VD-vas deferens.

able. Thus, the ratio diameter/height for 10

paratypes ranges from 0.99 to 1.19 with mean of

1.08 and standard deviation of 0.059. For these

paratypes, the following dimensions (in mm) and

counts were obtained (mean is before, range

within, and standard deviation after parenthe-

sis): shell diameter, 23.9 (21.5-27.5) 1.81; shell

height, 22.3 (20.1-24.1) 1.3; apertural width,

13.8 (12.5-15.8) 1.04; apertural height, 15.5

(14.0-16.7) 0.81; number of whorls, 4.21

(3.8-4.45)0.19.

Genitalia: (See Figs. 3 and 4). The penis is

relatively short for the genus (Solem, 1974:

Table 1). Internally, a verge occupies 0.4 to 0.65

of the length of the penis in four specimens dis-

sected. Internally, the verge is much like that

reported by Solem (1974:361) ior Humboldtiana

fullingtoni Cheatum, 1972, with an accessory,

inner membrane, covering four pilasters that

continue downward from the epiphallus. These

pilasters are deeply corrugated transversely and

continue to the lower end of the verge where

they terminate in four finger-like projections.

As in H. fullingtoni also, a circular pilaster sur-

rounds the lower part of the verge on the inner

wall of the penis in two specimens. Below this

circular pilaster arise 5 or 6 longitudinal

pilasters, which extend on to the atrium. The

epiphallus bears a well developed flagellum. The

lowermost, unaltered vagina is short. The

mucus gland ring is very close to the dart sacs

and both are large features. In some specimens

(as in Fig. 4) there is no externally discernible

separation between mucus glands and dart sacs,

while in others a separation not exceeding 0.5

mm is observed. The free oviduct above the

mucus glands is short. The spermathecal duct

bears a relatively long appendix shortly below

its terminal bulb. Measurements (in mm) of

genitalia for four specimens dissected (from

shells with diameters, respectively, of 26.3,

23.8, 22.5 and 22.5 mm) are: length of penis

from penial retractor insertion to atrium, 6.5,

7.0, 5.5, 7.9; length of vas deferens, 19.0, 18.5,

11.5, 16.3; length of epiphallic flagellum, 49.3,

53.7, 42.4, 48.4; length of unaltered vagina

below dart sacs, 2.2, 2.2, 1.4, 2.4; height of

largest dart sac, 3.0, 2.8, 2.2, 2.4; height of

smallest dart sac, 2.3, 2.1, 1.4, 1.5; height of

mucus glands, 2.7, 2.1, 2.0, 2.1; length of sper-

mathecal duct plus terminal bulb, 63.5, 59.8,

47.5, 68.6; length of spermathecal appendix, 9.8,

9.7, 7.9, 8.6; length of free oviduct, 2.7, 3.5, 1.3,

3.2. Clearly the relationship between shell size

and size of genitalia is not always closely cor-

related. Thus, the two shells with diameter of

22.5 mm vary considerably in dimensions of the

genitalia.

Types: Holotype: National Museum of Natural

History, USNM 784,768. Paratypes: USNM

784,769; Dallas Museum of Natural History

5,366; University of Arizona 19,045; University

of Texas at El Paso 4647 and 8786.

Type and only known locality: MEXICO,

Coahuila; 28°43'55"N, 102°30'39"W; in boun-

dary area between municipios of Ocampo and

Muzquiz; on Coahuila state highway 53, 6.2 km

slightly N of E from Tres Caminos (village);

above head of north branch of Canada la Virgen

72 THE NAUTILUS

April 6, 1983

Vol. 97(2)

(draining west); at summit of pass (about 1,625

m in elevation) variously termed La Cuesta, La

Cuesta de Malena and La Cuesta de Plomo,

where unimproved road diverges northerly from

Highway 53 towards the Mesa de los Fresnos

and 0.2 km NW of junction with road leading to

mining area to south. The above based on the

CETENAL 1:50,000 topographic quadrangle

for Sierra la Encantada, which, however, shows

La Cuesta de Malena 2.5 km east of the type

locality. Holotype and paratypes were collected

by the author on 7 May 1981 and additional

paratypes by David H. Riskind on 25 May 1975.

Etymology: (maleyiae = of Malena) The place

name Malena used in the area derives from the

given name (contraction of Maria Elena) of a

restaurateuse, who formerly maintained a cafe

in the area (David H. Riskind, pers. comm.).

Comparisons: Geographically, Humboldtiana

malenae occurs between the ranges of Humbold-

tiana taylori Drake, 1951, to the north in the

Sierra Maderas del Carmen, and of Humbold-

tiana plana Metcalf and Riskind, 1976, to the

southeast in the Sierra Santa Rosa (Metcalf and

Riskind, 1976, 1979). Shells of all 3 species are

granulose but those of H. malenae are less so

than in the other 2 species. The subglobose,

elevated shells of H. malenae differ trenchantly

from the low, depressed shells of H. plana. The

internal anatomy of H. plana is not known.

Mature shells of H. taylori reach well over 30

mm in both diameter and height, whereas speci-

mens of H. malenae observed do not reach 30

mm in either dimension. Although variable, the

shells. of if. taylori are consistently much darker

than those of H. malenae. Distinct, close-set,

minute granules occur already on whorls 0.5-1.5

in H. taylori. In H. malenae, fine growth lines

(which bear only widely separated tiny granules)

first appear at about 0.9 whorl.

A distinctive difference between H. taylori

and H. malenae involves the female genitalia,

with the mucus gland ring and dart sacs of the

latter being exceptionally widely separated, for

the genus. In H. malenae, on the other hand,

these structures are barely separated. In this

respect, the situation with H. malenae is like

that in the "first" group of Burch and Thompson

(1957:2) or the group of Humboldtiana buffo-

niana (Pfeiffer, 1845).

In Big Bend National Park 3 species of Hum-

boldtiana have been reported (Pratt, 1971:433,

434). A similar diversity of species probably

prevails in the complex of mountains south of

the park, which are dealt with here. In addition

to H. taylori and H. malenae, shells of small

Humboldtianas, possibly allied with H. malenae.,

have been collected from 2 localities in or north

of the Sierra Jardin. Shells from one locality are

similar to those of H. malenae. A single shell

from a second locality is distinctive in lacking

any indication of brown bands.

LITERATURE CITED

Burch, J. B. and F. G. Thompson. 1957. Three new Mexican

land snails of the genus Humboldtiana. Occns. Pap. Mus.

Zool. University Michigan 590:1-11, pis. 1-5.

Metcalf, A. L. and D. H. Riskind. 1976. A new Humbold-

tiana (Pulmonata: Helminthoglyptidae) from Coahuila,

Mexico. The Nautilus 90:99-100.

1979. New information concerning Humbold-

tiana taylori Drake, 1951 (Gastropoda: Pulmonata: Hel-

minthoglyptidae). Th£ Veliger 22:179-181.

Pratt, W. L. 1971. Humboldtiana agavophila. a new helmin-

thogl.vptid land snail from the Chisos Mountains, Big

Bend National Park, Texas. Thf Southwestern Naturalist

15:429-435.

Solem, A. 1974. On the affinities of Humboldtiana fulling-

toni Cheatum. 1972. The Veliger 16:359-365.

Vol. 97(2)

April 6, 1983

THE NAUTILUS 73

LIVING PROMENETUS EXACUOSUS (PULMONATA: PLANORBIDAE)

FROM NORTH CENTRAL TEXAS

William L. Pratt

Museum of Natural History

University of Nevada, Las Vegas

Las Vegas, Nevada 89154

ABSTRACT

Living Promenetus exacuosus (Say) occur in Tarrant County, Texas, slightly

south of latitude 33° N. The previous southern limit in the nfiid-continental region,

a disjunct station, was from approximately 36°N. The Texas population is be-

lie^'ed to be a naturally occurring Pleistocene relict.

The small planorbid snail, Proynenetus exacuo-

sus (Say, 1821), has been reported as far south

as latitude 32 °N in the great plains region

(Callahan County, Texas, Cheatum et al., 1972),

but all such southern records have been either

from river drift, or directly from Pleistocene

sediments. D. W. Taylor (in Hibbard and Taylor,

1960) surveyed American collections for live-

collected material and found that P. exacuosus

had been collected as far south as Valencia

County, New Mexico, in the Colorado Plateau,

but in the great plains did not occur alive south

of latitude 39°N, with a disjunct station in a

spring in Meade County, Kansas. Branson

(1961) reported the collection of living speci-

mens in Adair County, Oklahoma, at latitude

36° N in the Ozark region, "in a boggy area near

the creek."

It is, therefore, of some interest to find the

species alive slightly south of latitude 33° N, in

Tarrant County, Texas. The specimens were col-

lected in the course of an ecological study during

the spring of 1969 on the Fort Worth Nature

Center and Refuge, in the valley of the West

Fork Trinity River, on the west side of Lake

Worth reservoir, at its north end. P. exacuosus

was collected in a seasonal pond in bottomland

hardwood forest, occurring as a corridor along

the Trinity River, which at this point has cut a

deep valley in the limestones of the lower Creta-

ceous comanchean series. The uplands support

tall grass prairie, presently much altered by

grazing. The pond in which P. exacuosus was

collected is refilled annually by winter and

spring overflow of a small adjacent stream, and

dries completely during the summer and fall.

Other species with disjunct stations in the area

include Punctum minutissimMm (Lea, 1841)

(Pratt, 1971) and several plants (QUercus

muehlenbergii Engelm., Hexalectris spicata

(Walt.) Barnh., and Botrychium virginianum

(L.) Swartz). The occurrence of all is related to a

local microclimate which is more mesic, and

probably cooler, than the regional average.

The seasonal pond is deeply shaded, its bottom

covered by a layer of fallen leaves. The substrate

is arenaceous clay (14% sand, 7% silt, 79%

clays), with a 16% organic content. It has

developed in place on sandy Pleistocene terrace

deposits with little organic content. Water pH

was 7.3 on 5 January and 7.1 on 13 April. Total

hardness was 150 ppm and 290 ppm on the same

dates. At the time of the second sample the

stream was high with clear, brown-colored

water and connected with the pond by a single

channel about 20 cm deep. In the January collec-

tion, P. exacuosus occurred at a density of 13.5

per m\ and all were adults. On 13 April the den-

sity had increased to 139 per m\ all juvenile ex-

cept for a single large adult. I conclude that at

this locality P. exacuosus is probably semel-

parous, breeding in the early spring. Associated

species were Fossaria (Bakerilymnaea) techella

(Haldeman, 1867), Fossaria (Bakerilymnaea)

dalli (Baker, 1907), Physella (Costatella) virgata

(Gould, 1855), and Sphaerium (Musculium)

lacustre (Miiller, 1774).

It seems probable that the population repre-

74 THE NAUTILUS

April 6, 1983

Vol. 97(2)

sents a relict of a more extensive distribution

during the Wisconsin Pleistocene, but it does

not seem to have been found in the Pleistocene

deposits of the region, as yet. Cheatum and

Alien (1965) did not report it in their summary

paper of the region, although Promenetus um-

bilicatellus (Cockerell, 1887) was common in

several deposits. Branson (1961), who included

all river drift records, reported only a single ad-

ditional record for Oklahoma, collected by

Wallen and Dunlap (1953) in Texas County,

Oklahoma. (The precise locality and the nature

of the material were not specified in the original

report). Apparently the only Pleistocene locality

in the great plains south of Kansas is Singley's

(1893) record of material from a Pleistocene

deposit in Tule Canyon, Swisher County, Texas.

The alternative possibility is aerial dispersal on

flying aquatic animals, which has been repeated-

ly documented for various bassomatophoran

snails. It seems, however, that if P. exacuosus is

presently dispersing by this means in the south-

ern mid-continental region, it should be much

more common than it is. The few, widely scat-

tered stations from which it is known are

separated by considerably more than an average

single flight distance, with numerous suitable

resting habitats between. The station reported

here is not a good waterbird habitat, is not visi-

ble from the air, and is within a kilometer of

several hundred hectares of open marshland.

While the population probably was established

by aerial dispersal, it seems probable that the

dispersal occurred during the Pleistocene, at a

time when populations of P. exacuosus, although

scattered, were much more closely spaced in the

region.

The specimens have been catalogued in the

author's collection, as numbers 2691 (January

Collection) and 2707 (April Collection).

LITERATURE CITED

Branson, Branley A. 1961. Recent Gastropoda of Oklahoma.

Part II. Distribution, ecology, and taxonomy of fresh-

water species, with description of Helisoma travertina

sp. nov. Oklahoma State Univ. Pub.. Arts and Sci. Stud.,

Biol. Stud.. Ser., (6), 58(17):l-72.

Cheatum, E. P. and D. Allen. 196.5. Pleistocene Land and

Freshwater Mollusks from North Texas. Sterkiana

(18):1-16.

Cheatum, E. P., R. W. Fullington and W. L. Pratt. 1972.

Molluscan records from west Texas. Sterkiana (46):6-10.

Hibbard. C. W. and D. W. Taylor. 1960. Two late Pleistocene

faunas from southwestern Kansas. Contr. Univ. Michigan

Mus. Paleont. 16(l):l-223, pis. 1-16.

Pratt, W. L. 1 97 1 . The land snails of the Greer Island Nature

Center, Fort Worth, Texas. American Malac. Union,

Ann. Kept. 1970. (37):58-60.

Singley, J. A. 1893. Cont. to the natural history of Texas.

Part I. Texas Mollusca. Geol. Sum. Texas. l,th Ann. Rept.:

299-343.

Wallen, I. E. and P. Dunlap. 19.53. Further additions to the

snail fauna of Oklahoma. Oklahoma Acad. Sci., Proceed-

ings 34:76-80.

THE FRESHWATER SNAIL, MENETUS DILATATUS, (PLANORBIDAE)

IN NOVA SCOTIA

Derek S. Davis

Nova Scotia Museum, 1747 Summer Street, Halifax

Nova Scotia, B3H BAG, Canada

ABSTRACT

The small planorbid snail, Menetus dilatatus (Gould), is established as part of

the Canadian molluscan fauna through rediscovery of the species in Nova Scotia.

Since 1970 the Nova Scotia Museum has

undertaken collecting programs that have con-

siderably increased both the holdings of its

scientific collections and knowledge of the

Vol. 97(2)

April 6, 1983

THE NAUTILUS 75

natural history of Nova Scotia. Studies of land

and freshwater Mollusca have been included in

this work. A synopsis of the land and freshwater

mollusca of Nova Scotia is in preparation. The

family Planorbidae in Nova Scotia includes

Gyraulus circumstriatus (Tryon), Gyraulus

dejlectus (Say), GyTaulus parvus (Say), Heli-

soma anceps (Menke), Helisovia campanulatum

(Say), Helisovia trivolvis (Say), Planorbula

armigera (Say), Menetus exacuosus (Say) and

the recently rediscovered Menetus dilatatus

(Gould).

Meyietus dilatatus was first recorded from

"Dartmouth lakes", in Nova Scotia as "Planorbis

dilatata", and reported in 1863 by Robert Willis

(Ganong, 1890). In a catalogue of the recent

Mollusca of Canada prepared by LaRocque

(1954) this record was either overlooked or dis-

counted as a misidentification. Up to the present

time the species has not been considered as part

of the Canadian freshwater molluscan fauna

(Clarke, 1981) although an occurrence within 70

km of the Canadian border in the Niagara Fron-

tier region indicates that it could be expected

(Robertson and Blakeslee, 1948).

In Nova Scotia, M. dilatatus was rediscovered

by the author in Gold River at New Ross, Lunen-

burg County, in 1968, when a single living speci-

men was taken.

Twenty-six other specimens were obtained

from a pond at Waternish in the St. Mary's

River Valley, Guysborough County, in 1973 and

from Lake Fletcher, Lake Thomas, Lake

William and Grand Lake, in Halifax County, in

1973 and 1975. The Halifax County localities

correspond roughly with the original Dartmouth

Lakes locality given by Willis. These localities

are shown on the map (Fig. 1). The species was

not found at about 100 other freshwater locali-

ties sampled. The habitats where M. dilatatus

has been found are ponds, lakes and quiet sec-

tions of rivers. In all cases aquatic vegetation

was present and the snails were attached either

to the plants or to stones. These habitats gen-

erally correspond with the "quiet pools and

ponds" given as the habitat for this species in

New England (Emerson and Jacobson, 1970).

Some Nova Scotia specimens may attain a

larger size than the 2.00 mm diameter usually

given for this species. For example, the speci-

men illustrated in Fig. 2, which was collected at

Lake Thomas in August 1975, is 3.5 mm in

diameter.

FIG. 1. Localities for Menetus dilatatus (Gould) in Nova Scotia. 1. Gold River, New Ross,

Lunenburg Co. 2. Dartmouth Lakes, Halifax Co., and 3. Waternish, Guysborough Co.

76 THE NAUTILUS

April 6, 1983

Vol. 97(2)

FIG. 2. Menetus dilaUitus iGuuld). A tarye Kpecimen col-

lected from the outlet of Lake Thomas. Halifax Co., Nova

Scotia, 6 Aug. 1975. Diameter 3.5 mm. Nova Scotia Muxeuw

cat. no. l'J75Z.250.1,. Photo try A. Wihoti.

The occurrence ofM. dittitatun in Nova Scotia

is of some interest. It is apparently a disjunct

species,, beinj^ separated by several hundred

kilometers from the population in New England.

However, there are not yet sufficient records to

clearly indicate a pattern of distribution. The

Gold River and St. Mary's River watersheds

drain directly to the Atlantic coast, perhaps in-

dicating an association with a coastal plain

glacial refugium that contributed the New

England element to the flora of southwestern

Nova Scotia (Roland and Smith, 1969). The

lakes north of Dartmouth however, drain into

the Bay of Fundy by way of the Shubenacadie

watershed. Menetus dilatatus may have been in-

troduced into these lakes from lakes draining to

the Atlantic coast, following construction of a

canal during the middle part of the last century.

The possibility that the species was introduced

into Nova Scotia from New England was also

considered, as it has become locally established

in Britain following introduction about 1861.

(Ellis, 1951, Kerney, 1976). A similar introduc-

tion into Dartmouth Lakes, which are close to

the Hal if ax- Dartmouth urban area of Nova

Scotia, is conceivable, but the occurrence in the

Gold River and St. Mary's River watersheds

could not be explained in this way.

At the present time it is sufficient to establish

that M. dilatntus is part of the molluscan fauna

of Nova Scotia and of Canada. A study of non-

marine mollusca in Nova Scotia, based upon a 10

km^ grid system, as successfully used in Britain

(Kerney, 1976), is being contemplated. This will

eventually provide a clearer picture of the distri-

bution of M. dilatatus and other poorly known

species in the province.

LITERATURE CITED

Clarke, A. H. Jr. 1981. The Frc.'ihwater of Molluscs of

Canada. National Museums of Canada. Ottawa. 446 pp.

Ellis, A. E. 1951. Census of the distribution of British non-

marine Mollusca. 7th edition. J. Conch. 23:171-244.

Emerson, W. K. anci M. K. Jacobson. 1976. The American

Museum of Natural History Guide to Shells. Alfred A.

Knopff, New York. 482 pp.

Ganong, W. F. 1890. John Robert Willis, the first Nova

Scotian conchologist. Trans. N. S. Inst. Sci. 7(4):404-428.

Kerney. M. (). 1976. Atlas of the non-marine Mollusca of the

British Isles. Cambridge. 20fi pp.

LaRocque, A. 1953. Catalogue of the Recent Motlu.sca of

Canada. Bulletin No. 129. National Museums of Canada.

406 pp.

Roland, A. E. and E. C. Smith. 1969. The Flora of Nova

Sci}tia. Nova Scotia Museum. Halifax. 743 pp.

Robertson, I. C. S. and C. L. Blakeslee. 1948. The Mollusca

of the Niagara Frontier Region. Bull. Buffalo Society of

Natural Sciences 19(3):1-191.

Vol. 97(2)

April 6, 1983

THE NAUTILUS 77

VARIABILITY IN NORTH AMERICAN LYMNAEA STAGNALIS L.

(GASTROPODA: LYMNAEIDAE)

Eva Pip

Department of Biology, University of Winnipeg

Winnipeg, Manitoba, Canada R3B 2E9

ABSTRACT

The nomenclatural history of the pond snail, Lymnaea stagnalis L., in North

America is briefly reviewed. An examination of a. population of coexisting L. s.

appressa (=jugnlaris) and L. s. sanctaemariae shows continuous variation sug-

gesting that these two forms are ecophenotypes.

The pond snail, Lymnaea stagnalis L., is wide-

ly distributed in the northern hemisphere, oc-

curring in North America, Europe, parts of

North Africa and much of Asia (Hubendick,

1951). Its wide tolerance ranges for many water

chemistry parameters (e.g. Pip, 1978) and its ap-

parently effective dispersal have contributed

towards its extremely numerous populations.

The ecological success of L. stagnalis, combined

with the isolation of individual populations

characteristic of many freshwater organisms,

has been accompanied by a great variety of shell

form. The often striking morphological differ-

ences between various populations may be re-

garded in large part as a product of environ-

mental conditions acting on the fixed genetic

resources of a population which frequently

results from interbreeding among the descen-

dants of a small number of colonizers. Any addi-

tional variation in such populations must be in-

troduced through mutation or immigration of

new genotypes.

The variability of L. stagnalis has been

responsible for a long history of nomenclatural

excesses. During the latter part of the 19th cen-

tury a great number of European named varie-

ties were distinguished (e.g. Baker, 1898), many

of which were shown by Vignal (1911) to be ob-

tainable from average populations by altering

environmental conditions. Subsequent workers

(e.g. Zhadin, 1952; Fromming, 1956) suggested

that environmental quality and availability and

type of food were important factors in influenc-

ing variability in this species. Zhadin (1952) and

Germain (1969) declined to assign any names to

subspecific rank in their treatments on the basis

that continuous variation appeared to exist be-

tween different forms.

In North America, the great variability of L.

stagnalis was recognized by Walker (1892-3),

who figured several extreme forms. Baker

(1898) initially assigned the bulk of the North

American forms to L. s. appressa Say 1818 ( = L.

s. jugularis Say 1817), noting the substantial

variation in spire length. Subsequently (1911) he

recognized six forms that were differentiated on

the basis of relative spire length, and degree of

shouldering and lip expansion. Hubendick (1951)

suggested that the forms cited by Baker (1911)

have overlapping ranges of variation and there-

fore represent ecophenotypes. The former

worker pointed out that the full range of varia-

tion may be found in every geographical region

where this species occurs. La Rocque (1968)

presented a similar viewpoint and also noted

that Pleistocene examples of this species in

North America are primarily referable to L. s.

jugularis.

Clarke (1973) compared a number of popula-

tion samples from the Canadian Interior Basin

and found that the major characters distinguish-

ing some of the forms were not valid because of

the extent of variation that could be en-

countered within the same population. He did

retain L. s. sanctaemariae Walker 1892 since

the ratio of the aperture length to total shell

length in population samples of this form (>0.65)

appeared to show some discontinuity with the

ratios for L. s. appressa samples (<0.60). The

former variant also appeared to show some geo-

78 THE NAUTILUS

April 6, 1983

Vol. 97(2)

graphical unity, occurring in the Winnipeg

River, Lake Superior, Lake Huron and Wiscon-

sin River systems, where it is present as local

populations interspersed wnth populations of L.

s. appressa. However some single populations

on the periphery of the range of L. s. sanctae-

mariae were reported by Clarke (1973) to con-

sist of morphological intergrades between the

two forms.

A large population which contains both the

two forms and their intergrades (Fig. 1) is

present in Pike Lake, Cass Co., Minnesota

(47°18'N, 94°37'W), approximately 200 km

south of the present known boundary range of

L. s. sanctaemariae. The aperture length to

shell length ratios measured for a sample from

this population (Fig. 2) indicate that the general-

ly accepted ratios for distinguishing between

the two forms are not valid for this population,

which apparently shows continuous variation.

Shortness of the spire in L. s. sanctaemariae

is generally regarded as a result of selection

pressures in turbulent habitats (Baker, 1928;

Clarke, 1973). The coexistence of both forms

and their intergrades in the same habitat sug-

gests that such a population contains an un-

usually diverse range of genotypes, perhaps the

result of multiple immigrations from different

sources and/or endogenous origin of some

forms. The origin of the short-spired form in

Pike Lake is perplexing as a search of lakes in

the same and adjacent counties yielded only the

long-spired form.

That the morphological diversity of L. stag-

nalis in North America is an ecophenotypic one

is supported by observations that anatomical

sanctaemariae

0,54 0.56 058 060 062 064 0.66 0.68

APERTURE L / SHELL L.

FIG. 2. Distribtdioji of aperture/ shell length ratios rounded

off to nearest even number from a Pike Lake sample of L.

stagnalis.

differences between the forms are minor

(Baker, 1928) or doubtful (Clarke, 1973). How-

ever Alaskan forms of this species may consti-

tute a separate race (Baker, 1928; Hubendick,

1951).

It is interesting that Baker (1928) noted that

some populations of the short-spired form ex-

hibit a high proportion of scalariform-like abnor-

malities characterized by a forward slippage of

the affected whorls along the shell axis. In long-

spired forms this anomaly is very rare. In the

Pike Lake sample approximately 10% of the in-

dividuals exhibited this anomaly (Fig. 1).

Voucher specimens from this sample have

been deposited in the National Museum of

Canada (NMC 77376).

F'IG. 1. Speciiiteits of L. slagiialis _//■()»« a single population

in Pike Lake. Cass Co., Minnesota. First shell is SO mm.

Others to same scale.

LITERATURE CITED

Baker, F. C. 1898. The Mollusca of the Chicago area.

Chicago Acad. Sci. Nat. Hist. Survey, Bull. Ill, Part I.

418 pp.

1911. The Lymnaeidae of North and Middle

America. Chicago Acad. Sci. Spec. Pub. No. 3. 539 pp.

1928. The fresh water Mollusca of Wisconsin.

Part. Wis. Geol. Nat. Hist. Survey. Bull. 70. 507 pp.

Clarke, A. H. 1973. The freshwater molluscs of the Canadian

Interior Basin. MaZocoi. 13:1-509.

Fromming, E. 1956. Biologic der mitteleuropdischen

Silssuminser.schnecken. Dunker & Humblot, Berlin. 313 pp.

Germain, L. 1969. Faune de France. Vol. 22. Libr. Fac.

Sci., Paris. 897 pp.

Vol. 97(2)

April 6, 1983

THE NAUTILUS 79

Hubendick, B. 1951. Recent Lymnaeidae. Kiingl. Suenska

Vetenskap. Handl. Fjdrde Ser. 3: 223 pp.

La Rocque, A. 1968. Pleistocene Mollusca of Ohio. Part 3.

OhwDiv. Geol. Sun>ey. Bull. 62:357-553.

Pip, E. 1978. A survey of the ecology and composition of

submerged aquatic snail-plant communities. Can. J. Zool.

56:2263-2279.

Vig7ial, L. 1911. Quelques observations sur le Liynnaea

stmjmiHs L. F. J. Nat.. Paris. 157-158.

Walker, B. 1892-3. The shell-bearing Mollusca of Michigan.

The Nautilus 6:31-36, 135-141.

Zhadin, V. I. 1952. Moliusks of fresh and brackish waters

of the U.S.S.R. Keys to fauna ofU.S.S.R.. No. 46. U.S.S.R.

Acad. Sci. 369 pp.

UNEXPLAINED OCCURRENCE OF THE MACTRID BIVALVE,

RANGIA CUNEATA, FROM THE ARROWHEAD FARMS INDIAN SITE

NEAR LOUISVILLE, KENTUCKY

Frederick C. Hill

Department of Biology and Allied Health Sciences

Bloomsburg State College

Bloomsburg, PA 17815

ABSTRACT

Over three thousand valves of the Atlantic Coast brackish water clam Rangia

cuneata (Gray) were unearthed from the Arrowhead Farm archaeological site

near Louisville, Kentucky. These clams were brought to this site by Lake Wood-

land or Archaic inhabitants of the Ohio River Valley for unknown reasons. This

represents only the second mid-continent occurrence of these shells in an archaeol-

ogical site. The reasons that the Indians had for bringing them to the Arrowhead

Farm Site are obscure.

Marine mollusk shells in archaeological sites

far from the ocean no longer are startling dis-

coveries because of the rather large numbers of

them found associated with human living areas

(Parmalee 1958). These moliusks, however,

mainly share the common denomination of being

beautiful, unique or useful (Biggs, 1970).

It was thus with great interest that several

thousands of valves of the exceedingly common

estuarine clam, Rangia cuneata (Gray 1831),

were discovered at Arrowhead Farm, a multi-

component Late Archaic through Lake Wood-

land archaeological site in Jefferson County,

Kentucky (USGS Lanesville, Indiana-Kentucky

quadrangle 38° 10' 15' 'N 85° 53' 48' 'W). A

report of the archaeological investigation at the

Arrowhead Site is in print (Mocas 1976). Identi-

fication of the shells as Rangia cuneata (Gray),

the Common Rangia, was verified by Dr. Henry

vander Schalie of the University of Michigan

after an original determination by the author.

Also, Dr. Dee Dundee of Louisiana State Uni-

versity was kind enough to send several recently

collected shells of this clam for comparative pur-

poses.

Rangia cuneata is an old species, first appear-

ing in the Miocene of North America (Dall 1898).

During the Pleistocene it occurred on the East

Coast from New Jersey to northern South

America (Richards 1938, 1939, 1962; Moore

1969). Until 20 years ago R. ctmeata was re-

stricted to the Gulf of Mexico coast of the

United States and Mexico. However, since the

late 1950's it has re-invaded its former range,

occurring presently along the East Coast from

Florida to Maryland (Hopkins and Andrews

1970).

Hopkins et al. (1973) describe R. cuneata as

the most widely distributed and by far the most

abundant species of brackish water clam in its

80 THE NAUTILUS

April 6, 1983

Vol. 97(2)

habitat. Hopkins (1970) notes that the optimum

salinity for this clam is from 1 to 15 parts per

thousand. R. cuneata cannot maintain popula-

tions outside this salinity range, thus it would

not survive in an entirely riverine habitat.

Population densities are high in many places

where this clam exists. Pfitzenmeyer (1970)

recorded a maximum density of 10,000 individ-

uals per square meter, however, this is an ex-

ceedingly high figure reflecting an extreme of

the tremendous reproductive potential of the

species. Normal densities are in the range of 4 to

30 clams per square meter (Hopkins et al. 1973).

However, since the dead shells accumulate over

the years in a productive area, the high repro-

ductive rate and large populations produce huge

piles of easily accessible dead shells. Shells are

presently so abundant they are mined with bull-

dozers, etc. for use in place of gravel in road

building as well as various industrial and water

purification processes (Hopkins et al. 1973).

Living shells certainly are not known for their

beauty. Adult R. cuneata are small, less than 4

cm long. The periostracum is fairly smooth and

grayish brown while the interior of the shell is

glossy white with a tinge of blue-gray.

Results and Discussion

During the excavation of the Arrowhead Site,

Mr. Steve Mocas (personal communication) esti-

mated that over 10,000 shells of Rangia littered

the surface at the site. However, 3,298 shell

fragments of which 1,629 were nearly complete,

were available for this report (Table 1). This lat-

ter group was also complete enough to recog-

nize the specific characteristics (Abbott 1954)

used to identify them (Fig. 1). The 1,669 unsided

TABLE 1. Animal remains from the Arrowhead Site, Ken-

Lucky.

Rangia cuneata valves 767 left, 362 right, 1669 fragments.

1 Indeterminable marine mussel 1 shell fragment.

1 Indeterminable freshwater mussel 1 shell fragment.

cf. Odocoileus virginianux, whitetail deer ■ 1 tooth

fragment.

Siui scrofa, domestic pig - 1 tooth fragment.

Canis familiaris, domestic dog - 1 right upper carnassial.

Sylvilagus floridanus, cottontail ■ 1 left tibiofibula adult.

FIG. 1. Rangia cuneata ro/res/Vom the Arrowhead Farms

site, Louisville. Kentucky. Upper I'alves are about 4 ct« in

size.

fragments could only tentatively be assigned to

R. cuneata. No other mussel species was identi-

fied with these shells, a condition not unlike that

in the natural habitat of the Common Rangia

(Hopkins et al. 1973). Only one shell fragment

was unique enough to suggest the presence of a

second marine species. None of the shells was

modified by humans in any recognizable man-

ner, thus presumably precluding their collection

for the purpose of artifact manufacture.

Since nearly all of the shells were badly

eroded and quite chalky, length measurements

were not made. Comparisons of the shells with

those sent by Dr. Dundee show the Arrowhead

Farms specimens to be within the maximum of

2V2 inches (6 cm) total length suggested for this

species by Abbott (1954). In fact, the largest ap-

pears to have been no longer than 4 cm. Most

look to be no larger than about 2.5 cm.

A limited number of other species was found

as remains associated with these shells. These

are reported in Mocas' (1976) report. A small

number of additional species are listed here in

Table 1. All of the other faunal remains are from

forms which clearly existed in the Louisville

Area throughout the time of occupation of this

site. The pig and dog remains are modern in-

trusives found on the surface.

The cultural affiliation of the shells is unfor-

tunately not possible to determine since none of

them was found in a culturally definible feature

(Mocas 1976). However, they appear to have I

Vol. 97(2)

April 6, 1983

THE NAUTILUS 81

been introduced by a Middle Woodland or later

group (Mocas 1976).

Many of the shells were present in the undis-

turbed sub-plowzone. Mocas (1976) indicates

that the shells must have been deposited in

shallow pits, most of which were subsequently

destroyed by plowing. These latter shells were

scattered across the surface of the Arrowhead

Farms site.

The only other archaeological site in which the

Common Rangia appears in the Midwest is

Cahokia. Baker (1941), Parmalee (1958) and

Chmurny (1973) report its presence. However,

these authors report only a small number of un-

modified shells and offer no insight in unravel-

ing the mystery of the Arrowhead Farms cache.

The origin of the shells is clearly somewhere

along the Atlantic or Gulf Coast. The possibility

exists that these shells were part of a collection

brought from the East Coast as trade items.

Why they ended up concentrated in the Arrow-

head Farms site still remains a mystery.

LITERATURE CITED

Abbott. R. T. 1954. American seashells. D. Van Nostrand

Co., New York, N.Y. XVII & pg. 541.

Baker, F. C. 1941. The use of MoUuscan Shells by the

Cahokia Mound Builders. Trans. Amer. Phil. Soc, 32,

part 2. 51-77 pp.

Biggs, H. E. 1970. Mollusca from human habitation sites and

the problem of ethnological interpretation. In: Science in

Archaeology, D. Brothwell and E. Higgs, 2n(i F^dition.

Praeger Pub., New York, N.Y. pp. 423-427.

Chmurny, W. W. 1973. The ecologj' of the Middle Mississip-

pian occupations of the American Bottom. Ph.D. Disser-

tation, Univ. Illinois, IJrbana, ILL.

Dall, VV. H. 1898. Contributions to the tertiary fauna of

Florida with a special reference to the Miocene silex-beds

of Tampa and the Pliocene beds of the Caloosahatchie

River including in many cases, a complete revision of the

generic groups treated of and their American species.

Trans. Wagner Free Inst. Sci. Phil., 3(4):571-947.

Hopkins, S. H. 1970. Studies on brackish water clams of the

genus Rangia in Texas. (Abstract). Proc. Nat. Shellfish

Assn.. 60:.5-6.

Hopkins, S. H. and .J. D. Andrews. 1970. Rangia cioieata

on the East Coast: thousand mile range extension, or

resurgence? Science 167:868-869.

Hopkins, S. H., J. W. Anderson, K. Horvath. 1973. The

brackish water clam R(uigia cuneata as indicator of ecolo-

gical effects of salinity changes in coastal waters. Con-

tract Report H-73-1 U.S. Army engineers waterways

experiment station, Vicksburg, Mississippi. 250 pp.

Mocas, S. T. 1976. Excavations at Arrowhead Farm (IS Jf

237). Lfnpublished manuscript. University of Louisville

Archaeological Survey, Louisville, Ky. 72 pp.

Moore, R. C, ed. 1969. Treatise on invertebrate paleontol-

ogy, Part N, Mollusca 6 Bivalvia, Vol. 2. pp. N491-N952.

Parmalee, P. W. 1958. Marine shells of Illinois Indian sites.

r;(e/V(M/i)7MS 71:132-139.

Pfitzenmeyer, H. T. 1970. Project C. Benthos. Pg. 26-38.

In: Cronin L. E.. ed. Gross physical and biological effects

of overboard spoil disposal in upper Chesapeake Bay.

LIniv. Maryland Natural Resources Inst., Spec. Rep. 3.

IV & 66 pp.

Richards, H. G. 1938. Animals of the seashore. Bruce

Humphries, Inc. Boston, Mass. 273 pp.

1939. Marine Pleistocene of Texas. Bull Biol.

Soc. Amer. 50(12-1):1885-1898.

1962. Studies in the marine Pleistocene. Trans.

Am. Philos. Soc. N.S. 52(3):1-141, 21 pis.

AUTO-DRILLING IN THE OYSTER DRILL

THAIS HAEMASTOMA (MURICIDAE)

Robert S. Prezant

Department of Biology

University of Southern Mississippi

Hattiesburg, Mississippi 39406-5018

ABSTRACT

Abnormal drilling behavior is reported in the muricid gastropod Thais

haemastoma canaliculata (Gray). A single starved oyster drill ivas observed bor-

ing into its own operculum front the pedal side. This misplaced drilling may he a

response to st:im.uli received within a mixed fauna! aquarium.

Theoyster drill, T/iais/iaCTwasioma (Linnaeus, 1758), is a well-known, predatory, boring gas-

82 THE NAUTILUS

April 6, 1983

Vol. 97(2)

tropod found throughout the Caribbean and in

the Gulf of Mexico. This gastropod may devour

up to 50% of a given oyster population in a

year's time (St. Amant, 1938) and, in higher

salinities, up to 85% (May and Bland, 1969). The

muricid's primary food is obtained by drilling a

small hole in the prey's calcareous valves using

corrosive secretions of the accessory boring

organ and mechanical raspings of the radula.

Gunter (1968) suggests that large T. haevia-

stoma bore through the shell in order to release

a paralytic substance. Once the hole is drilled in

the oyster, for instance, the snail releases a

paralytic agent through the aperture and this

causes relaxation of the bivalve's adductor

muscles. The predator then consumes the soft

parts through the gaping valves. The exact

nature of predation by this species is still in dis-

pute (Breithaupt and Dugas, 1979). Neverthe-

less, the extensible proboscis is used to ingest

the meat.

During mid-March 1982, a single specimen of

Thais haemastoma [subspecies canaliculata

(Gray, 1839) (Synonym: haysae Clench, 1927)]

from northwest Florida, 64 mm-long, initiated

an unusual type of drilling behavior. The snail,

while located alone on the wall of an aquarium,

bored a hole through its own operculum.

The snail had been placed in an artificial sea

water (26.6 ppt, 21°C), 20 gallon aquarium with

five other similarly sized snails and was fed

oysters (Crassostrea virginica) at irregular in-

tervals not exceeding four weeks between feed-

ings. In nature Thais haeynastoma feed heavily

about every three weeks (Demoran and Gunter,

1956).

After being in the tank for a period of 21

weeks and with the previous four weeks under

starvation conditions, the snail extended its pro-

boscis along the midventral region of its foot

and rasped at a small portion of the operculum

(Fig. 1). The proboscis was partially obscured

along the anterior region because of infolding of

pedal musculature. Along the posterior half of

the foot, however, the proboscis was clearly evi-

dent and the radula was seen, through the ex-

tended, translucent proboscis wall, rasping at

the operculum. Gunter (1968) reported that in T.

haemastoma, during an attack on an oyster.

FIG. 1. Proboscis of Thais haemastoma canaliculata ex-

tended between folds of foot upward to pedal side of oper-

culum. Operculum (0) shows as dark semilunar plate at tip

of proboscis. Horizontal field width = i.5 cm.

"The foot is folded around the proboscis and the

latter is never seen."

After about four and one-half hours, activity

ceased and a borehole was evident in the opercu-

lum. This hole was located 8.9 mm from the an-

terior edge of the operculum, 8.4 mm from the

posterior edge, 3.5 mm from the right edge and

5.7 mm from the left opercular edge (Fig. 2).

The hole itself has a diameter of 0.75 mm. Dur-

ing these observations the snail did not apply

the accessory boring organ (ABO) to the drilled

region.

Within five months after the event, the hole

had been transferred to the very edge of the

operculum as natural opercular growth dis-

placed it. At this date (eight months later), the

snail still feeds and shows no further aberrant

behavior nor any obvious indication of physi-

ological distress.

Carriker et al. (1978) and Carriker and

Williams (1978) suspect that secretions from the

accessory boring organ, of at least Urosalpinx

cinerea, are involved in the breakdown of calci-

fied shell and organic matrix in molluscan prey.

Jensen (1951) reports that some naticid gastro-

pods can bore into skate egg cases. Thus, the

Vol. 97(2)

April 6, 1983

THE NAUTILUS 83

FIG. 2. Diagram of the operculum of Thais haemastoma

eanaliculata shoudng site and size of auto-driUed hole.

Horizontal field width = 9.5 mm.

ABO is likely responsible for at least partial

penetration of partly and fully organic sub-

strates. Drilling of a poorly calcified, proteina-

ceous operculum without use of the ABO is like-

ly to be relatively "easy" for a large oyster drill.

The well-developed radula, used consistently,

penetrated the thin horny operculum within five

hours. Carriker (Pers. Comm.) has supportive

evidence for the use of only the radula in some

events of drilling. This support is based on

observations of raspings by U. cinerea on the

periostracum of Mytilus edulis. The relatively

thin, noncalcified operculum of T. haemastoma

may offer little resistance to purely mechanical

boring.

The significance of this anomolous activity is

uncertain. No other snails starved for the same

duration showed any signs of aberrant drilling.

In one instance, another snail did extend its pro-

boscis partly within a fold of its foot but did not

attempt to rasp or drill the operculum. It is

possible that starved drills may periodically ex-

tend their proboscises in response to chemical

stimuli. Gunter (1968) reports that hungry T.

haemastoma will evert their proboscis in

response to oyster flesh stimulus. Carriker and

Yochelson (1968) and Hancock (1959) report

that in laboratory populations of drills (both

Urosalpinx and Eupleura) kept in tanks with

both dead (empty valves) and live oysters, the

snails would occasionally drill into an unoc-

cupied shell. It has been suggested that the snail

could not distinguish live from dead because of

high concentrations of dispersed chemoattrac-

tants in the tank (Carriker and Yochelson,

1968). The possibility exists that a chemical

stimulus within the aquarium was detected by

our specimen. The aquarium at the time was

also occupied by several hermit crabs [Clibana-

rius vittatus (Bosc, 1801)] and one large Dolly

Vardin crab [Hepatus epheliticits (Linnaeus,

1763)]. The proboscis of the drill may have ex-

tended in response to some metabolite released

by one of these organisms or perhaps by another

snail. The tip of the probing proboscis may have

encountered the operculum and "mistaken" this

for the shell of potential prey. Recent and fossil

boreholes of Thais haeinastoma Jloridana (Con-

rad, 1837) reported by Carriker and Yochelson

(1968) from calcified oyster valves had dia-

meters greater than 1.4 mm, twice that found in

the operculum. In the former cases the borehole

reflects the size and shape of the ABO. The

small size of the hole drilled in the operculum

may be a result of the animal becoming "aware"

of the fact that it was not drilling into food, and

thus terminating the purely radulate drilling.

It is unlikely that the exact cause of this

behavior will be uncovered; however, it is impor-

tant to note that aberrant proboscidal activity

and drill behavior may be a i-esult of keeping

thaids in laboratory conditions in mixed faunal

aquaria.

Acknowledgments

I am grateful to Drs. R. Tucker Abbott and M.

R. Carriker for reviewing and improving the

manuscript, and Mrs. E. Henderson for her

careful secretarial assistance in preparation of

the manuscript.

LITERATURE CITED

Breithaupt, R. L. and R. J. Dugas. 1979. A study of the

southern oyster drill (Thais haeviastoma) distributions and

density on the oyster seed grounds. Louisiana Wildlf

Fish. Comm. Tech. Bull. 30: 20 pp.

Carriker, M. R., D. Van Zandt and T. J. Grant. 1978. Pene-

tration of molluscan and non-molluscan minerals by the

boring gastropod Urosalpinx cinerea. Biol. Bull.

155:511-526.

Carriker, M. R. and L. G. Williams. 1978. The chemical

mechanisms of shell dissolution by predatory boring

gastropods: a review and an hypothesis. Malacologia

17:143-156.

Carriker. M. R. and E. L. Yochelson. 1968. Recent gastro-

84 THE NAUTILUS

April 6, 1983

Vol. 97(2)

pod boreholes and Ordovician cylindrical borings. Geol.

Sun>ey Prof. Paper 593-B:l-26, .5 plates.

Demoran, W. J. and G. Gunter. 19.56. Ability of Thais

haemastoma to regenerate its drilling mechanism. Scievce

123:1126.

Gunter, G. 1968. Some factors concerning the drilling

apparatus and the feeding and predation of Prosobran-

chiate gastropods especially on other molluscs. Mar. Biol.

Assoc. India, 1968. Proc. Symp. Mollusca, Pt. 1:370-378.

Jensen, A. S. 19.51. Do the Naticidae (Gastropods Proso-

branchia) drill by chemical or by mechanical means?

Vidensk. Medd. Dan. Naturhist. Form. 113:251-261.

May, E. B. and D. G. Bland. 1969. Survival of young oysters

in areas of different salinity in Mobile Bay. Proc. 23rd

Ann. Conf. Southeast. Assoc. Game Fish Co?n?N.. -519-521.

St. Amant, L. 1938. Studies on the distribution of the

Louisiana oyster drill, Thais floridana haysae Clench.

Master's Thesis, Louisiana State University, Baton Rouge.

116 pp.

RECENT DEATH

Wendel Phillips Woodring, paleontologist,

died January 29, 1983, in Santa Barbara, Cali-

fornia, at age 92. Born in Reading, PA, June 13,

1891, he obtained his Ph.D. at Johns Hopkins

University in 1916. He did extensive field work

in the West Indies and Central America while

employed by the U. S. Geological Survey. His

first outstanding publication was on "The

Miocene Mollusks from Bowden, Jamaica," but

throughout his 67 productive years of research

he became the leading authority on the taxon-

omy and stratigraphy of the Tertiary mollusks

of the tropical New World. He was a past-

President of the Paleontological Society of

America and the Geological Society of America,

as well as receiving many honors and medals for

his work. He is survived by his wife. Merle

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American Malacnlogist^, first edition, 1973,

p. 487 and Who'a w'ho.

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Vol. 97(2)

April 6, 1983

THE NAUTILUS 85

KATHERINE V. W. PALMER - AN OBITUARY

1895 - 1982

On September 12, 1982 death brought to an

end the distinguished career of Dr. Katherine V.

W. Palmer, one of America's foremost paleon-

tologists. Katherine Evangeline Hilton Van

Winkle was born February 4, 1895, in Oakville,

Washington, the daughter of Dr. Jacob Out-

water Van Winkle and Edith Hilton Van

Winkle. After high school she enrolled in the

University of Washington where she came

under the influence of Dr. Charles E. Weaver

who urged her to work in paleontology. An ex-

cellent student, she published her first scientific

paper before graduation. Having completed re-

quirements for graduation at the University of

Washington, she spent the last two quarters of

her senior year as Assistant in Geology at the

University of Oregon. After receiving her B. S.

in 1918 she went to study under Professor

Gilbert D. Harris at Cornell where she was

Goldwin Smith Fellow in Geology for 1918-19

and 1919-20. Her plans to return to Washington

to work with Dr. Weaver changed after she

married Dr. Ephraim Laurence Palmer on

December 24, 1921. Dr. Palmer, a professor at

Cornell who later became an eminent naturalist,

encouraged his wife in her studies. She received

her Ph.D. from Cornell in 1925.

Marriage and the subsequent birth of two sons

did not stop Dr. Katherine Palmer's paleon-

tological research. She continued to work with

Professor Harris who had his own printing

press. He and his students set type, made plates,

ran the press and folded signatures. From this

modest beginning came the Paleontological

Research Institution, with Katherine Palmer as

one of the Founding Members in 1932. After the

death of Professor Harris in 1951 she became

Director of the Paleontological Research Insti-

tution and served in that position until her

retirement in 1978. The story of this unique and

important institution is told in her last publica-

tion which details the organization and func-

tions of the P.R.I, during its first fifty years.

Katherine Palmer was active and admired in

many circles. In 1917 she was Chapter Presi-

dent (and Charter President) of Alpha Delta Pi,

an undergraduate social sorority. The lead in its

publication. The Adelphian, of April, 1920, a

two-page article plus a fetching photograph,

described her as "all-around", "wonderfully

good-natured", with a sense of humor and "a

droll way that every one loves." This assessment

of her disposition and wit remained valid

throughout her life. Her popularity with one and

all is reflected in many newspaper articles, one

in particular referring to the Palmers as one of

only two couples of whom both were listed in

Who':> Who; "The other couple was Charles and

Anne Morrow Lindbergh. Quite an honor for the

Lindberghs, wasn't it?"

She was also a member of Sigma Xi; Phi Kap-

pa Phi; a co-founder of Sigma Delta Epsilon, a

women's graduate scientific organization of

which she was later National President; a

Charter Member and President of the Cornell

86 THE NAUTILUS

April 6, 1983

Vol. 97(2)

Chapter of the women's geological organization,

Chi Upsilon.

Katharine Palmer was a member, either ac-

tive or honorary, of many scientific organiza-

tions. Among these were the American Mala-

cological Union, of which she was a Life Member

and President in 1959-60; she was a Fellow of

the Geological Society of America; a Fellow of

the American Association for the Advancement

of Science; a member of the Society of Systema-

tic Zoology; the American Association of Petro-

leum Geologists; the Society of Economic Pale-

ontologists and Mineralogists; a Life Member of

both the Geological Society of France and the

Societe Linneenne du Lyon; and a Fellow of the

Paleontological Society of America, which

presented her with its prestigious Paleontolo-

gical Society Medal in 1972.

As the seventh recipient of the Paleontolo-

gical Society Medal, Katherine Palmer estab-

lished a number of 'firsts': The first recipient to

have worked with mollusks, the first institu-

tional administrator to receive the award, and is

still the only woman to have been so honored.

On presenting the Medal to her, Dr. Kenneth E.

Caster referred to "the Harrisian kind of paleon-

tological instruction . . . where all students were

treated as 'zealous companions in research'".

Whether inate or acquired, treatment of every

interested person as a companion in research

was characteristic of Katherine Palmer. Al-

though always engaged in her own research and

the affairs of the P.R.L she gave freely of her

time and advice, encouraging amateurs, stu-

dents and fellow scientists alike. She made all

feel that what they were doing was important.

She especially enjoyed helping build up their

libraries and was always alert for titles needed

by various workers.

The excellent library of the P.R.L and her own

impressive library served her well, making it

possible to work at home or the Institution.

Recognizing the importance of the literature

and bibliographic detail she continued reprint-

ing unavailable, old and rare publications by the

P.R.L

Her interests were many and varied. Her

work with Tertiary mollusks inevitably led to

work with Recent mollusks, and her search for

completeness of detail in her taxonomic publica-

tions led to additional publications in biography

and bibliography. She had a strong interest in

systematic nomenclature and attended the

Copenhagen Colloquium on Zoological Nomen-

clature in 1953 and the London Colloquium in

1958. She was an active participant in the delib-

erations at both of these important conferences.

In 1978 Tuiane University conferred upon

Katherine Palmer an honorary Doctor of

Science degree during a symposium that was

held in her honor. The citation for the honorary

degree stated in part, ". . . all the praise and

recognition she has received could not be

enough to match the extent of her contribution

to Tertiary paleontology. The brilliance of her

work, the precision of her method, the wisdom

of her leadership of the Paleontological

Research Institution will stand as the greatest

monument to Dr. Palmer and as the highest ex-

ample for future generations of what it means to

be a scientist."

In her acceptance of the Paleontological

Society Medal, she commented that "our ideas

have not come spontaneously, but were stimu-

lated and nurtured by the tomes of the giants

before us . . ." Katherine Palmer's name must

now be added to this list of giants who have left

a legacy of solid scientific research to be utilized

and expanded by future generations. She pub-

lished over 70 books and papers. A complete

bibliography appears immediately following her

last scientific publication in Tuiane Studies in

Geology and Paleontology (1979, i5(l-4):74, 94,

104, 128). In May of 1982 she completed what

was to be her final paper, a history of the first

fifty years of the Paleontological Research In-

stitution which has now been published by that

Institution.

Katherine Palmer will continue to be known

through her work by future generations of

paleontologists and malacologists. To those of

us who knew her, she will be remembered not

only for those contributions, but also for being

the special person she was. Dr. Katherine V. W.

Palmer is survived by one son, Richard Robin

Palmer.

Richard E. Petit

North Myrtle Beach

South Carolina

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JULY 29, 1983

THE

NAUTILUS

ISSN 0028-1344

Vol. 97

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Dr. Joseph Rosewater

Division of Mollusks

U.S. National Museum

Washington, D.C. 20560

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Department of Invertebrates

Field Museum of Natural History

Chicago, Illinois 60605

Dr. Aurele La Rocque

Department of Geology

The Ohio State University

Columbus, Ohio 43210

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Museum of Zoology

The Ohio State University

Columbus, Ohio 43210

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Los Angeles County Museum of Natural History

900 Exposition Boulevard

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Department of Mollusks

Museum of Comparative Zoology

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THE

NAUTILUS

Volume 97, number 3 - July 29, 1983

ISSN 0028-1344

CONTENTS

Nancy E. Brandauer

Malacological Collection of H. P. Mera at the University of Colorado Museum 89

Winston F. Ponder

Reclassification of Some American Species Assigned to the Rissoidae (sensu lato) 90

James D. Lazell, Jr.

Rediscovery of the Palm Snail, Hemitrochus nemoralinus intensus Pilsbry (Cepolinidae) 91

William Miller, III

Survey of the Pyramidellid Gastropods in the Wassaw Sound Area, Coastal Georgia 93

Robert Robertson

Observations on the Life History of the Wentletrap Epitonium echinaticostum

in the Bahamas 98

Bretton W. Kent

Diet Expansion of Busycon contrarium in the Absence of Triplofusus giganteus

(Gastropoda: Buccinacea) 103

K. Elaine Hoagland

Ecology and Larval Development of Crepidula protea (Prosobranchia: Crepidulidae)

from Southern Brasil: A New Type of Egg Capsule for the Genus 105

Ralph W. Taylor

The Freshwater Naiad (Mussel) Fauna of the Nolin River in the Green River Drainage

of Central Kentucky (Mollusca: Bivalvia) 109

Jack R. Davis

An Additional Record of Living Orygoceras (Hydrobiidae) from Texas 112

Glenn A. Long

The Unionids (Bivalvia) of Loch Raven Reservoir, Maryland 114

Robert Robertson

Axial Shell Rib Counts as Systematic Characters in Epitonium 116

Deaths 113 Zoological Record Up-date 88

88 THE NAUTILUS

July 29, 1983

Vol. 97(3)

Zoological Record UP-DATE

MOLLUSCA Section 9

Now printed and distributed in the United

States, and under a new speed-up project aided

by the non-profit organization, BIOSIS.

The Mollusca Section, (vol. 117) so essential to

all research workers, now covers the 1980 litera-

ture. IV2 volumes will be issued during each of

the next three years. Thus by 1987 the Zoolog-

ical Record will be covering the very latest

molluscan literature. It is something every

mollusk research department should have.

Order vol. 117 before September 30, 1983, and

receive a 5% discount. Send $120.00 to BIOSIS

Customer Services, 2100 Arch Street, Phila-

delphia, PA 19103-1399. Computer diskettes or

tapes available for years 1978-80. Coming soon:

up-dated to 1984, with monthly up-dates.

Search on your home computer. Write for free

Zoological Record Online brochure.

NEWS

PUBLICATIONS RECEIVED

Inaba, Akihiko. 1982. Molluscan Fauna of the Inland Sea,

Japan. Hiroshima Shell Club. 180 pp., 4 pis. Paperback.

An annotated checklist, containing geographical and hab-

itudinal information on 1005 marine mollusks in the Seto

Inland Sea. Illustrations of 72 uncommon species. A useful

guide to one of Japan's richest collecting areas between

Honshu and Shikoku Islands. Send International Postal

Money order for U.S. $13.00 to Hiroshima Kairui Danwa-

kai (Hiroshima Shell Club), c7o Mukaishinia Marine Biol.

Station, Onomichi P. 0., Hiroshima Pref., .Japan 722.

.JECOLN Annual Reports (Japanese Expert Consultation on

Living Nautilus, Kanagawa). 1979-1982. 56 collected

reprints on the Nautilus, maily by Japanese authors.

Covers years 1977-78; 1978-79; 1979-82. ISSN 0396-

054X. Yokosuka City Museum, Fukadadai, Yokosuka 238,

Japan.

Bogan, Arthur E. and Paul W. Parmalee, 1983. The Mol-

lusks in Vol. 2 of Tennessee's Rare Wildlife. 123 pp. Spiral

bound. Tenn. Wildlife Resources Agency, Nashville, TN

37204. Names, illustrations, maps, descriptions, synony-

mies, distributions, and natural history information on 33

bivalves and 20 gastropods considered to be rare or en-

dangered.

Garcia - Cubas, Antonio. 1981. Moluscos de un Sistema

Lagunar Tropical en el sur del Golfo de Mexico (Laguna de

Terminos, Campeche). Special Publ. no. 5, Inst. Cienc. del

Mar y Limnol. Univ. Nal. Auton. Mexico. 182 pp. 175

species described, illustrated and ecologically discussed.

A.M.U. SEATTLE MEETING

American Malacological Union's 49th Annual

Meeting, Aug. 7-13, 1983, at the University of

Washington in Seattle. Symposia on "Molluscan

Nerve Cells," "Molluscan Extinctions" and

"Avian Molluscivores." Papers on Cephalopods,

Pacific Northwest Molluscs and recent studies.

Workshops, field trips. For information write

Alan J. Kohn, Dept. of Zoology, Univ. of Wash-

ington, Seattle, WA 98195 or Tel. (206) 543-

1629. The Western Society of Malacologists

will have a joint program and auction with the

A.M.U.

C.O.A. FLORIDA MEETING

The Conchologists of America will hold their

annual meeting in Sarasota, Florida, Sept. 21

through 25. For details see the C.O.A. June

Bulletin, or contact Dick Forbush, 1104 Sklar

Drive East, Venice, FL 33595.

Specimen Shells

Offering microscopic and miniature (to '/t inch) shells

from the Florida Keys, with accurate locality data Also

unsorted grunge; write for list.

Margaret Teskey

P.O. Box 273

Big Pine Key, Fl. 3301,3

SHELL SHOWS 1983

Jacksonville (Florida) Shell Show. July 29-31,

1983.

Midwest Regional Shell Show (Indianapolis).

Aug. 13-14, 1983.

Philadelphia Shell Show. Oct. 1-2. 1983.

Astronaut Trail Shell Show. (Merritt Id., FL).

Nov. 4-6, 1983.

Vol. 97(3)

July 29, 1983

THE NAUTILUS 89

MALACOLOGICAL COLLECTION OF H. P. MERA AT THE UNIVERSITY

OF COLORADO MUSEUM

Dr. Harry P. Mera's collection of land and

fresh water molluscan shells, which was donated

to the University of Colorado Museum in the

early 1950's, was rediscovered here in the spring

of 1980 by Dr. Dwight W. Taylor, who was

visiting the Museum. The University had been

without a curator in malacology for 40 years un-

til the arrival of Dr. Shi-Kuei Wu in 1973, and so

much work remained to be done that the Mera

Collection had not yet been recognized for what

it was.

Dr. Mera, a practising physician in Abilene,

Kansas, began collecting shells in about 1912

and continued for at least 15 years. According

to an obituary and bibliography published in

American Antiquity (Vol. 17, July 1951), he

moved to New Mexico in 1922 and later became

widely known as an anthropologist and staff ar-

chaeologist at the Laboratory of Anthropology

in Santa Fe, where he specialized in the artifacts

of southwestern Indians. His bibliography con-

tains no malacological publications.

He supplemented his shell collecting efforts in

Kansas and New Mexico with extensive pur-

chases from W. F. Webb and Ward's Natural

Science Establishment in Rochester, New York,

and from the dealers Sowerby and Fulton in

London and Geret in Paris. His collection also

contains shells from the collections of Bryant

Walker, Junius Henderson and Victor Sterki,

who all helped Dr. Mera with identification

problems, and from F. C. Ackers, C. M. F.

Ancey, Miss Bowman, H. C. Brooks, J. Y.

Crans, Lorenzo E. Daniels, Joseph L. Goodell,

E. E. Hand, I. B. Hardy, Esther Haskell, Ethel

Haynes, Anson A. Hinkley, Yoichiro Hirase,

Florence Ives, Julia Ives, P. B. James, Norman

W. Lermond, Herbert N. Lowe, F. E. Mera,

Mrs. H. P. Mera Sr., Amanda Nixon, Mary

Olney, Charles R. Orcutt, G. W. Pepper, J. F.

Quadras, H. E. Sargent, E. R. Schowalter,

Charles T. Simpson, John H. Thompson,

Charles Townsend, R. H. Turver, Robert

Walton and Edna Worley.

The collection has now been put into useful

order. There are 1380 lots; 947 are land snails

and 433 are fresh water mollusks. Numerous

paratypes are included. The filing system is both

numerical and alphabetical by genus. The land

snails have a worldwide distribution and include

representatives of over 90 genera. Nearly half

the specimens are island forms, and two thirds

of those are from islands in the Pacific. The

fresh water lots, which include 30 genera, are

dominated by Pisidium, Sphaerium and Muscu-

lium, many of which came from Bryant Walker.

In addition to Dr. Mera's private collection,

which is housed separately at the University of

Colorado Museum, the Museum collection con-

tains 139 lots of shells which were sent by Dr.

Mera to Junius Henderson, the original curator,

between 1912 and 1921.

The collection is all dry material in good

physical condition and predominantly with com-

plete data. Specimens are sometimes identified

only to the genus level. The nomenclature is

often years out of date, but it is hoped that this

would not be an overwhelming obstacle to an ex-

perienced worker.

The author wishes to acknowledge the helpful

comments and suggestions of Dr. Dwight W.

Taylor and Dr. Shi-Kuei Wu in the preparation

of this manuscript.

Nancy E. Brandauer

University of Colorado Museum

Campus Box 218

Boulder, Colorado 80309

90 THE NAUTILUS

July 29, 1983

Vol. 97(3)

RECLASSIFICATION OF SOME AMERICAN SPECIES ASSIGNED

TO THE RISSOIDAE (SENSU LATO)

Winston F. Ponder

The Australian Museum

College Street

Sydney, N.S.W. Australia 2000

ABSTRACT

Twenty five species oftaxa previously assigned to the Rissoidae fsensu latoj are,

from examination of their type material, considered to belong to other families.

During the examination of type material of

the Rissoidae in several museums in the U.S.A.

it was found that a number of species described

as rissoids had been wrongly assigned to the

family. As most of these are still included in the

Rissoidae (Abbott, 1974; Keen, 1971), a list of

the species, together with an assessment of the

family to which they properly belong is given

below.

Abbreviations - ANSP- Academy of Natural

Sciences of Philadelphia; CAS-California

Academy of Sciences, San Francisco; MCZ-

Museum of Comparative Zoology, Cambridge,

Mass.; USNM-National Museum of Natural

History, Washington, D.C.

List of Non-rissoid Taxa

apicina Verrill, 1S84, Ali'nnia. Southeast of Nantucket,

1608 fathoms. Epitoniidae (holotype, USNM) (also noted

by Waren (1974)) who says that it is close to, if not identi-

cal with Epitontu7n frwlei {DaW. 1889).

athymorhyssa Dall, 1892, Rissoa. Pliocene, Florida. Hydro-

biidae (holotype, USNM).

berryi Baker, Hanna & Strong, 1930, Rissuina. Cape San

Lucas, Baja California. ?Epitoniidae (holotype, CAS). This

species is based on a juvenile specimen with a damaged

aperture.

catlistrophia Dall, 1892, Rissoa (Onoba). Pliocene, Florida.

Hydrobiidae (holotype, USNM).

campid Dall, 1927, Rissoa (Cinyula). Off Georgia. Family?,

possibly Fossariidae or Epitoniidae. The shell has a pauci-

spiral protoconch, weak a.xial sculpture and no varices

(type, USNM).

conica C. B. Adams, 1850, CinguLaJ('!). Jamaica. Cerithiidae

(holotype, MCZ). Figured by Clench & Turner (1950)

without comment. The holotype is a worn, juvenile

Ceriihium (s.l).

curta Dall, 1927, Rissoa (Cingulina). Off Fernandina,

Florida, 294 fathoms. Skeneidae (syntypes, USNM). Pre-

occupied by Rissoa curta Dujardin, 1837.

eulimoides C. B. Adams, 1850, Rissoa. Jamaica. Eulimidae,

as shown by Lyons (1977).

femandinae Dall, 1927, Rissoa (Nodulus). Off Fernandina,

Florida, 294 fathoms. Family?, possibly Skeneidae (syn-

types, USNM).

fragilis Wade, 1926, Rissoina. Upper Cretaceous, Tennes-

see. Eulimidae (holotype, USNM). Sohl (1960) has already

pointed out that this species is "melanellid in character".

gallegosi Baker, Hanna & Strong, 1930, Alvania. Cape St.

Lucas, Baja California. Cerithiidae (holotype, CAS).

infrequens C. B. Adams, 1852, Rissoa. Panama. Epitoniidae

(holotype, MCZ). Bartsch (1915) placed this species in

Pliciscala but Keen (1971) rejected this location in favour

of Rissoina. Examination of the holotype supports the

placement of this species in the Epitoniidae.

Ificunatus Carpenter, 1865, .Amphithalamus. California.

Family? (holotype, USNM). The type is a broken juvenile

and could belong in one of several families. This species

should be regarded as a nomen dubium. It is certainly

not an Amphithalamiis.

knnpra Dall, 1927, Rissoa. Off Fernandina, Florida, 294

fathoms. Fossariidae (Couthouyia - Zeradina group).

Preoccupied by /? . lampra Suter, 1908. R. lampra Dall is,

however, very similar to R. saridfrsoni Verrill and may l)e

conspecific so a replacement name is not provided.

mayori Dall, 1927, Rissoina. Off Georgia & Miami. Epitoni-

idae (type, USNM).

microcharia Dall, 1892, Rissoa (Onoba). Pliocene, Florida.

Pyramidellidae (holotype, USNM).

pom.pholyx Dall, 1927, Rissoa. Off Georgia. Fossariidae

(Couthouyia - Zeradina group) (type, USNM).

portoricana Dall & Simpson, 1901, Rissoa. Mayaguez Har-

bor, Puerto Rico. Cerithiidae (Finella (Caloosalaba))

(holotype, USNM),

sandersoni Verrill, 1884, (yingula. Off Cape Hatteras, North

Carolina, 142 fathoms. Fossariidae (? syntypes, USNM,

one lot ANSP). Waren (1974) states from examination of

the lot in the USNM that they are Rissoellidae or, possibly

Aclididae. They appear to me to belong to the Couthouyia -

Zeradina group in the Fossariidae. R. lampra Dall is

very similar.

Vol. 97(3)

July 29, 1983

THE NAUTILUS 91

solida C. B. Adams, 1850, Cingvia (?). Jamaica. Trochidae

('! = Hiili^tylus pupoides (Carpenter, 1864)) (holotype,

MCZ). Fibred by Clench & Turner (UtSO) without com-

ment. This is a juvenile shell with a broken aperture that

matches specimens of the West Coast H. pupoides rather

well, indicating that the original locality is probably in

error.

stephe)is<u' Baker, Hanna & Strong, 1930. Rissoina. Cape

St. Lucas, Baja California. Cerithiidae (probably Bittivm)

(holotype, CAS).

stewardsnni Vanatta, 1909, Rissoa (Nodulus). Fairyland,

near Hamilton, Bermuda. Hydrobiidae (holotype and para-

types, ANSP).

subomata Wade, 1926, Rissoiyia. Upper Cretaceous, Ten-

nessee. Fossariidae (Couthouyia - Zeradina group) (holo-

type, USNM).

texana Stanton, 1947, Rissoa! Lower Cretaceous, Texas.

Family?, possibly Cerithiidae or Trichotropidae (types,

USNM).

toroensis Olsson & McGinty, 1958. Rissoa. Boeas Islanil,

N.E. Panama. Pyramidellidae (holotype and paratypes,

ANSP).

Acknowledgments

I would like to thank Drs. G. M. Davis and R.

Robertson (ANSP), Dr. B. Roth (CAS), Prof. R.

D. Turner (MCZ) and Drs. R. S. Houbrick and J.

Rosewater (USNM) for gfiving me access to the

type collections in their care and the facilities

with which to study them.

LITERATURE CITED

Abbott, R. T. 1974. American seasheHs. The marine Mol-

lusca of the AtUmtic and Pacific coasts of North America.

Van Nostrand Reinhold, New York.

Bartsch, P. 1915. The Recent and fossil mollusks of genus

Rissoina from the west coast of America. Proc. U.S.

Natn. M«.s-. 49:33-62.

Clench, W. and R. D. Turner. 1950. The Western Atlantic

marine MoUusca described by C. B. Adams. Occ. Pap.

Mollusks. Harvard l(15):233-404.

Keen, A. M. 1971. Sea shells of tropical West America,

Marine mallusks from Baja California to Peru. Stan-

ford LIniv. Press. Stanford.

Lyons, W. G. 1977. Comments on three Jamaican melanel-

lid species described by C. B. Adams (Gastropoda: Mela-

nellidae). Occ. Pap. Mollusks. Harvard 4(55):149-157.

Sohl, N. F. 1960. Archeogastropoda, Mesogastropoda and

stratigraphy of the Ripley, Owl Creek, and Prairie Bluff

Formations. U.S. Geol, Survey Prof Pap. 331-A:1-151,

18 pis.

Waren, A. 1974. Revision of the Arctic-Atlantic Rissoidae

(Gastropoda, Prosobranchia). Zoologica Scripta

3:121-135.

REDISCOVERY OF THE PALM SNAIL, HEMITROCHUS

NEMORALINUS INTENSUS PILSBRY (CEPOLINAE)

James D. Lazell, Jr.

8 Swinburne Street

Conanicut Island, RI 02835

During a biological reconnaissance of Guana

Island, just north of Tortola in the British Virgin

Islands, a small, arboreal snail was found asso-

ciated with the palm, Thrinax morrisii. The

snail appears to be H. n. intensus Pilsbry, 1889,

described without type-locality and apparently

not mentioned in the literature since. In life, the

shell has a lightly striped, axial pattern; the shell

has shades of brown, from ivory or ochraceous

buff to russet. The soft parts are pinkish grey. A

dozen specimens were preserved in ethanol and

deposited in Florida State Museum, Gainesville

(UF 40044).

Although empty shells of H. n, intensus were

occasionally found at several points on Guana

Island in association with scattered Thrinax

palms, the concentration of both snails and

palms is in the moist ravine on the northern side

of the island at ca 30-100 m (Fig. 2). This was

during the dry season, from February to April,

92 THE NAUTILUS

July 29, 1983

Vol. 97(3)

FIG. 2. Guana Island in the British Virgins. Stippling indi-

cates the zone of palms. Thrinax morrisii, in which H. n. in-

tensus was concentrated in March, 1982. Dots indicate posi-

tions of empty shells. Contours are 100 m. Inset shows Puer-

to Rico (PR) and the Virgin Islands. Arrow indicates the

po.sition of Guana Island (G). Fine line is the approximate

100 m submarine contour, proljable maximum land edge dur-

ing Wurm glaciation.

when I was present on the island. At this time

there were only a few, small, scattered pools of

fresh water in the ravine. No other ravine on the

island retained any standing fresh water in

March, 1982.

The palm itself is described as "unusual" in the

Virgin Islands by D'Arcy (1971), although Read

(in Howard, 1979) indicates these islands are

well-within the general distribution. Although I

have done extensive survey work in the British

Virgins (Lazell, 1980), I have not found stands of

Thrinax morrisii elsewhere. It does occur wide-

ly as single, isolated individuals, and in small

stands on Anegada (D'Arcy, 1971). The ravine

stands on Guana form a dense undercover domi-

nated by the plants, Bursera simaruba, Pimnia

subcordata, and Hippomane mancinella. The

palm is widely harvested in the Virgin Islands to

make brooms and is often called "broom palm"

or "broom tyre."

The biological relationship of//, n. inteyisus to

nominate nemoraiinus Petit of Puerto Rico

should be investigated. Moisture-dependent ver-

tebrates, such as frogs of the genus Eleuthero-

dactylus, fossorial reptiles such as Ampkisbaima

FKI. 1- liiuiiiii palm siiiiil. Heniitrochus

nemoralinus intensus Pikbry, 1889, from

Guana Island in the British Virgins. Photo by

Robert Ginsberg.

and Typhlops, and some terrestrial geckos of the

genus Sphaerodactylus, show greater tenden-

cies for speciation in the Virgin Islands than do

more xeric-adapted forms (Lazell, 1983). This is

presumably true because, even during the

height of the Wiirm glacial maximum when the

land areas were united, moist areas in the

eastern, low portions of the Puerto Rico Bank

(now the Virgin Islands) were still isolated by in-

tervening xerophytic regions.

I am indebted to Dr. Fred G. Thompson,

Florida State Museum, for identification of the

snail, and to Dr. Robert W. Read, U.S. National

Museum, for identification of the palm. I was

assisted in the field by Gerald Durrell, Dr. Lee

Durrell, Robert Ginsberg (who provided Fig. 1),

Jan Soderquist, and members of the Jarecki

family -owners of Guana Island. I am deeply

grateful to the Jareckis, and Mary Randall and

Albert Penn, who manage the island, for their

help and hospitality.

LITERATURE CITED

D'Arcy, W. G. 1971. The mystery Sabiil of Anegada. Prin-

cipes, J. Palm Soc. 15(4):l"31-133.

Howard, R. A. 1979. Flora of the Lesser Antilles Leeward

and Windward Islands 3. Arnold Arboretum, Harvard

Univ., Jamaica Plain, Mass.

Lazell, J. D. 1980. Report: British Virgin Islands, 1980.

Privately printed, Jamestown, RI.

1982. Biogeography of the herpetofauna of the

British Virgin Islands with description of a new Anolis

(Sauria, Iguanidae). Spec. Publ. Mus. Comp. Zool., in

press.

Pilsbry, H. A. 1889. Manual of Conchology, Ser. 2, vol. .5;22;

pi. 31, figs. 6, 7.

Vol. 97(3)

July 29, 1983

THE NAUTILUS 93

SURVEY OF THE PYRAMIDELLID GASTROPODS IN THE

WASSAW SOUND AREA, COASTAL GEORGIA

William Miller, III

Department of Geology

Tulane University

New Orleans, LA 70118

ABSTRACT

Four taxonomic associations of pyramidelli.d snails were identified in the

Wilmington River-Wassaw Sound estuary and on the adjacent continental shelf,

including: 1) an estuarine margin association, consisting mostly of Odostomia

impressa (Say) occurring in and around intertidal oyster banks; 2) an offshore

estuarine association, with sparse Turbonilla interrupta (Totten) occurring in

subtidal areas with abundant Mulinia lateralis (Say); 3) an open marine associa-

tion, characterized by a few T. incisa incisa Bush also associated the valves o/M.

lateralis; and Jt) a mixture of odostomes, turbonills. and pyrams derived from.

several stratigra^hic horizons and original environments, and associated with a

species-rich assemblage of fossil mollusks occurring for the most part within tidal

channels.

The purpose of this paper is to report the com-

position of the pyramidellid snail fauna of the

Wilmington River-Wassaw Sound estuary and

adjacent shelf, and to document the distribution

of the fauna in the estuarine and nearshore

marine environments of the area. An unexpect-

edly taxonomically rich fauna was discovered,

particularly where fossil and recent shells had

been mixed within tidal channels; high abun-

dance of individuals, however, was attained only

by one species. Members of the Pyramidellidae

inhabiting the environmental mosaic of the

Wassaw Sound area could be readily segregated

into four taxonomic associations (terminology of

Kauffman and Scott, 1976), and the distribu-

tional and paleontologic aspects of pyramidellid

species will be discussed in terms of these recur-

rent groups of taxa.

Methods

During the summer of 1981, fifteen bulk sam-

ples of shelly sediments were collected from a

variety of coastal environments, wet-sieved to

separate shells from matrix, and carefully ex-

amined for pyramidellid shells. The sample sta-

tions are shown in Fig. 1. Samples labelled with

KJ were collected in deeper subtidal locations

using a Reineck box corer deployed from the

deck of the R/V Kit Jones. Samples labelled with

W and H-1 through H-4 were collected at low

tide from intertidal locations, and H-8 was col-

lected in shallow water with a bottom grab-

sampler deployed from a small whaler. Six ma-

jor types of estuarine and nearshore marine en-

FIG. 1. Index map showing location of study area in north-

em coastal Georgia (inset) and map of the Wassaw Sound

area showing location of sample stations.

94 THE NAUTILUS

July 29, 1983

Vol. 97(3)

vironments were sampled, including: 1) seaward

flank of the ebb tidal shoal located at the mouth

of the sound; 2) surface of the ebb tidal shoal;

3) the tidal inlet floor; 4) estuarine channel bot-

tom within the sound; 5) the flank of the estu-

arine channel; and 6) intertidal oyster banks or

reefs along the margins of the sound and lower

stretches of the river (Table 1, Fig. 1).

All samples were sieved, dried, and hand-

picked to remove all pyramidellids larger than

about 1 mm in largest dimension. Published de-

scriptions and illustrations were used to identify

specimens (Maryland Geological Survey, 1904;

Bartsch, 1909; Henderson and Bartsch, 1914;

Morris, 1973; Abbott, 1974). All specific deter-

minations were made using a dissecting micro-

scope and all identifications were double-

checked at least once. Other mollusks co-occur-

ring with the pyramidellids in large numbers

were also identified.

TABLE 1. Environmental setting, water depth, and volume

of samples before processing. Sample numbers are keyed to

Fig. 1.

- - collected just .-ibove mean low fide level

Taxonomic Practice

A total of 1859 specimens was recovered,

representing at least fourteen species belonging

to three genera (Table 2). Taxonomic conserva-

tism was practiced wherever possible. For ex-

ample, samples from tidal channel areas con-

tained small turbonills that resembled very

closely Turbonilla puncta (C. B. Adams, 1850),

but graded toward larger shell sizes into definite

T. interrupta (Totten, 1835). Although to my

knowledge T. puncta and T. interrupta have not

as yet been shown to be the same species, I iden-

tified all of these specimens as T. interrupta.

However, within the same series of samples, a

number of different species of Turboyiilla

definitely co-occur, and some of these seem to be

best assigned to the species-groups erected by

Henderson and Bartsch (1914) for Virginia tur-

bonills, which were evidently based upon very

small collections. Taxonomic conservatism has

its natural limitations in the study of the

pyramidellids because there really is an extraor-

dinary richness to the family (cf. Ode, 1981),

probably owing to the mode of evolution within

the group.

Habitat heterogeneity, very limited contact

between populations of adult individuals, small

populations isolated on resource islands (host

organisms), and ability to self-fertilize may have

promoted a kind of accelerated parapatric spe-

ciation among some pyramidellids, with the re-

sult that every geographic segment or compart-

ment of the coast might actually contain its own

recently evolved set of species. Other pyramidel-

lids have evolved to exploit exceedingly wide-

spread prey, and through the Late Cenozoic

have "tracked" their abundant hosts in space

and time to become very widespread and abun-

dant themselves (e. 17.. Odostomia impressa (Say)

with Crassostrea virginica (Gmelin)). Both rarer

species like 0. gibbosa (Bush), and the abundant

and variable forms like 0. iynpret^i^a have with

time become distributed interprovincially.

Nevertheless, if we remember that most pyra-

midellids are parasites, it should not surprise us

to find small numbers of narrowly distributed

species or subspecies, which are actually or

potentially reproductively isolated from other

Vol. 97(3)

July 29, 1983

THE NAUTILUS 95

TABLE 2. Pynuii idcll id sna itf: Jhim the Wii.snaw Sound area, coastal Georgia. Sample numbern are keyed to sampie sta-

tions shown tn Fiy. 1.

SAMPLES

T A X A

■X.

•J- Lr> >X)

Odostomia impressa (Say, 1821)

0. conoidea (Brocchi , 1814)

0. laevigata (d'Orbigny, 1842)

0^ glbbosa Bush, 1909

0^ seminuda (C. B. Adams, 1837)

0^ engonia Bush, 1885

0^ dux Dall and Bartsch, 1906

0. pocahontasae Henderson

and Bartsch, 1914

Odostomia sp. indet .

Turbonilla interrupta (Totten, 1835)

T^ conradi Bush, 1899

T. incisa incisa Bush. 1899

T. powhatani Henderson

and Bartsch , 1914

T. toyatani Henderson

and Bartsch, 1914

Turbonilla sp . indet.

Pyramidella sp. indet.

722

1

1

1

1

80 553 22 13 217 92

5 - 6

- - 2

6 - 1

- 1 -

2 - 1

4 - 2

5 - 3

- 1 -

- 2 2

23 10 IJ

3 - ;

3 - -

6 15

- - 2

'■'Samples composed wholly or partially of reworked Pleistocene shells .

morphologically similar forms, co-occurring

with the more widespread and abundant taxa.

We should expect to find abundant, widely