JANUARY 31, 1983 THE NAUTILUS ISSN 0028-1344 Vol. 97 No. 1 A quarterly devoted to malacology and the interests of conchologists Founded 1889 by Henry A. Pilsbry. Continued by H. Burrington Baker. Editor-in-Chief: R. Tucker Abbott EDITORIAL COMMITTEE CONSULTING EDITORS Dr. William J. Clench Curator Emeritus Museum of Comparative Zoology Cambridge, Mass. 02138 Dr. William K. Emerson Department of Living Invertebrates The American Museum of Natural History New York, New York 10024 Dr. Aurele La Rocque Department of Geology The Ohio State University Columbus, Ohio 43210 Dr. James H. McLean Los Angeles County Museum of Natural History 900 Exposition Boulevard Los Angeles, California 90007 Dr. Arthur S. Merrill 103 West 8th Avenue Cudjoe Gardens Summerland Key, Florida 33043 Dr. Donald R. Moore Division of Marine Geology School of Marine and Atmospheric Science 10 Rickenbacker Causeway Miami, Florida 33149 Dr. Joseph Rosewater Division of Mollusks U.S. National Museum Washington, D.C. 20560 Dr. G. Alan Solem Department of Invertebrates Field Museum of Natural History Chicago, Illinois 60605 Dr. David H. Stansbery Museum of Zoology The Ohio State University Columbus, Ohio 43210 Dr. Ruth D. Turner Department of Mollusks Museum of Comparative Zoology Cambridge, Mass. 02138 Dr. Gilbert L. Voss Division of Biology School of Marine and Atmospheric Science 10 Rickenbacker Causeway Miami, Florida 33149 EDITOR-IN-CHIEF Dr. R. Tucker Abbott American Malacologists, Inc. Box 2255, Melbourne, Florida 32901 Mrs. Cecelia W. Abbott Business and Subscription Manager P.O. Box 22.55 Melbourne. Florida 32901 Tlie Nautilus (USPS 374-980) ISSN 0028-1344 OFFICE OF PUBLICATION American Malacologists, Inc. (United Parcel Address: 2208 South Colonial Drive, Melbourne, FL 32901) Mail: Box 2255, Melbourne, FL 32901 Second Qass Postage paid at Melbourne, Florida and other post offices Subscription Price: $13.00 (see inside back cover) $15.00 (foreign); institutions $18.00 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 of medical and veterinary importance. An invaluable reference work for malacologists, epidemiologists, parasitologists, freshwater biologists and biogeographers. Published by TAYLOR & FRANCIS LTD, LONDON. Distributed in the Americas by AMERICAN MALACOLOGISTS, INC. FLORIDA. 450 pp. 230 X 152mni (9 x 6 0 85066 145 5 Cloth 153 Figs. Regular price $55.00 (plus postage) NOW $27.50 The knowledge resulting from intensive study in recent years is brought together for the first time in this book. Dr. Brown presents a systematic synopsis of nearly 400 species, most of them illustrated, together with chapters on host/parasite relations, snail control, ecology, distribution and biogeography, accompanied by many maps, photographs of shells and drawings. (plus postage) $2. QP NEW: SUPPLEMENT 2 Wagner and Abbotts STANDARD CATALOG OF SHELLS Complete species and generic catalogs of six major families, a revision of 1200 World Size Records, and an Index to genera and subgenera. With seven New guide tabs. Fits into your pres- ent binder. 116 pages, illus. $13.00, plus $1.00 postage (foreign postage $1.50). Families covered: Coralliophilidae {Latiaxis. Magilus. etc.); Triviidae; Ovulidae; Nassariidae (by W. 0. Cernohorsky, about 2,600 names); Glycymerididae (650 names); Spondylidae (250 names). Order now from AMERICAN MALACOLOGISTS, INC., P.O. Box 2255, Melbourne, FL 32901, (ISA. We accept VISA or MASTER CARD payment.s. Give your name, card numher. and the month and year of expiration. 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. THE NAUTILUS 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 ■y D — vr i^ _^''' ov "■♦S'^ ".^^ — ^ — J 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. LITERATURE CITED Baha. K. 1974. Some comments on Lobi.ger souverhii Fischer. 1856, re-identified, of Japan. Veliger 16:253-257. Baba, K. and I. Hamatani. 1970. Occurrences of specimens presumably identifiable with Stitiqer ontutuf! Ehrenl>erg, 1831, at Seto, Kii, Middle Japan (Opisthanz, D. R. 1968. (Occurrence and distribution of New Jersey Opisthobranchia. The Na.utilu.-< 82:7-12. Gascoigne, T. 1978. The internal anatomy of Stiliger fu.fra- lus (A. Gould, 1870) (Opisthobranchia: Sacoglossa). Zool. J. Linn. Soc. 63:265-274. 1979. A redescription of Caliphylla mediter- ranea Costa, 1867 (Opisthobranchia: Ascoglossa). J. Moll. Stud. 45:300-311. Gilbert, S. and K. B. Clark. 1981. Seasonal variation in standing crop of the seagrass Syringodium fiiiforme and associated macro|)hytes in the northern Indian River, Florida. Estuaries 4:223-225. Gould, A. A. and W. G. Binney. 1870. Report on the Inverte- brata of Massachusetts, 524 pp. Grahame, J. 1969. The biology of Berthelinia caribbea Edmunds. Bull. Mar. Sci. 19:868-879. Horsoe, K. 1956. Notas biologicas sobre Eiysia serca Mar- cus, 1955. Contr. avuls. Inst. Oceanogr. Sao Paulo, Biol. 2:1-7. Houbrick, J. R. 1968. A survey of the littoral marine mol- luscs of the Caribbean coast of Costa Rica. Veliger 11:4-23. Jaume, M. L. 1945. El genero Oxynoe en Cuba occidental. Revista de la Sociedad Malacologia 3:18-25. Jensen, K. R. 1980a. A review of sacoglossan diets, with comparative notes on radular and buccal anatomy. Malacol. Rev. 13:5.5-77. 1980b. Oxynoe azuropunctata, n. sp., a new sacoglossan from the Florida Keys (Mollusca: Opistho- branchia). J. Molt. Stud. 46:282-292. 1981a. Observations on feeding methods in some Florida ascoglossans. J. Moll Stud. 47:190-199. 1981b. Influence of filament diameter on food preference and growth in Oxynoe azuropunctata Jensen, 1980 and Eiysia cauze Marcus, 1957 (Opisthobranchia: Ascoglossa). Ophelia 20:127-135. 1982. Chemoreception as a factor in food loca- tion of Eiysia cauze Marcus (Opisthobranchia: Asco- glossa). Mar. Behav. Physiol. 8:205-218. In press a. Occurrence oi Eiysia serca Marcus in Florida, with notes on the synonymy and biology of the species. J. Conch., Lond. In press b. Factors affecting feeding selectivity in herbivorous Ascoglossa (Mollusca: Opisthobranchia). Jour. Exp. Mar. Biol. Ecol. 66. Kay, A. 1964. A new species of Berthelinia. and its asso- ciated sacoglossans in the Hawaiian Islands. Proc. Malac. Soc. London 36:191-197. Lance, J. R. 1962. A new Stiliger and a new Corambella (Mollusca: Opisthobranchia) from the Northwestern Paci- fic. Veliger 5:33-38. Miirch, 0. 1863. Contributions a la faune malacologique des Antilles Danoises. J. Conchyliol. 11:21-43. Marcus. E. 1955. Opi.sthobranchia from Brazil. Bol. Fac, Filos. Cienc. Univ. S. Pauki, Zoologia 20:89-262, pis. 1-30. 1957. On Opisthobranchia from Brazil. J. Linn. Soc. Zool. 43:390-486. 1959. Lameliariacea und Opisthobranchia. Lunds Univ. Aarsskr. N.F. Avd. 2, 55:1-135. 1972a. On some opisthobranchs from Florida. Bull. Mar. Sci. 22:284-308. 1972b. Notes on some opistholiranch gastropods from the Chesapeake Bay. ChesapeaJte Sci. 13:300-317. 1973. On the genus Bosellia (Mollusca: Gastro- poda: Ascoglossa). Bull. Mar Sci. 23:811-823. 1976. Opisthobranchia von Santa Marta, Colom- bia. Stud. Neotrop. Fauna Environni. 11:119-150. 1977. An annotated checklist of Western Atlan- tic warm water opisthobranchs. J. Moll. Stud.. Suppl. 4:1-21. 1980. Review of western Atlantic Elvsiidae (Opisthobranchia Ascoglossa) with a description of a new Vol. 97(1) January 31, 1983 THE NAUTILUS 13 Etysia species. BulL Mar. Sri. 30:.'54-7;). Marcus, E. and E. Marcus. 19(i0. Opisthobraiichs from American Atlantic warm waters. Bull. Mitr. Sri. 10:129-203. 1963. Opisthcibranchs from the Les.ser Antilles. Stud. Fauna Curat;iu.i 19:1-76. 1967. Tropical American upisthnhraiichs. Stud. 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 (Hainm: 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 /lake superior /^y^ . ^\ MICHIGAN r '- ( / i ^\ ^ f MINNESOTA / < I IOWA A, UJ < 1^ ILLINOIS 0 km 80 ^ ^ 0 miies 50 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. Locality Number Faraneter 1 2 3 li 5 6 7 pH 6.7 6.8 6.6 6.8 6.6 7.9 7.5 Tamporature (°C) 23 23 2k 26 19 18. 5 18 Turbidity (PTU) 9 26 20 25 30 7 3 Dissolved Oxygen (mg/l) U.O 9.5 U.O 12.0 10.0 9.k 8.9 Total Hardness (mg/l CaC03) 55 70 95 PO 50 128 88 Total Alkalinity (ib«/1 CaC03) 30 20 30 20 20 102 82 Carbon Dioxide (mg/l) 2.0 2.0 3.0 2.0 2.0 k.7 6.7 Sulfate (mg/l) — 7.0 — . 3.0 2.0 19.6 6.3 Chloride («g/l) 12.5 12.5 10.0 15.0 5.0 13.6 3-2 Silica (mg/l SIO2) 3.0 2.6 1*.0 2.8 1.7 '*.5 11.6 Secchl Disc (cm) — 51 30 '*5 90 — ~ 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.) Dapth (») ' Substrate SpecloBt 1.0-1.-) 1.1-1.7 1.7-2.0 2.7-1.3 Type lt(2) IB-S A. pllcata 2(1) 26(18) m-B-e, 53(32) '.7(28) 31(19) 9-K-b 2(2) »-9 E. dllatatuB 3(3) 7(7) m-a-g "2(1.3) 19(19) 25(26) 3-ff-b 1(1' m-a F. flava SCt) 12(10) n-8-« wci) '*2(37) 6(5) 8-ft-b Q. Botanerva 8(7) 11(0) n-3-K B4( 72 ^ \'i{\2) a-«-b Q. custuloaa 11(6) 16(9) »-»-g TiltO) 83('.1) 3(2) 8-K-b T. vemjcoaa 5(3) 20(13) •-B-g 6q(«t) t>2('«)) 3-K-6 A. n&r^nata 111 20) »-B-g 8(57) 2(U) s-K-b t(66) m-3 A. ffrandlB 1(17) 1(17) 8-«-b L. costata 8(100) B-g-b S. undulatUB 1(8) 3(25) »-•-« 6(50) 2(17) .-g-b A. carlnata 3(10) •-• 6(20) 21(70) .-g-b 18(1»0) B-S L. radlata 1(2) m-s-g 5(11) 21(1.7) s-g-b L. ventrlcoaa 5W 12(10) »-8-g '•1(33) 66(53) S-g-b L. fraMllo IC*) •-a-g 12(52) I0(lt4) .-g-b L. recta 2(2) 1(1) .-a-g W(55) 37('.2) B-g-b 0. reflaxa !*(1U) 2(7) n-.-g M72) 2(7) •-g-b 0. ollvarla 5(7) 2(3) •-.-g w.(59) 22(30) 1(1} »-g-b P. alata 1(8) 1(8) 8(62) 3(22) .-a-g .-g-b P. laBvlBslna 1(25) 3(75) .-a-g B-g-b T. donaclforids 1(17) 1(17) 1.(66) a-.-g B-g-b T. tmncata 7(70) 3(30) B-g-b 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.) Depth (») Sub.trate 1 .0-1 .3 1.3-1.7 1.7-2.0 2.7-3.3 trpei 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. Baker 3tarn Baker Storn Faally Unionldae Subfaally Unlonlnaa Amblena pllcata e.l. X X X X CvclonaAas tub«rculata X X Elliptic dUatatus X X FuBconala abena X Pusconala flava s.l. X X X X Plathobaaue cyphyus X X Plaurobema cocci neum X X Quadrula met&nerva X X Quadrula pustulosa X X • X ^u&dnila quadrula X X TrltOKonla verrucosa X X • Subfamily Anodontlnae Alaamldonta aarxlnata X X Anodonta candle s.l. X X • X AnodoDta Imbeclllls X • X Arcldons confra^osus X Lasalftona costata X • X SlMpsonlconcha aabl^ua X Strophltue undulatue X X Subfamily Lampelllnae Actlnonalaa carlnata X X X X Caruncullna parva • — LanpslllB anodontoldea X Lampsllls radlata X X X X Laiipsllls ventrlcosa X X • X Leptodea fra^llls X X Ll^mla recta X X • X Obllquaria rsflexa X X • X Obovarla ollvarla X X Proptora alata X X X Proptera laevlaelma • X X Truncllla donaclformla X X Truncllla truncata X X Wisconsin and Havlik and Stansbery (1977) 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 Molluslwing 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 Paleoenvironment Species % of samples containing each species Abundance (average no. ot specimens/ liter ot sample) Backbarrier esCuarine (11 samples) Odostonila impressa Odostomia virginica Turbonilla powhatani 81,8% 36,4 45.6 20.5 (abundant) 4.3 (rare) 7,5 (common) Transitional (1 sample) Odostomia impressa 100 . 0% 34.0 (abundant) 66 . 77. 2.0 (rare) 33.3 2.0 (rare) acutidens 33.3 2.0 (rare) 33.3 2.0 (rare) 33.3 4.0 (rare) ita 100.0 44.0 (abundant) 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) Previously reported as fossils (and oldest documented occurrence-') Not previously reported from fossil material 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). Mean shell Initial shell Days Final shell length increments/ New ribs Mean ribs/ Sex length (mini I'lapsed length (mm) day (mm) grown day CT 4.1 4.1 4.5 0.10 3 0.7 cr 5.0 4.1 5.1 0.02 1 0.2 o- 5.5 4.1 5.6 0.02 0 0 o- 5.7 4.1 5.9 0.05 2 0.5 Cf 6.0 4.1 6.4 0.10 2 0.5 cr 6.2 7.3 6.9 0.10 4 0.5 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, Al0.65) 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 Crisher (Foshay). Additional information in American Malacnlogist^, first edition, 1973, p. 487 and Who'a w'ho. american malacologists (3U.5) 725-22fill P O BOX 2255 MELBOURNE, FLORIDA 32901, U.S.A. Prm.lSHERS OF DISTINCTIVE BOOA'.S' (W MOLLHSK."^ Abbott, R Tucker, editor THE BEST OF THE NAUTILUS 1976 288 pp An exciting and nostalgic anthology of shell collecting In the Nineteenth Century ISBN 0-915726-02 X Cloth $13 95 Arakawa, K Y SHELLS ON STAMPS OF THE WOHLD, 1979 234 pp., 15 color pis ; text figs lilus Catalog and classifications ISBN 0 915826-07-0 Cloth $15 95 Bouchet. P SEASHELLSOF WESTERN EUROPE 1980 (Jan) 144 pp, ail color, living animals, habitats ISSN 0-915826-05-4 Paperback $ 8 95 Houbrick, Richard S THE FAIyllLY CERITHIDAE IN THE INDO-PACiFlC 1979 130 pp , 98 pis , 3 in color l^lonographs of Marine Moiiusca no 1, with buchram iooseieaf binder ISSN 0162-8321 $26 00 Wagner, R J L- Abbott, R Tucker STANDARD CATALOG OF SHELLS 1978, with supplement no 1 (1978) 450 pp Values, world size records, lists, maps, hundreds of figs , some in color 15,000 entries ISBN 0-915826-03-8 Buchram postblnder $40 00 McDonald and Nybbaken, 1980 GUIDE TO THE NUDIBRANCHS OF CALIFORNIA - 112 color plates, 64 pp., ISBN 0-91582608-1 Soft $13 50 Brown, D S FRESHWATER SNAILS OF AFRICA AND THEIR MEDICAL IMPORTANCE 487 pp , lllus Cloth $55 00 THE NAUTILUS quarterly MONOGRAPHS OF MARINE MOLLUSCA .Reprints of out-of-print volumes of nfiUTILUS Vols. 1-40 and Gen. Ind. 1-34. Philadelphia, Pa., 1886-1926/27. clothbound $842.00 paperbound $716.00 Kraus Reprint Millwood, New York 10546 AVAILABLE FROM THE ORIGINAL PUBLISHER Vols. 73 to date (vol. 94 in 1980) per vol. paper $12.00 A 25% discount will be honored if 10 or more vols, are purchased between vols. 73 and 94. Individual numbers are not available. Index to Authors for vols. 1 -75 (with the titles to their articles) paper $ 8.00 Geographical Index to vols. 1-90 and Index to Scientific Names to vols. 61-90 in 1 vol. $24 00 Domestic and foreign customers, please direct orders to. AMERICAN MALACOLOGISTS, INC. P.O. Box 2255, Melbourne, FL 32901 USA ^ / 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 INFORMATION FOR SUBSCRIBERS The annual subscription rate for The Nautilus is $13.00 for individuals (foreign $15.00) and $18.00 for institutions (domestic or foreign). Subscriptions may begin in January. Send check or money order made out to "American Mala- cologists" to the Business Manager, P.O. Box 2255, Melbourne, Florida 32901, U.S.A. Back issues from volume 72 to date are ob- tainable from the Business Manager. 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Continued by H. Burrington Baker. Editor-in-Chief: R. Tucker Abbott EDITORIAL COMMITTEE CONSULTING EDITORS Dr. William J. Clench Curator Emeritus Museum of Comparative Zoology Cambridge, Mass. 02138 Dr. William K. Emerson Department of Living Invertebrates The American Museum of Natural History New York, New York 10024 Dr. Joseph Rosewater Division of Mollusks U.S. National Museum Washington, D.C. 20560 Dr. G. Alan Solem 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 Dr. David H. Stansbery Museum of Zoology The Ohio State University Columbus, Ohio 43210 Dr. James H. McLean Los Angeles County Museum of Natural History 900 Exposition Boulevard Los Angeles, California 90007 Dr. Ruth D. Turner Department of Mollusks Museum of Comparative Zoology Cambridge, Mass. 02138 Dr. Arthur S. Merrill 103 West 8th Avenue Cudjoe Gardens Summerland Key, Florida 33043 i)r. Donald R. Moore Division of Marine Geology School of Marine and Atmospheric Science 10 Rickenbacker Causeway Miami, Florida 33149 Dr. Gilbert L. Voss Division of Biology School of Marine and Atmospheric Science 10 Rickenbacker Causeway Miami, Florida 33149 EDITOR-IN-CHIEF Dr. R. Tucker Abbott American Malacologists, Inc. Box 2255, Melbourne, Florida 32901-0328 Mrs. Cecelia W. Abbott Business and Subscription Manager P.O. Box 2255 Melbourne, Florida 32901-0328 The Nautilus (USPS 374-980) ISSN 0028-1344 A quarterly magazine devoted to malacology. Copyright ^1983 by American Malacologists, Inc. OFFICE OF PUBLICATION American Malacologists, Inc. (United Parcel Address: 2208 South Colonial Drive, Melbourne, FL 32901) Mail: Box 2255, Melbourne, FL 32901-0328 POSTMASTER: Send address changes to above. Second Class Postage paid at Melbourne, Florida and other post offices Subscription Price: $13.00 (see inside back cover) $15.00 (foreign); institutions $18.00 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 >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 distributed taxa with long stratigraphic dura- tions; together with rare, widespread taxa also having long stratigraphic ranges; as well as the rare, very poorly known endemic species or sub- species with short or unknown evolutionary his- tories. The more infrequent case of abundance and widespread geographic distribution versus the apparently more common case of rarity among the Pyramidellidae is probably related to degree of host specificity (see Price, 1980); breadth of distribution is probably a function of host deployment patterns and the age of pyra- midellid lineages. Therefore, the taxonomic philosophies of the liberal and the conservative label-users, excluding of course the methodolo- gic blunders which have multiplied synonyms to the point of absurdity, should both contain an element of natural reality: there are both species that have been overnamed (e.g.. T. in- terrupta) and species that have been unjustifi- ably ignored (e.g.. T. toyatani Henderson and Bartsch). A biologically meaningful taxonomy of pyra- midellids must be developed from an under- standing of how the family has evolved, and not merely from a less extravagant nomenclatural practice. In the words of Price (1980, p. 43), "For such small, short-lived, precisely adapted organisms as parasites, evolution will operate in minature-in short times, in small spaces, but with impressive results." Survey Results Four taxonomic associations of pyramidellid species were readily delineated in the samples: 1) an intertidal to very shallow subtidal estua- rine margin association with abundant shells and fragments of Odostomia impressa; 2) a deeper subtidal estuarine association with few 96 THE NAUTILUS July 29, 1983 Vol. 97(3) TABLE 3. Tnxoiiomir rnmposition ofpyramidellid o.ssdn'a- tions. Wdssdir Sound (iri'd. (Georgia. MAJOR COMPONENTS MINOR COMPONENTS ESTUARINE MARGIN Odostomia impressa OFFSHORE ESTUARINE Turbonilla interrupca OPEN MARINE T. incisa incisa 0. conoidea 0. laevigata 0. Rjbbosa 0. seminuda* 0. enfionia* 0. enROnla T. conradi ENVIRONMENTALLY AND TEMPORALLY MIXED ASSEMBLAGES 0. impressa 0. gibbosa 0. seminuda 0. pocahontasae T. interrupta 0- dux T. toyatani 0. conoidea T . incisa incisa T- powhatani Pyramidella sp . ""allochchonous , probably washed in from deeper water Turbonilla interrupta; 3) a subtidal open marine association with few T. incisa incisa Bush; and 4) a temporally and environmentally mixed as- semblage of fossil and modern odostomes, tur- bonills, and pyrams, confined to tidal channels and the ebb tidal shoal surface (see Table 3). Distribution of the modern associations of pyra- midellid species not only coincides with the dis- tribution of host organisms, but also appears to closely coincide with environmental gradients (a predictable result of membership in physically- accomodated nearshore benthic communities in which environment-organism interactions generally determine community structure; see Johnson, 1972). Estuarine Margin Ansociation - Odostomia irnitressa is a well-known ectoparasite of oysters (Allen, 1958; Wells, 1959, 1961; Robertson, 1978). Its shells are found in great numbers in modern shell deposits surrounding beds and isolated clumps of Crassostrea virginica along the edges of the lower Wilmington River and Wassaw Sound. Nearly all of the shells of 0. im- pressa showed evidence of attack and peeling by crabs, both as repaired and unrepaired shell damage. Although some of the broken shells and small fragments of 0. impressa are the results of physical destruction by waves, in all likelihood most of the fragments are the by-products of crab predation. Crab predation on intertidal populations of 0. impressa may be an important cause of mortality among these snails, and ap- pears to be unreported in the literature. In addi- tion to signs of predation, nearly all shells and fragments were at least slightly damaged due to dissolution of calcium carbonate. (A detailed report on the ecologic significance of the condi- tion of 0. impressa shells from the estuarine margin association is in preparation). Other rare pyramidellids occurring in samples from the estuarine margin are listed in Table 3. Mollusks commonly co-occurring with 0. impressa in- cluded: C. virginica, Mulinia lateralis (Say), Brachidontes exustus (Linne), Geukensia demissa (Dillwyn), and Ilyanassa obsoleta (Say). Offshore Estuarine Association - Samples recovered from Wassaw Sound, which did not contain a mixture of fossil and recent mollusks, were characterized by a few shells of Turbonilla interrupta. the second most abundant pyrami- dellid collected in the survey (Table 2). T. inter- rupta is a common ectoparasite of shallow- water bivalves in areas of near-normal marine salinities (Morton, 1967). In addition to about half of all shells of T. interrupta showing signs of peeling and breakage by crabs, one-third of all shells had been bored by predatory gastro- pods. Mollusk species commonly co-occurring with T. interrupta included: Mulinia lateralis, Tellina agilis Stimpson, Abra lioica (Dall), and Acteocina canaliculata (Say). Open Marine Association - Only sample KJ-1 contained modern shells of Turbonilla incisa in- cisa and T. conradi Bush. The discovery of these two snails in a modern shell deposit off the Geor- gia coast represents a northern extension of geographic ranges for both species, which nor- mally inhabit waters south of Georgia (Abbott, 1974). To my knowledge, both species are bio- logically unknown. The single specimen of T. Vol. 97(3) July 29, 1983 THE NAUTILUS 97 conradi showed signs of repaired crab damage. Mollusks co-occurring with these two species in- cluded Mulinia lateniU^ and Tellinn agiliy:. Mixed Assemblages - The most species-rich collections of pyramidellids came from samples taken in the estuary channel, inlet, and from the ebb tidal shoal surface (Fig. 1, Table 2). Most of the shells were broken and abraded, and many were chalky or stained with iron oxide. The most common pyramidellid in these samples was Turbonilla inter rupta. mostly derived through reworking of the older shelly Pleistocene deposits that underlie the area (Frey ct (»/.. 1975). Other fossil pyramidellids found in the samples were the distinctive shells of Odostomia seminuda (C. B. Adams) and 0. dux (Dall and Bartsch) (which were absent from the open marine and offshore estuarine associations), and a variety of turbonills referrable to T. incisa in- cisa. T. toyatani. and T. powhatan i CHenderson and Bartsch) (Table 3). These samples also con- tained the only specimens of Pyramidella en- countered in this survey. Associated with the pyramidellids were large numbers of the follow- ing mollusk species, most of which showed signs of being derived from Pleistocene deposits: Mulinia lateralis. Crassostrea virginica. Tellina agilis, Spisula solidissima (Dillwyn), Anadara transversa (Say), Anadara oualis (Bruguiere), Anadara brasiliana (Lamarck), Donax variabilis Say, Abra aequalis (Say), Crassinella lunulata Conrad, Corbula contracta Say, Nucula proxima Say, Acteocina canalicu- lata. and Mitrella lunata (Say). This bewildering assemblage of broken, chalky, stained shells is of especial interest to paleontologists, as it represents a mixed fossil assemblage "in the making." Not only have several stratigraphic horizons contributed shells to the assemblage, but a variety of estuarine and marine benthic communities are represented. If entombed at the base of a regressive sequence of estuarine sediments, the mixed pyramidellid assemblage would be easy to recognize (even if individuals cannot be identified to species) because of the condition of shells, association with an environmental mixture of other mollus- can taxa, and envelopment within a coarse- grained matrix of channel sand. This assem- blage would grade upward in the sequence into the offshore estuarine association contained in a finer-textured matrix of muddy sand containing M. lateralis shells, which in turn could be overlain by an oyster biostrome containing the estuarine margin association. A transgressive sequence would, on the other hand, consist of either a tidal channel deposit containing a mixed assemblage of shells, the offshore estuarine association, or oyster bank deposits containing abundant 0. impressa. grading upward into either the open marine association or the ebb tidal shoal sediments also containing a mixed assemblage of pyramidellids. ACKNOWLEDGMENTS The use of research vessels and laboratory space at the Marine Extension Service of the University of Georgia, at Skidaway Island, Georgia, is gratefully acknowledged. My stay at Skidaway Island was supported by a grant from the Administrators of the Tulane Educational Fund, expedited by Dr. Emily H. Yokes. Dr. Robert W. Frey assisted with the fieldwork and Dr. R. Tucker Abbott gave valuable advice on the literature of pyramidellids. Mrs. Elizabeth Seale skillfully typed the manuscript. LITERATURE CITED Abbott. R. T. 1974. American Seanhells (2nd Ed,). Van Nostrand Reinhold, New York, 66.3 pp. Allen, J. F. 1958. Feeding habits of two species of Odosto- mia, The Nautilus 72:11-15. Bartsch, P. 1909. Pyramidellidae of New England and the adjacent region. Proc. Boston Soe. Nat. Hist. 34(4):67-113 + plates 11-14. Frey, R. W.. M. R. Voorhies and J. D. Howard. 1975. Estu- aries of the Georgia coast, U.S.A.: sedimentology and biology. VII. Fossil and recent skeletal remains in Georgia estuaries. Senckenherginna martt. 7:257-295. Henderson, J. B. and P. Bartsch. 1914. Littoral marine mollusks of Chincoteague Island, Virginia. Proc. U. S. Nat. Mus. 47(2055):41 1-422 + plates 13-14. .Johnson. R. G. 1972. Conceptual models of benthic marine communities, In Schopf, T. J. M. (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- 98 THE NAUTILUS July 29, 1983 Vol. 97(3) communities. Dowden, Hutchinson and Ross, Strouds- burg. Penn., p. 1-28. Maryland Geological Survey. 1904. Miocene Plates. Johns Hopkins Press, Baltimore, 127 pp. + 125 plates. Morris, P. A. 1973. A Field Guide to Shells of the Atlantic and Gulf Coasts and the West Indies (3rd. Ed.). Houghton Mifflin, Boston, 330 pp. + 76 plates. Morton, J. E. 1967. MoHmsc.s. Hutchinson Univ. Library, London, 244 pp. Ode, H. 1981. A catalogue of Pyramidellidae. Texa^ Con- chologist 17(4):99-104. Price, P. W. 1980. Evolutionary Biology of Parasites. Mono- graphs Pop. Biol. 15, Princeton Univ. Press, 237 pp. Robertson. R. 1978. Spermatophores of six eastern North American pyramidellid gastropods and their systematic significance (with the new genus Boonea). Biol. Bull. 155:360-382. 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. Monographs 31: 239-266. OBSERVATIONS ON THE LIFE HISTORY OF THE WENTLETRAP EPITONIUM ECHINATICOSTUM IN THE BAHAMAS Robert Robertson Academy of Natural Sciences Nineteenth and the Parkway Philadelphia, PA 19103 ABSTRACT The wentletrap Epitonium echinaticostum (Orbigny, 18Jt2) lives with the ac- tiniarian sea aneinone Bunodeopsis g\oh\)i\\iera{Dnchassaing, 1850) at Freeport, Grand Bahama Island. Both were collected in a canal from turtle grass (Thalassia testudinum Banks ex Konig) leaves. The Epitonium feeds on portions of the anemone's tentacles. Further observations are reported on E. echinaticostum's habitat, host, feeding, sexuality, spermatozeugmata, egg capsules, eggs, planktotrophic veligers, protoconchs and purple dye. As expected, there is protan- dry. but. puzzlingly. squashes of 27 whole animals of all representative sizes failed to reveal any oocytes or eggs - although some of the individuals had just laid egg capsules in the laboratory. Comparisons are m,ade with the larger, Caribbean species E. albidum (Orbigny, 18i2) and the Western Pacific E. millecostatum (Pease, 1860-1861), the only other tropical wentletraps on which similar life history data have been published. Wentletraps (Gastropoda: Epitoniidae) are now well-known to live with or to crawl in search of coelenterates, on which they all ap- parently feed (Robertson, 1981a: 13). However, the only tropical western Atlantic species whose host has so far been reported is Epitonium albidum (Orbigny, 1842), which feeds on Sticho- dactyla helianthus (Ellis, 1768) (Robertson, 1963, 1983a and 1983b). It was therefore with much interest that I learned from Jack Worsfold (an avid naturalist and shell collector) that he had found E. echinaticostum (Orbigny, 1842) associated with a sea anemone at Freeport, Grand Bahama Island, northern Bahamas. In early and mid-September 1982 I was at Free- port to study the association myself. Methods Salinity was determined with an American Optical Hand Refractometer. The gamete deter- minations in Figs. 3B and 3C (and the observa- tions on a dearth of females) are based on scjuashes of whole live animals (unstained and Vol. 97(3) July 29, 1983 THE NAUTILUS 99 3 •- U) 0) a 3 10 8 6 4 2 0 3 many ' -Q some h I , 1,1., I I ih II III I ,11 II ill egg capsules mature spermatozeugmata ,aC developing q spermatozeugmata lllil.lli . I .il 3.0 3,5 4 0 SHELL LENGTH ( m m ) 4.5 6,0 FIG. 3. Epitonium echinaticcistum; A. Shell length frequency distribution of a randomly collected sample obtained at Freeport, Grand Bahama Is. in early- and mid-Septembei^ 1982. B. Lengths of the freshly collected animals containing developing sper- matozeugmata and their frequencies in mid-September. C. Same, but mature sperTnatozeugmata. No developing or mature spermatozeugmata were found in specimens l.U, 5.8 and 5.9 mm long. D. Lengths of specimens that laid eggs in the laboratory in early- and mid-September 1982, and the numbers of capsules in each cluster. The data, clearly show protandry. uncleared) that were studied with a compound microscope. Voucher Specimens Epitomwn echinaticostum: Acad. Nat. Sci. Philadelphia A9513 (portion of sample in alco- hol); 355872 (dry shell [Fig. 2]). Bunodmpsis globulifera (Duchassaing, 1850): California Acad. Sci. 031663. Locality and Habitat Worsfold showed me the best locality that he had found for Epitonium echinaticostum which is near the southwest end of the about 20-year- old canal between Oceanhill Boulevard and Bamboo Cay (Street), southwest Freeport (0.4 km E.S.E. of Xanadu Beach Hotel and Marina; 26°30'N.; 78°43'W.), Grand Bahama Island, Bahamas. The host anemone lives on many of the hard substrates in the habitat but is most readily collected from the distal halves of turtle grass leaves {Thalassia testudinum Banks ex Konig), the plants of which grow patchily on the canal's soft substrates at a depth at low tide of about 0.5 to 2 m. The bottom salinity was tested once and was approximately 34 "Uo- Also present in the sheltered canal habitat are such conspicuous organisms as the algae Peni- cillus, Halimeda, Acetabularia and Caulerpa, the large, benthic scyphozoan Cassiopea xamachana Bigelow, 1892, the gastropods Ceri- thium litteratum (Born, 1778), Modulus modulus (Linne, 1758), Strombus gigas Linne, FIG. 2. Epitonium echinaticostum: repre.senliilii'e shell. Freeport, Bahama,^. 100 THE NAUTILUS July 29, 1983 Vol. 97(3) 1758, Fasciolaria tulipa (Linne, 1758), and the fishes Lutjamts apodus (Walbaum, 1792) and Eupomacentrus spp. The bivalve Isognomon alatus (Gmelin, 1791) is common on the canal walls. Some of these organisms commonly live on or near mangroves, but no mangroves are present at the Oceanhill Boulevard-Bamboo Cay locality. The Host The sea anemone host of Epitonium echina- ticostum at Freeport is Bunodeopsis globulifera (Duchassaing, 1850)' (Fig. 1). This species is unusual in having relatively large, conspicuously colored and patterned ovoid vesicles on the col- umn. B. globulifera can turn itself outside in so that its tentacles go through the mouth and become hidden in the coelenteron (Duerden, 1902:299 made similar observations); the mouth can also invert. This behavior may afford Buno- deopsis some protection against animals that at- tack its tentacles. When the tentacles are ex- tended, Bunodeopsis is able to detach its pedal disc and swim weakly or drift to another sub- strate. With the tentacles spread radially, Buno- deopsis globulifera at Freeport attains a max- imum diameter of about 3 cm; the pedal disc diameter is up to about 2 cm. The tentacles of Bunodeopsis globulifera are translucent, faintly tinted with orange-brown. The column and pedal disc are pale orange- brown. The vesicles are cream-colored (some- times pale-brown or tinted with green) with brown or purple-brown spots or stripes. Bunodeopsis globulifera has hitherto been recorded only from Bermuda, Florida, Jamaica, Guadeloupe, Barbados and Curasao (Duchas- saing and Michelotti, 1861: 320, as Viatrix globulifera; Verrill, 1900; Duerden, 1902; Carlgren, 1949, 1952; Lewis and HoUingworth, 1982). The Parasite The shell oi Epitonium echinaticostum (Fig. 2) has from 6 to 15 wavy axial ribs per whorl 'Probable synonyms: B. antilliensis Ducnlun. 1897; B. globulifera Verrill, IDOO. iranitrttuiiiiiMiM. FIG. 1. Bunodeopsis globulifera on (i Thalassia testudinum leaf. Note the vesicles on the column and the juveniles on the leaf. Hope Toum, Ahaco. Bahamas, mm scale. (Clench and Turner, 1951:253-255; Robertson, in press). The maximum observed shell length is 9.5 mm (Clench and Turner, 1951). The largest observed at Freeport was 5.9 mm long; the modal length at the time of the observations was about 2.4 mm (Fig. 3A). The shell is refigured here because Clench and Turner's three illustra- tions (copied by Abbott, 1974:120) all show shells with unusually disjunct whorls (see Rex and Boss, 1976, about "open" coiling of this species). The distinctive shell characters of E. echinaticostum led to its becoming the type- species of Cycloscala Dall, 1889, ranked as a subgenus of Epitonium by Clench and Turner (1951:253). In color, the overall aspect of living Epito- nium echinaticostum is cream-white, with much of the soft body surface finely speckled with pale brick-red and cream. The spire becomes covered with cream-colored detritus. Thus the Epito- nium is not cryptically colored either with its anemone or on Thalassia leaves. The known geographic range of Epitonium echinaticostum is from Bermuda, the Bahamas and Florida south through the West Indies to the Caribbean coast of Panama and the state of Espi'rito Santo, Brazil (Clench and Turner, 1951:255; Olsson and McGinty, 1958:13; Rios, 1975:56, pi. 15, fig. 216). It has been dredged as deep as 200 fathoms [ = 366 m] (Clench and Turner, 1951), but this record may have been based on an empty shell. Porter (1974:167) records the species from S.E. of Cape Lookout, North Carolina, in 200 m (not live-collected), but Vol. 97(3) July 29, 1983 THE NAUTILUS 101 this identification needs to be checked. Abbott (1974:120) remarked that "Dan Steger reports it common in Florida Bays and inlets" - sheltered areas similar to the Freeport canals. At the Freeport locality it was not feasible in the time available to quantify the abundance of Epitonium echinaticostum except to record that an average of about 11.3 specimens was col- lected per man hour (138 specimens in a total of about 12V4 hours). The Epitonium in all cases was found on Thala^sia leaves near Bunodeopsis globulifera. Thus the species is epifaunal, not in- faunal as speculated by Rex and Boss (1976:295, the "axial ribs. . . may function to anchor in- dividuals . . . firmly in the substratum adjacent to their hosts."). Feeding In the laboratory, Epitonium echinaticostum. was seen to feed both during the day (five times) and at night (six times), always on Bunodeopsis tentacles. Feeding was most readily observed after the wentletraps had been starved for several days. A tentacle was ingested either starting from the tip and extending proximally (once) or by being bitten off somewhere along its length and thence being ingested either towards the tip or the base (7 times). The tentacles are too big to be swallowed whole (they can be 10 mm or more long, and the wentletrap at Free- port was less than 6 mm long). When the acrem- bolic proboscis of E. echinaticostum. is fully everted it is about V2 or % the length of the shell. After a piece of Bunodeopsis tentacle of manageable length is nipped off, it can be seen passing proximally in the inverting proboscis. The dishes in which feeding took place became littered with discarded tentacle tips. Two feeding attacks by moderate-sized wentletraps were seen to be thwarted by an anemone pulling away the threatened tentacle in time, after it had been touched by an Epitonium. A 1.4 mm long Epitonium was seen repeatedly trying to attach the tip of its tiny proboscis onto a much larger diameter anemone tentacle, but without success. There was no indication that Bunodeop- sis vesicles or columns were ever attacked. Sexuality Squashes of whole animals were made of 27 freshly collected specimens of all representative sizes. As expected, Epitonium echinaticostum is protandric, with developing spermatozeugmata present in the gonads of individuals 1.5 to 3.8 mm in length (Fig. 3B), and with mature sper- matozeugmata in individuals 2.2 to 4.9 mm in length (Fig. 3C). Puzzlingly, not a single oocyte or egg was seen in any of the 27 specimens, although specimens 3.8 to 4.8 mm-long laid eggs in the laboratory (Fig. 3D). Animals that had just laid eggs contained only spermatozeug- mata. After initial protandry there is a relative- ly long stage of simultaneous hermaphroditism during which eggs perhaps are grown and ex- pelled rapidly by a few individuals. (Growth can be very rapid in fipi^oniMm- Robertson, 1983a). Conditions possibly are different in seasons other than later summer. There is no evidence yet for an exclusively female stage in E. echina- ticostum, as there is in E. albidum (Robertson, 1981b). The largest specimens squashed (5.8 and 5.9 mm long) contained gametes of neither sex. Spermatozeugmata The spermatozeugmata of Epitonium. echina- ticostum are unusual in having two postero- lateral projections on each side of the hind part of the lamellar end. Egg Capsules and Eggs Egg capsules (Fig. 4) were first observed several days after the wentletraps had fed voraciously. Clusters of from 2 to 11 capsules (mean of 7 counts: 5) were seen to have been laid by animals 3.8 to 4.8 mm long (Fig. 3D). The capsules are connected one to another by an elastic mucous thread, one end of which comes from the median ventral groove in the posterior three- fifths of the foot of the parent (the pedal pore from which the thread is secreted lies in this groove). Thus the parent drags a cluster of egg capsules behind it. The capsules are elliptical in outline, with several rounded mucoid projections around its periphery. There is no sand agglutination, but 102 THE NAUTILUS July 29, 1983 Vol. 97(3) FIG. 4. Epitonium echinaticostum egg capsule. Freeport. Bahamas. detritus may be attached here and there. In lengths the capsules range from 0.9 to 1.2 mm. The number of contained eggs ranged from 28 to 65 (mean of 8 counts: 43). It was not possible to determine whether small capsules are laid by small females and contain fewer eggs than large capsules, as in Epitonium albidum (Robertson, 1983b). The eggs are cream-white, and their un- cleaved diameters measured 98 to 106 ^m (mean of 10 measurements: 102 ^m). The surrounding medium within the capsule initially is albumi- nous, and causes the eggs to cluster along the capsule's longitudinal axis. Viscosity decreases during development. In the laboratory, all but three of the capsules became infected interiorly with bacteria and protozoans. Planktotrophic veligers hatched from two of the uninfected capsules but were ab- normal. The hatching shell diameter is about 170 fim, i.e. much larger than the egg. The veliger has a coiled shell, a pair of small velar lobes, an operculum, a purplish black pigmented mantle organ, and a pair of eyes but no ten- tacles. Protoconch The protoconch is about 0.3 to 0.4 mm long, i.e. it is much larger than the hatching veliger shell-proof that the larva is planktotrophic and grows substantially while it is in the plankton. Unfortunately, the protoconch could not be studied with a scanning electron microscope, but at X 50 with a dissecting microscope it was seen to have less inflated whorls than Epito- nium albidum. Microsculpture could not be discerned but may be present. Purple Dye No purple was seen to be released when Epitonium. echinaticostum fed, only when it was molested or dying. I have suggested elsewhere (Robertson, 1983b) that Epitonium purple is repugnatorial. Discussion and Conclusions Epitonium echinaticostum is a smaller species than E. m illecostatum or E. albidum, the only other tropical wentletraps on which there are similar published life history data (Robertson, 1981a and 1983b). Like many (but not all) wen- tletraps, each lives with, or crawls in search of, a particular genus or species of coelenterate upon which it feeds. There is little information on the sexuality of Epitonium millecostatum.. Although both E. echinaticostum and E. albidum are protandric, the two species differ considerably in other respects. At least in late summer at Freeport there seems to be no exclusively female stage in E. echinaticostum; animals producing eggs are curiously rare, and large individuals seem to be in a post-reproductive stage. The egg capsules of Epitonium. echinaticos- tum are relatively large, being about the same size as those of small-or medium-sized female E. albidum., a much bigger animal. Differences be- tween the egg capsules of £■. echinaticostum and E. albidum are that the former are much thinner-walled, have mucoid projections, and have the mucous thread attached terminally in- stead of laterally. With their mucoid projec- tions, the egg capsules of E. millecostatum are more like those ofE. echinaticostum but those of the former have thicker walls. Sand agglutina- tion occurs in E. albidum but not in the other two species, which lack loose sand in their microhabitats. The uncleaved eggs of Epitonium, echinaticos- tum. averaging 102 pim in diameter, are bigger than those of £■. millecostatum (mean: 73 ^m) and E. albidum. (mean: 68 ^m)- E. echinati- costum's eggs are not so large, however, as to Vol. 97(3) July 29, 1983 THE NAUTILUS 103 make the species lecithotrophic; it is still planktotrophic. Both in E. echinaticostum and E. alhidum, and possibly also in E. millecusta- tum, the developing larvae appear to subsist in part on the albuminous material initially present inside the capsules. Hatching sizes are larger than egg sizes. ACKNOWLEDGMENTS I am most grateful to Jack Worsfold for all his help and kindness at Freeport. Dr. Daphne Fautin Dunn (California Academy of Sciences) identified the Bunodeopsis and shared her knowledge of sea anemones. Tracy D. Pedersen drew Fig. 2. The following kindly read and criticized various drafts of the manuscript: Dr. Arthur E. Bogan, Dr. George M. Davis, Dr. D. F. Dunn, Virginia Orr Maes and Dr. Joseph Rosewater. My personal bank account helped to support this research. LITERATURE CITED Abbott, R. T. 1974. AmeHcan Seashells. 2nd Ed., Van Nostrand Reinhold Co., N.Y., vii + 663 pp., 24 color pis. Carlgren, 0. 1949. A survey of the Ptychodactiaria, Coralli- morpharia and Actiniaria. Kmgl. Svenska Vetenskap- sakad Handl. ser. 4, 1(1):1-121, 4 pis. 1952. Actiniaria from North America. Arkir for Zoologi. ser. 2, 3(30):373-390. Clench, W. .J. and R. D. Turner. 1951. The genus Epitonium in the western Atlantic. Part. I. Johnsojua 2(30):249-288, pis. 108-130. Duchassaing [de Fonbressin], P. [1850]. Animaux radinires des Antilles. Paris, 33 pp., 2 pis. Not seen. and J. Michelotti. 1861 [1860? 1862?]. Memoire sur les Coralliaires des Antilles. Mevi. Reale Accad. Sci. Tonnii. ser. 2, Classe Sci. Fis. Matemat. 19:279-365, 10 pis. Duerden, J. E. 1902. On the actinian Bunodeopsis globuli- fera. Verrill. Trans. Linn. Soc. London, ser. 2, Zool., 8(9):297-317, pis. 25-26. Lewis, J. B. and C. E. Hollingworth. 1982. Leaf epifauna of the seagrass Thaln.'isia te.->tndinum. Marine Biol. 71(1): 41-49. 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. Bulls. Ameri- can Paleont. 39(177):l-58, 5 figs., 5 pis. Porter, H. J. 1974. The North Carolina marine and estua- rin.e Mollusca-an atlas of occurrence. Univ. North Caro- lina Inst. Marine Sci., Morehead City, vi -t- 351 pp. Rex, M. A. and K. J. Boss. 1976. Open coiling in Recent gas- tropods. Malacologin 15(2):289-297. 2 figs. Rios, E. C. 1975. Brazilian marine mMusks iconography. Museu Oceanografico, Centre de Ciencias do Mar, Funda^ao L'niversidade do Rio Grande, 331 pp., 91 pis. Robertson, R. 1963. Wentletraps (Epitoniidae) feeding on sea anemones and corals. Proc. Malac. Soc. London 35:51-63, pis. 5-7. 1981a ["1980"]. Epitonium millecostatum and Corcdliophila clathrata: two prosobranch gastropods sym- biotic with Indo-Pacific Palythoa (Coelenterata: Zoan- thidae). Pacific Sci. 34(1):1-17, 13 figs. 1981b. Protandry with only one sex change in an Epitonium (Ptenoglossa). The Nautilus 95(4):184-186. 1 fig. 1983a. Extraordinarily rapid postlarval growth of a tropical wentletrap (Epitonium albidum). The Nauti- lus 97(2):60-66, 5 figs. 1983b. Observations on the life history of the went\eir3.'p Epitonium alhidum in the West Indies. Amer. Malac. Bull. 1. In press. Axial shell rib counts as systematic characters in Epitonium. The Nautilus. Verrill, A. E. 1900. Additions to the Anthozoa and Hydrozoa of the Bermudas. Trans. Connecticut Acad. Arts Sci. 10(14):551-572, pis. 67-69. DIET EXPANSION OF BUSYCON CONTRARIUM IN THE ABSENCE OF TRIPLOFUSUS GIGANTEUS (GASTROPODA: BUCCINACEA) Bretton W. Kent Department of Zoology University of Maryland College Park, MD 20742 Biisycon contrarium (Conrad) is a large, melongenid, predatory gastropod occurring along the Atlantic and Gulf coasts of North America (Abbott, 1974). This species feeds 104 THE NAUTILUS July 29, 1983 Vol. 97(3) almost exclusively on sand-dwelling bivalves (Paine, 1963; Kent, 1982), but Paine (1963) found that B. contrarium was not eating two large and apparently poorly defended pen shells (Bivalvia: Pinnidae), Atrina rigida (Lightfoot) and A. serrata (Sowerby) in northwestern Florida. Of the eight predatory gastropods Paine (1963) studied, only the very large fasciolariid Triplofusus^ giganteus (Kiener) (Olsson & Harbison, 1953) preyed on Atrina. The absence of Atrina from the diet of bivalve- specializing B. contrarium, when sympatric with T. giganteus, suggests that an interaction may be occurring between these two gastro- pods. During a study of competitive interactions between busyconine whelks in 1979, I made diet observations on B. contrarium in an area near Paine's study site. These observations are useful for testing the hypothesized interaction between B. contrarium and T. giganteus, since during 1979 T. giganteus was virtually absent from in- tertidal grass flats. Paine (1963) reported a mean density of 3.6 T. giganteuslhr. (from Fig. 1 in Paine, 1963); in 1979 only eight T. giganteus were observed on grass flats in 11 months of in- tensive study (Kent, 1983). In Paine's (1963) study A^rma spp. accounted for 20% of the diet of" T. giganteus (13 of 66 prey); the remainder of the diet consisted entire- ly of gastropods (13 Fasciolaria tulipa, 6 F. lilium hunteria, 17 B. contrarium, 4 Bu^ycoty- pu^ spiratum, 12 Chicoretis dilectus, and 1 Poli- nices duplicatu^). In 1979, six of the eight T. giganteus were feeding and one of these six feedings was on Atrina ( = 17% of diet). As in Paine's study, the remainder of the diet was composed of gastropods (1 F. tulipa, 2 B. con- trarium, 1 C. dilectus, and 1 Melongena corona). Despite the very small sample size for 1979, the diets are surprisingly similar; however the very low densities of T. giganteus in 1979 meant that it was not a numerically important predator. The diet of B. contrarium changed markedly 'Considered a subgenus of Fieumploca by some recent authors -editor. when T. giganteus was rare. Paine (1963) found that no Atrina were eaten in 172 feedings, while in 1979 eight of 63 feedings were on Atrina ( = 13% of diet; Kent, 1983). The change is even more dramatic if only larger (>120 mn) B. con- trarium are considered; eight of 32 feedings were on A trina ( = 25% of diet). This comparison is more suitable than using all B. contrarium because; 1) small B. contrarium were rare dur- ing Paine's study and 2) it compares B. contra- rium and T. giganteus of similar sizes, thus avoiding size-related differences in diet (Kent, 1983). A second sympatric busyconine whelk, Busy- cotypus spiratum (Lamarck), does not prey on Atrina in the absence of T. giganteus. Apparent- ly this is due to a strong preference by B. spira- tum for active bivalves (e.g. - scallops and cockles) rather than passive bivalves such as Atrina (Kent, 1983). While these data suggest that an interaction is occurring between T. giganteus and large B. contrarium, the exact mechanism of this inter- action is unclear. The dietary expansion of B. contrarium may simply represent competitive release, although the situation is complicated by B. contrarium being an important prey of T. giganteus (28% of diet in Paine's study and 33% during 1979). Detailed observations and experi- mental manipulations are needed both to clarify the mechanism for this interaction and to deter- mine the effects this change has had on the selective pressures acting on Atrina. LITERATURE CITED Abbott, R. T. 1974. Am.eHcan Seashells. Van Nostrand Rein- hold Co., N.Y., 663 pp. Kent, B. W. 1983. Patterns of coexistence in busyconine whelits. J. Exp. Mar. Biol. Ecol. 66:257-283. Olsson, A. A. and A. Harbison. 1953. Pliocene Mollusca of southern Florida. Acad. Natl. Sri.. Phila.. Monogr. 8: 172 pp. Paine, R. T. 1962. Ecological diversification in the genus Busycon. Evolution 16:515-523. 1963. Trophic relationships of eight sympatric gastropods. Ecology 44:63-73. Vol. 97(3) July 29, 1983 THE NAUTILUS 105 ECOLOGY AND LARVAL DEVELOPMENT OF CREPIDULA PROTEA (PROSOBRANCHIA: CREPIDULIDAE) FROM SOUTHERN BRASIL: A NEW TYPE OF EGG CAPSULE FOR THE GENUS K. Elaine Hoagland Department of Malacology Academy of Natural Sciences Philadelphia, PA 19103 ABSTRACT Examination of museum specimens verified the existence of at lea^t three distinct species of the marine gastropod gentis Crepidula off the coast of southern Brasil: C. protea, C. plana, and C. aculeata. Speciinens ofC. protea were collected alive by dredging. They carried broods of larvae that appeared to be ready to be released in the veliger stage. Unlike other Crepidula, the larvae were imbedded in the matrix of the egg capsule material. Other character states distinguishing C. protea and C. plana are discussed, as well as the population structure of C. protea. The distribution ofC. protea is limited coinpared with that ofC. plana. Crepidula protea d'Orbigny, 1841, is a com- mon subtidal gastropod found on shelly sub- strates off the coast of South America. Because living specimens are obtained most often by dredging, little is known about the species. Even the limits of variation of the shell phenotype are poorly defined. D'Orbigny (1841, 1842) commented that his description could en- compass North American specimens, presum- ably C. plana Say, 1822. The original species localities were the Antilles, the coast of Patagonia, and "Maldonado, Brasil" [Uruguay]. Dall (1889) synonymized C. protea with C. unguiformis Lamarck, although most other workers have maintained it as a separate species (Rios, 1982 ms.; Hoagland, 1977). In December, 1981, I examined museum re- cords and specimens of Crepidula protea in the Museu Oceanografico de Rio Grande do Sul, Brasil. Once assured on the basis of shell char- acters that C. protea is distinct from C unguiformis and C. plana, I arranged to dredge off the coast of southern Brasil in an attempt to collect living specimens. This paper describes the specimens obtained, including characteris- tics of their brooded embryos and their popula- tion structure. Confirmed localities for C. protea and C. plana, in South America are compiled from examination of the museum specimens. Methods In December, 1981, Prof. E. Rios arranged for me to examine the collection of Crepidula at the Museu Oceanografico, Rio Grande. He also kind- ly arranged for the use of their oceanographic vessel equipped with bongo nets. The nets were lowered and dragged on the bottom approx- imately 10 miles off Barra, S.E. of Rio Grande, in the state of Rio Grande do Sul, southern Brasil. Shelly substrates were sampled at 25, 30, and 40 m. C. protea was found at 30 m. The shell debris was sorted and specimens of Crepidula were returned to the laboratory still attached to their substrates, which were either shell frag- ments or, more commonly, living specimens of the trochid gastropod Photinula blakei (Clench and Aguayo). The specimens were separated from one an- other, sexed, and examined for broods of em- bryos. Shell characters, including size and color, were recorded. All brood capsules were re- moved from the females, examined in seawater, and their embryos counted using a dissecting microscope. The adult specimens were then frozen for electrophoresis, the results of which are reported elsewhere (Hoagland, 1984). Voucher specimens of C. protea are deposited at the Academy of Natural Sciences (ANSP 106 THE NAUTILUS July 29, 1983 Vol. 97(3) #355327). The type specimens (shells only) of Crepidula protea were examined at the British Museum (Natural History), in order to confirm the identity of the Brasilian material. Results There are 16 syn types of Crepidula protea d'OrbigTiy in the British Museum [BM(NH) reg. no. 1854.12.4]. The locality is given only as Patagonia. One lot of 5 specimens (#573) con- tains one white and four tan specimens, all arched, with no muscle scar, and with the shelf margin broken. Length of the shells varied from 16 to 32 mm. A second lot of 11 white specimens (#574) contained shells of length 11-29 mm. They agreed in shell phenotype with the first lot. One specimen did have a septal margin intact; it had a notch at the center and at the left corner. The larger specimens were flatter, approaching C. plana in appearance. The shell phenotypes of Brasilian Crepidula protea were in agreement with the type material. However, the shell characters that I found to distinguish the Museu Oceanografico specimens of C. protea. from those of C. plana are few. The internal septum of C. plana has a deep notch on the left side near where it joins the shell proper, while Crepidula protea has smaller notches located there and at the center of the septum (Fig. lA). The shell of C. plana is thin, white, occasionally with faint orange lines, and flat or slightly convex. The shell of C. protea is slightly convex to highly arched, generally smaller but thicker than C. plana, and most often tan with reddish-brown lines, although pure white specimens occur (Fig. IB). Fifteen percent of the 60 live-collected specimens were white. Table 1 gives the known localities for Crepi- dula protea and C plana based on the Museu Oceanografico collection. The data support the claim of Rios (personal communication and 1982 ms.) that C. protea is found only as far north as Rio de Janeiro, not in the Antilles as thought by Castellanos (1970) and d'Orbigny (1842). The southernmo.st locality represented in the collec- tion for either .species is Miramar, Argentina, although localities to the south are likely. B FIG. 1. The shell of the tmijtsl sj)e.cim.en o/Crepidula protea collected off Rio Grande. Brasil. Shell length = 15.1 mm. A, ventral view. B, dorsal view. One observation distinguishes Crepidula pro- tea from all other Crepidula I have examined. The egg capsules consisted of an outer mem- brane containing a sticky matrix in which the embryos were imbedded, making the embryos difficult to remove from the capsules. Other species of Crepidula, both species with a plank- tonic larval stage (e.g. C. fomicata (Linnaeus), C. plana, C. onyx Sowerby) and those which brood through metamorphosis (C. convexa Say, C. aculeata (Gmelin), C. adunca Sowerby), hold the embryos loosely within the thin egg capsule membrane so that the embryos move freely in a watery medium. Table 2 gives the sex structure of the sample of 60 snails collected off Rio Grande. Most com- monly, a male-female pair was found on one piece of substrate, with the male sitting on the right side of the shell of the female in mating position. Specimens of opposite color did form male-female pairs. There were 2 juveniles, 25 males, 2 intermediates undergoing sex change, Vol. 97(3) July 29, 1983 THE NAUTILUS 107 TABLE 1. Localities of specimens o/Crepidula protea nnd C. plana deposited and cataloged in the Museu Oceano- grdfico. Localities are arranged approximately north to south. Locality Species Depth (rn) BRASIL lie de Itamaraca C. plana Cassipore C. plana Sepetiba, R.J. C. plana Off Juatinga. R.J. C. plana 40-50 Cabo Ig^iape, Sao Paulo C. protea lmbitul)a, S.C. C. protea Off Amapa C. plana 45 Off Rio Grande do Sul C. protea, C. plana 20 Farol Conceicjao, R.G.S. C. protea. C. plana 18 Sul da Barra. R.G.S. C. protea. C. plana 15 Cassino, R.G.S. C. protea Farol de Albardao, R.G.S. C. protea. C. plana 25.35 Chui, R.G.S. C. protea 166 Uruguay Cabo Tolouio C. plana 40 La Coronilla C. protea La Paloma C. protea Ilha dos Lobos C. plana Argentina San Antonio C. protea. C. plana 40 370 Parallel C. plana Miramar C. protea. C. plana TABLE 2. Sex structure 0 (dredged sample o/Crepidula pro- ten from a single locality. Sex Category Number ofSp ecimens Juvenile alone 2 Intermediate alone 2 Male alone 5 Male on Female 3M + 3F Male on Female with embryos 17 M + 17 F Female alone 3 (1 parasitized) Female alone with embryos 8 60 total with a range of 26 to 48. The largest specimen contained the largest number of egg capsules. One capsule from each of 10 females was counted to determine the average number of embryos per capsule: the mean and standard er- ror were 61 and 8, respectively. The largest specimen contributed greatly to the high vari- ability. It contained capsules of 120 embryos, whereas the range without that specimen was 33-70 embryos per capsule. Electrophoretic study (Hoagland, 1982) confirmed that all specimens belonged to the same species, so one is not justified to remove the larger specimen from the analysis. The average number of em- bryos per brood was 1,927. All of the embryos were in the veliger stage or earlier. All embryos in each brood were at a similar developmental stage. Because none of the broods examined were in the egg stage, the diameter of the uncleaved egg of Crepidula pro- tea could not be determined. The diameter of a developing embryo was approximately 0.15 mm. The shell diameters of the brooded veligers were not significantly greater than the diameter of earlier development stages, nor did there ap- pear to be less capsule matrix in capsules of more advanced development. The veligers were pinkish in color, unlike the nearly transparent veligers reported for Crepidula plana (Hoagland, 1977, p. 390). Facilities for rearing larvae were unavailable, but it appeared that the larvae would be released as veligers. There was no sign of resorption of the velum or enlarge- ment of the foot and shell in any of the 25 broods examined. and 31 females of which 25 (81%) were brooding embryos. The sex ratio of the mature individuals was 45% males. The size range of females was 7.3 to 15.1 mm in length and 4.8 to 9.7 mm in width. Males were 4.6 to 8.0 mm long and 3.8 to 5.7 mm wide. The largest female (Fig. 1) was on a dead shell of C. protea that was 20 mm long. The size of each of the other specimens was con- strained by the size of the shells used as sub- strate. The average number of egg capsules per brood was 31.6 with standard error of 1.1, and Discussion Crepidula protea as seen in Brasil is the same as d'Orbign/s type material. The dredged specimens were especially like smaller in- dividuals of BM(NH) #574. The species is distinct from C. platia primarily on the basis of its egg capsule composition and the shape of the septum. C. protea is more variable in shell shape and color than is C. plana. Specimens of C. pro- tea constrained to a small substrate become highly arched; they reproduce at a small size and have smaller broods than unconstrained speci- mens. Both the number of egg capsules per 108 THE NAUTILUS July 29, 1983 Vol. 97(3) brood and the number of embryos per capsule are reduced. In fact, these specimens of C. pro- tea are the smallest of the genus that I have seen to produce larvae that are apparently plankto- trophic. An average brood size of nearly 2,000 for C. protea can be compared with C. plana of New England, about 5,000-8,000 planktonic larvae, and C. convexa, about 200 non-planktonic larvae (Hoagland, 1975). Both the number of embryos per capsule and the number of capsules per female C. protea were of the order of magnitude expected for a species of Crepidula with plank- totrophic development. Because one large speci- men contained the largest brood, and because the sizes of the dredged specimens were on the low end of the size range seen in museums, I ex- pect that the ~2,000 embryos per brood reported here is at the low end of the range of brood sizes for C. protea. The embryo diameter of 0.15 mm is similar to the 0.136 mm reported for eggs of C plana (Conklin, 1897). The sticky capsule matrix must be investi- gated chemically to determine if it is a nutritive material that is ingested by veliger-stage or earlier larvae remaining in the capsule. The observation on the broods from southern Brasil do not support the hypothesis, because the cap- sule matrix did not appear to break down or decrease in volume with the advancement of lar- val development. Measurement tools available did not detect size increase of embryos within the capsules. However, a full range of larval sizes were not seen, and it is still possible that the matrix is used late in development, or that its composition changes with development. The size ranges of adult males and females support the notion that, like all other known Crepidula, C. protea is protandrous. From the scarcity of juveniles present and the large percentage of females brooding embryos, one would expect December to be near the begin- ning of a breeding season. The presence of several brooding females without males shows that mated pairs are not permanent, although stacking of at least 3 specimens (one a dead specimen) was observed. No obvious growth lines were found on the shells to indicate age of the large stacked specimens, although most of the smaller specimens had a single growth discontinuity near the shell margin. The distribution of all species of Crepidula in the southern Atlantic is limited because of the lack of suitable substrate. Only in mudflats or on scallop or oyster beds (as in Mar del Plata and San Antonio, Argentina) are Crepidula abun- dant and in shallow water. Dredging off Rio Grande demonstrated that specimens of C. pro- tea are patchy. No specimens of C. plana were collected, so anatomical comparisons of the two could not be done. However, museum records do document the sympatry of the two species off Rio Grande. Four living specimens of Crepidula aculeata were collected together with those of C. protea. Body pigment patterns, shell characters, and electrophoretic patterns confirmed that this is the same species as C. aculeata from Florida. Brasilian museum records document a nearly continuous distribution of C. aculeata from Florida to Mar del Plata, Argentina. No other species of Crepidula besides C. plana and C. aculeata are known to coexist with C. protea in southern Brasil; however, at least two species, C. onyx and C. dilatata Lamarck from the Chilean fauna, overlap at Mar del Plata (Parodiz, 1939). The factors that limit the north-south distri- bution of Crepidula protea relative to other species of Crepidula are unknown, especially in the light of the discovery of a planktonic larval form. One would expect phoresis of adults in ad- dition to larval dispersal, because of the small size and sedentary habits of C. protea. and its apparent preference for settling on other living gastropods. However, the relatively greater known range of C. plana may be revised down- ward if some southern populations turn out to be sibling species when they are examined bio- logically. At this time it must be concluded that Crepidula protea is a South American species endemic to southern Brasil, Uruguay, and Argentina. ACKNOWLEDGMENTS This work, particularly the dredging of live specimens, would have been impossible without Vol. 97(3) July 29, 1983 THE NAUTILUS 109 the arrangements made by Prof. E. Rios of the Museu Oceanografico Rio Grande do Sul. R. Capitoli and G. M. Davis assisted in collecting the specimens. P. Mordan of British Museum (Natural History) made available the types of Crepidula protea. The manuscript was read and criticized by G. M. Davis and R. Robertson. Funding was provided in part by a Fleischmann Foundation grant to the Wetlands Institute, Stone Harbor New Jersey. I appreciate the hospitality of S. and R. Capitoli during my stay in Rio Grande. LITERATURE CITED Castellanos, Z. J. A. de. 1970. Catalogo de Los Moluscos Marines Bonaerenses. La Plata, Anales Comisioti de In- restigacion Cientifica, Provincia de Buenos Aires 8:9-36.5. Dall, W. H. 1889. Reports on the results of dredging, under the supervision of Alexander Agassiz. in the Gulf of Mex- ico (1877-78) and in the Caribbean Sea (1879-80), by the U.S. Coast Survey Steamer "Blake," Report on the Mol- lusca. Part 2: Gastropoda and Scaphopoda. Bulletin of the Museum of Comparati re Zoology 18: 492 pp., 31 pis. Hoagland, K. E. 1975. Reproductive strategies and evolu- tion in the genus Crepidula (Gastropoda: Calyptraeidae). Ph.D. Diss. Harvard University, 360 pp. 1977. Systematic review of fossil and recent Crepidula and discussion of evolution of the Calyptrae- idae. Malacologia 16(2):353-420. 1984. Use of molecular genetics to distinguish species of the gastropod genus Crepidula. Malacologia, in press. Orhigny, A. D. d'. 1841. Mollusques. Voyage dans I'Amerique Meridionale, 1826-1833. Ministre de I'lnstruction publique, Paris, 5(3): 758 p. 1842. Mollusques. In: Histoire Physique, Poli- tique et Naturelle de File de Cuba, vol. 2, Sagra, R. (ed.). Bertrand. Paris, 264 p., 28 pis. Parodiz, J. J. 1939. Las especies de Crepidula de las costas Argentinas. Physis, Buenos Aires, 17:685-709, 1 pi. Rios, E. 1982. South America Sea.'ihells [manuscript]. Say, T. 1822. An account of the marine shells of the United States. Jour. Acad. Natural Sciences Philadelphia, 2: 221-227. THE FRESHWATER NAIAD (MUSSEL) FAUNA OF THE NOLIN RIVER IN THE GREEN RIVER DRAINAGE OF CENTRAL KENTUCKY (MOLLUSCA: BIVALVIA) Ralph W. Taylor Department of Biological Sciences Marshall University Huntington, West Virginia 25701 ABSTRACT A survey of the freshwater mussels (naiads) ofNolin River of the Green River, Kentv/^ky, drainage, produced specim,ens of twenty-one species ofyiaiads, and the Asian Clam, Corbicula fluminea. Six of the species reported from the Nolin are considered Endangered or of Special Concern by the Kentucky Academy of Science. Villosa ortmanni is considered endangered by most malacologists because it is restricted in distribution to the Green River drainage. At present, however, it is the most commoyily found species in the Nolin River. Ortmann (1926) recognized the importance of the Green River as a freshwater mussel stream. Individuals were abundant, species were numerous. The typical Ohioan fauna of this stream was identical with the Kentucky River and other Ohio River tributaries to the north, but distinctly different from the Cumberlandian fauna of the Cumberland and Tennessee Rivers just to the south. Clench and van der Schalie (1944) did addi- tional work on the mainstem Green River and made extensive collections in several major tributaries, such as the Nolin, Barren and Rough Rivers. Additionally, during the 1950's no THE NAUTILUS July 29, 1983 Vol. 97(3) and 1960's David Stansbery (1965) and others from Ohio State University collected extensive- ly in the Green River. Their work, including that of their predecessors, produced a list of 64 species resident in the Green River drainage and led Stansbery (1965) to state that the mussels in the Green River . . ."comprise the finest representative Ohioan naiad fauna yet in existence." Within the last twenty-five years adverse ef- fects of damming have come to the unique Green River. At the present time all the major tributary headwaters, including the mainstem, have been dammed. While no studies document- ing a concommitant decline in mussel popula- tions have been forthcoming, it must be as- sumed, based on numerous other studies, that the dams will have deleterious effects. The dams and the large lakes behind them, which are unsuitable habitat for most stream forms, are formidable barriers and restrict the free flow of host fish species between the main- stem Green River and tributary headwaters. The full long-term effect of such barriers has not yet been documented. Baseline data on the mainstem Green River are readily available through the works of the previously-mentioned authors. There is, however, a dearth of recent information available on some of the major tributaries. Clench and van der Schalie (1944) collected the Barren River extensively and composed a list of 36 resident species. Their work at two sta- tions on the Rough River netted 22 species, and limited collecting on the Nolin River produced only 8 species. I have been unable to find addi- tional papers on this drainage, and in this paper an attempt is made to fill a gap in the data base. The only previous collections on the Nolin re- ported in the literature were made nearly forty years ago and prior to impoundment of the river. This paper reports on extensive collecting in the river, above Lake Nolin, during the sum- mer of 1981. The Nolin River is one of two major tribu- taries of the Green River which enter from the north. The Nolin originates in the Mississippian Plateau Province of central Kentucky. The origin in Larue Co. is located a few miles south c^ of the Abraham Lincoln Birthplace National Historical Site. The river then flows through Hardin, Grayson and Edmonson Counties to its confluence with the Green River within the con- fines of Mammoth Cave National Park. Collecting Stations 1. Nolin River at White Mills, Hardin County, Kentucky, two miles E of State Rt. 84 on Coun- ty Rd. 1904. 2. Nolin River, four miles due west of village of Flint Hill on State Rd. 720 on Hardin/ Grayson Co. line. (This site roughly corresponds to one site reported by Clench and van der Schalie.) 3. Nolin River at Millerstown, at intersection of State Rt. 224 and 479. (Extremely rich area for several hundred meters above and below the bridge.) Numerous other sites were visited, but these three were by far the most productive. The Millerstown site is to the Nolin River what Mun- fordville is to the Green River, the site where all conditions seem to be prime for mussel habita- tion. Naiad Species Recorded from the Nolin River 1944 = Previously recorded (Clench and van der Schalie, 1944). 1981 = Recently collected (Taylor, 1981). Sh-ophitUH u. undulatu.'i (Siiy. 1817) 1944; 1981 Alnxtmd inch) shells from the Florida Keys, with accurate locality data. Also unsorted grange: write for list. Margaret Teskey P.O. Box 273 Big Pine Key. Fl. 330U3 132 THE NAUTILUS October 28, 1983 A SEMELE STORY (BIVALVIA: SEMELIDAE) Eugene Coan Research Associate Department of Invertebrate Zoology California Academy of Sciences San Francisco, California 94118 Vol. 97(4) In her review of the eastern Pacific molluscan taxa proposed by Morch (1859-1861), Keen (1966) discovered that the type material of the previously unillustrated Semele verrucosa Morch, 1860, was something very different from what most authors had assumed. Morch's taxon, she concluded, was probably conspecific with S. margarita Olsson, 1961, and later she syno- nymized the two without question (Keen, 1971). This discovery was presumed to leave without a name the species that had been illustrated by previous authors as S. verrucosa Morch. For this, she proposed a new species, S. verruculas- tra Keen, 1966, selecting as holotype a specimen in the collection of the California Academy of Sciences that had been illustrated by Hertlein & Strong (1949) as S. verrucosa. Unfortunately, her description of Semele ver- r^iculastra was predicated upon the distinctions drawn by Hertlein & Strong and other earlier workers between their "S. verrucosa" and a similar species, 5. formosa (Sowerby, 1833). These distinctions prove illusory, and S. verru- culastra falls into the synonymy of S. formosa. Semele formosa (Sowerby) was only briefly dis- cussed by Hertlein & Strong (1949), who evi- dently believed that the material they assigned to Semele verrucosa was separable from S. for- mosa by means of the same features with which Morch had originally differentiated his taxon - more elongate, more subtruncate posteriorly, and with a more gently arcuate ventral margin -and pronounced, scaly, verrucose sculpture. Olsson (1961) figured a specimen of Semele formosa from the BM(NH), and he apparently had no material of his own. He also discussed and illustrated material he believed to be Semele verrucosa, and he indicated that the latter dif- fered from S. formosa in its less convex valves, and its coarser concentric ribs that become divided and scalloped, frilled, or scalelike at their posterior and anterior ends. I have examined a number of specimens of this uncommon species and find that none of these distinctions stands up under close scrutiny. In addition to the two type specimens, I have ex- amined 3 lots in the United States National Museum of Natural History, 2 lots in the Cali- fornia Academy of Sciences, and 3 lots from the collection of Mrs. Carol Skoglund of Phoenix, Arizona. Olsson (1961) provides measurements of two additional specimens, and Draper (1980) of two more. Semele formosa becomes higher and more in- flated as it matures. Its length ranges from 1.23 to 1.33 times its height, with young specimens more elongate. Similarly, its thickness is be- tween 0.43 and 0.52 times its height, the valves becoming more convex as individuals mature. Large specimens may expand somewhat antero- ventrally, making that margin appear more curved. Nor can I see any distinction in the degree of posterior truncation among material that has been illustrated or that has been available to me. The feature that has been relied upon to the greatest degree in differentiating two species is the nature of the concentric ribbing. Contrary to Olsson's assertion, the material that had been assigned to "S. verrucosa" has ribbing that is no coarser than that on the lectotype (designated here) of Amphidesma forraosum. Moreover, growth and the sculptural differences between right and left valves account for the supposed distinction in rib roughness. Small specimens have rougher sculpture, as do right valves. The holotype of S. verruculastra is a right valve, as is the specimen figured by Olsson (1961) as "S. verrucosa. " whereas he illustrated a left valve of S. formosa. Vol. 97(4) October 28, 1983 THE NAUTILUS 133 The following is a partial synonymy of this species: Semele formosa (Sowerby, 1833) Amphidesma fonnosum Sowerby. 1833 Sowerby, 1833a: 7; pit. 19, figs. 8; Sowerby, 1833b: 199; Hanley, 1843: 44; 7 (pit. expL); pit. 12, fig. 48; Reeve, 1853: pit. 4, fig. 27 [as A. yormosa'l Semele formoxa (Sowerby) Hertlein & Strong, 1949: 249; Keen, 1958: 196, 197; fig. 482; Olsson, 1961: 365; 558 (pit. expl.); pit. 85, fig. 8; Keen, 1971: 251, 252; fig. 631; Abbott & Dance, 1983, p. 350 (in color). Semele verrucosa Morch, auctt.. non Morch, 1860 Hertlein & Strong, 1949: 249; 258 (pit. expl.); pit. 1, figs. 21, 24; Keen, 1958: 202, 203; fig. 504; Olsson, 1961: 366; 538 (pit. expl.); pit. 65, figs. 1-lb; Emerson & Hertlein, 1964: 359-360; 357 (pit. expl.); 356. figs, i, j; {non Morch, 1860: 190-191]. Semele I'erruculastra Keen. 1966 Keen, 1966: 32-33; Keen, 1971: 255-257; fig. 653. Type Material: Amphidesma formosuvt - BM(NH) 1907.10.28.20, lectotype (herein), a right valve, the specimen in Sowerby's upper- most figure; length, 50.7 mm; height, 41.2 mm; thickness, 9.6 mm (Fig. 1). Sovv'erby (1833b) FIG. 1. Semele formosa (Sowertyy). lectotype therein), BM(NH) 1907.10.28.20: length. 50.7 mm; Bahia Santa Elena. Ecuador. FIG. 2. Semele formosa (Sowerby). Holotype o/ Semele ver- ruculastra Keen. CASGTC 9256. length. J,2.8 mm: Hannibal Bank, Panama. specifically mentions that only two single valves were collected, of which the lectotype matches the original measurements. Thus, the specimen figured as "holotype" by Olsson (1961), BM(NH) 198224/1, cannot be from the original lot because it is a pair of matched valves, 64.4 mm in length. It is the specimen figured by Reeve (1853). Bahia Santa Elena, Guayas Prov., Ecuador (2°10'S, 80°50'W); H. Cuming; 13 meters. Semele verniculastra - CASI2 036679 (formerly CASGTC 9256), holotype, a right valve; length, 42.8 mm; height, 32.8 mm; thick- ness, 7.1 mm (Fig. 2). Hannibal Bank, Panama (7°23'30"N, 82°3'W); CAS Loc. 17996; about 68 meters. This striking species occurs from the Gulf of California to Bahia Santa Elena, Ecuador. The largest specimen I have seen is in the collection of Carol Skoglund of Phoenix, Arizona. It was collected at Isla Catalina, Bahia San Carlos, Sonora, Mexico, in about 20 meters of water and measures 73.3 mm in length, 59.1 mm in height, and 29.7 mm in thickness. ACKNOWLEDGMENTS I appreciate the loan of the lectotype of Am,- phidesma formosum by the British Museum (Natural History) through the courtesy of 134 THE NAUTILUS October 28, 1983 Vol. 97(4) Solene Morris and the loan of specimens by Carol Skoglund. I also appreciate the advice of Carol Skoglund, James McLean, Barry Roth, Helen DuShane, and Myra Keen on this project. LITERATURE CITED Abbott, R. Tucker and S. Peter Dance. 1982. Compendium ofSeashdts. E. P. Button, New York, x + 411 pp.; illus. Draper, Bertram C. 1980. Lost operculum club list of cham- pions. Los Angeles, Calif. (Conch. Club Southern Calif.) 32 pp. Emerson, William Keith and Leo George Hertlein. 1964. Invertebrate megafossils of the Belvedere Expedition to the Gulf of California. San Diego Soc. Natur. Hist., Trans. 13(7):333-368; 6 figs. (30 Dec.) Hanley, Sylvanus Charles Thorp. 1843 [1842-1856]. An illus- trated and descriptive catalogue of Recent bivalve shells. London (Williams & Norgate). xviii + 392 + 24 pp.; pits. 9-24 [dating: pp. v-vi & Reynell (1918)] [pp. 1-32 (late 1842); pp. 1-32 (second issue), 33-144; pits. 9-13; pp. 1-8 (early 1843); pp. 14.5-272 (late 1843); pits. 14-16; pp. 9-12 (late 1844); pp. 13-18 (1846); pits. 20-24; pp. 19-24 (26 July 1855); pp. i-xviii + 273-392 (1856)]. Hertlein, Leo George and Archibald McClure Strong. 1949. Eastern Pacific Expeditions of the New York Zoological Society. XLI. Mollusks from the west coast of Me.xico and Central America. Part VHL Zoologica 34(4):239-258; pit. 1. (30 Dec.) Keen, A. Myra. 1958. Sea shells of tropical west America; marine mollusks from Lower California to Colombia, 1st ed. Stanford, Calif. (Stanford Univ.) .xi -i- 624 pp.; illust. (8 Dec.) 1966. Moerch's west Central America molluscan types with proposal of a new name for a species oiSernele. Calif. Acad. Sci., Occ. Papers 59:33 pp.; 41 figs. (30 June) 1971. Sea shells of tropical west America; ma- rine mollusks from Baja California to Peru, 2nd ed. Stan- ford, Calif. (Stanford Univ.) xiv + 1064 pp.; illust. (1 Sept.) Morch, Otto Andreas Lowson. 1859-1861. Beitrage zur MoUuskenfauna Central-Amerika's. Malak. Blatter 6(4): 102-126 (Oct. 1859); 7(2):66-96 (July 1860); 7(3):97-106 (Aug. 1860); 7(4):170-192 (Dec. 1860); 7(5):193-213 (Jan. 1861) Olsson, Axel Adolf. 1961. Mollusks of the tropical eastern Pacific particularly from the southern half of the Panamic- Pacific faunal province (Panama to Peru). Panamic-Pacific Pelecypoda. Ithaca, New York (Paleo. Resh. Inst.) 574 pp.; 86 pits. (10 March) Reeve, Lovell Augustus. 1853. Monograph of the genus Amphicksma. In: "Conchologia Iconica; or, illustrations of the shells of molluscous animals" 8. pits. 1-7 [pits. 1-4 (Oct. 1853); pits. 5-7 + title page & index. (Nov. 1853)] Reynell, Alexander. 1910. Further notes on the dates of issue of the parts of Sowerby's Conchological Illustra- tions. Malac. Soc. London, Proc. 9(3):212-213. (26 Sept.) 1918. The Index Testaceologicus of W. Wood & S. P. Hanley. Malac. Soc. London, Proc. 13(l/2):26-27. (9 Sept.) Shaw, Henry Otho Nicholson. 1909. On the dates of issue of Sowerby's "Conchological Illustrations," from the copy preserved in the Radcliffe Library, Oxford. Malax: Soc. London. Proc. 8(6):333-340 (5 Oct.) Sherborn, Charles Davies. 1909. On "The Conchological Illustrations," by George Brettingham Sowerby, Jun., London, 1832-1841, and the "Descriptive Catalogue of Shells," by John Edward Gray, 1832. Malac. Soc. London. Proc. 8(6):331-332 (5 Oct.) Sowerby, George Brettingham, II and George Brettingham Sowerby, I. 1833a [1832-1841]. The conchological illustra- tions; or coloured figures of all the hitherto unfigured Recent shells, . . . London (Sowerby) 200 plates in 200 parts, each with its own text, paginated separately and usually also reissued in 1841; species by G. B. Sowerby, I, except in certain groups. Concerning this work: Sherborn (1909). Shaw (1909), & Reynell (1910) [The section on Aniphidesma was first issued between 18 Jan. and 8 March 1833; the text was by G. B. Sowerby, I] Sowerby, George Brettingham, I. 1833b. Characters of new species of Mollusca and Conchifera, collected by Hugh Cuming. Zool. Soc. London. Proc. for 1832 [pt. 2] (25): 194-202 (13 March) NOTES ON NIDIFICATION AND OVULATION IN DRYMAEUS MULTILINEATUS (SAY) (PULMONATA, BULIMULIDAE) Jane E. Deisler Department of General Biology University of Arizona Tucson, Arizona 85721 During field work near Key West, Florida, Dryinaeuii muttilineatus (Say, 1825), an arbo- real snail native to Florida and the Caribbean, was observed nesting and laying eggs. No previ- ous reports on the reproductive behavior of this species are known, although observations on D. dormani (Binney, 1857), a species of northern and central Florida, have been reported (Muma Vol. 97(4) October 28, 1983 THE NAUTILUS 135 1955). The systematics and anatomy of the Florida species of Drymaeus are summarized by Pilsbry (1946, p. 21) and Breure and Eskens (1981). On 26 September 1982, during a heavy rain- storm, four specimens from a small colony of D. multilineatits were seen descending two trees (Bursera simaruha) on which they had been seen feeding earlier at the study site on Stock Island. The snails burrowed into the upper layers of the leaf mold immediately at the base of the trees once they had reached the ground. Burrowing continued until all but the extreme tip of the shells were covered loosely by leaf fragments, a depth of approximately 2 cm. Ovulation occurred over a period ranging from 18 to 22 hours subsequent to burrowing. The eggs were small and round with a maximum diameter of 2 mm. They were yellow-white and moderately hard but not calcareous and were deposited in a slightly sticky mass of 40-99 eggs. Two weeks later, on 9 October 1982, the nests were re-examined. It was found that all of the egg masses had shriveled up due to dessication. Similar shriveled egg masses were found throughout the study area, indicating that this may be a major source of mortality. This work was supported by funding from the Division of Sponsored Research, University of Florida (DSR Seed Grant A-1-26) and the U. S. Fish and Wildlife Service (Contract No. 85910- 0759), extended to Fred G. Thompson, Florida State Museum. LITERATURE CITED Breure. A. S. H. and A. A. C. Eskens. 1981. Notes on and descriptions of Bulimulidae (Mollusca, Gastropoda), II. Zoologische Verhandhngeru Leiden. No. 186(1981):1-111, pis. 1-8. IVIunia, M. H. 1955. Observations on the biology of the citrus tree snail. The Citrus Indushy 36(l):6-9 & 21. Pilsbry, H. A. 1946. Land Mollusca of North America (north of Mexico). Acad. Nat. Sci. Philad. Monogr. No. 3(2(1)): i-vi, 1-520. [pp. 21-29]. A NEW SINISTRAL TURRID FROM BRAZIL (GASTROPODA: TURRIDAE) Donn L. Tippett 10281 Gainsborough Rd. Potomac, Maryland 20854 ABSTRACT A new turrid species, Borsonia brasiliana, characterized by sinistral coiling is described and figured. A columellar plait and other features places it in the sub- family Borsoniinae and the genus Borsonia. The sinistral turrids are reviewed briefly. The discovery of a lot consisting of 144 speci- mens of an undescribed and unusual turrid in the collection of Recent mollusks of the U. S. National Museum of Natural History, Washing- ton, D. C, warrants the establishment of a new species. This is by virtue of its distinctive features including sinistral coiling, the presence of a columellar plait, and other aspects of shell morphology. The proposed taxon, Borsonia bra- siliana, is based on conchological characters as no animal material was available. A few dried animals were present, but attempts to recover 136 THE NAUTILUS October 28, 1983 Vol. 97(4) radular teeth by dissolving in KOH were un- successful. Subfamily: Borsoniinae Bellardi, 1875 Genus: Borsonia Bellardi, 1839 Borsonia brasiliana new species (Figs. 1-5) Description: Shell small (largest specimen 12.9 mm), sinistrally coiled, biconic fusiform, spire with a somewhat blunt apex, body whorl large and tapering gently to a moderately elon- gate open, weakly notched anterior canal. A small umbilical chink and a rounded anterior siphonal fasciole of variable strength are usually present in larger specimens. Columella slightly twisted to the right and bearing a narrow, mod- FKiS. l-.'j. Bor.sonia brasiliana new specicn. 1, 2, 3, Anterior, lateral and posterior niews of Holotype, USNM 810567, 11.9 mm length, Jt.3 mm width. 4, Specimen mth Up broken back showing columellar plait. 5, Specimen with enlarged siphonal fasciole. (Photos courle.'iy Smithsonian In- stitution. V. Krantz). erately elevated plait just above the center. The plait tends to be placed well back in the aperture but is usually easily visible. Occasionally it can be seen to terminate or fade within the recess of the aperture. It is occasionally weak or, rarely, rudimentary. The plait can usually be seen on earlier whorls when breaks or drilled holes per- mit viewing. A suggestion of a second, broader, fold below the plait is present in about 10% of specimens. The columellar lip shows a thin glaze of callus, margined below in large specimens. Protoconch of 2 to 2V2 smooth whorls with a slightly immersed and somewhat laterally placed tip. Protoconch usually preserved. Adult sculpture marked by the appearance of small ax- ial ribbing. Post-nuclear whorls 5 to 6, sharply angled just above mid-whorl by the ends of the axial ribs, and with a concave sulcus extending to the preceding suture. Suture minutely chan- neled producing the impression of a spiral sub- sutural thread. Sculpture of oblique axial ribs, 12 to 14 on the penultimate whorl. These are regularly spaced, about the same width of that of the interspaces, broader at the shoulder where they are angled and often finely noded or even slightly cusped. Although rather constant, the axials are reduced in strength occasionally, approaching peripheral nodulation only. This is never to the degree seen in Borsonia ceroplasta or B. silicea however. In some large specimens the axials become nearly obsolete on the last whorl. Posterior sinus occupying the whole of the shoulder slope, moderately deep, U-shaped. Lip thin, fragile, usually broken. No "stromboid notch" or varix. Color uniformly milky white, glistening and somewhat translucent when fresh. A few specimens contained dried animals retracted too deeply within the shell to observe for an operculum. Two specimens were sacri- ficed to attempt recovery of radular teeth. No operculum was identified on the broken pieces of animal thus obtained. Type locality: 200 miles north of Sao Luis, Brasil, on the edge of the continental shelf at 150 fathoms; lat. 00°18'N, long. 004°17'W. R/V Oregon, station 4226, Mar. 9, 1963, in one dredge haul using a six foot dredge. No record of substrate. Types: Holotype: USNM 810567, 11.9 mm Vol. 97(4) October 28, 1983 THE NAUTILUS 137 TABLE 1. Ratios of major shell diniensions in percentages. Based on measurements of 20 largest, in- tact (presumably adult) specimens. Range of measurements: Total length 10.9-12.9 mm: Maximum width i.0-5,1 mm: Body whorl length 6.2-8.5 mm; Length aperture plus canal 5.0-6.0 mm. Spire angle 27°-35°, mean 30°. standard deviation 2.1°. Max. width to Body whorl to A pert, plus canal total length total length to total length Range 34-40 55-69 42-52 Mean 36 64 46 Standard deviation 1.7 3.9 3.2 length, 4.3 mm width. Paratypes: USNM 818743 (119 specimens). Two additional para- types deposited at each of the following institu- tions: Academy of Natural Sciences of Phila- delphia, Pennsylvania; American Museum of Natural History, New York; Auckland Institute and Museum, New Zealand; British Museum (Natural History), London; California Academy of Sciences, San Francisco; Delaware Museum of Natural History, Greenville, Delaware; Los Angeles County Museum, Los Angeles; Museum of Comparative Zoology, Cambridge, Massachu- setts; Museum National d'Histoire Naturelle, Paris; Zoological Museum, Copenhagen; and the Museu Oceanografico, Rio Grande, Brasil. Remarks: The museum lot apparently repre- sents sampling of a homogeneous population. There is little variation among individuals. Range of variation of major dimensions is noted in Table 1. As can be seen, the greatest vari- ability is in the ratio of body whorl length to total length, although this is not evident simply by inspection. The most obvious variation, be- sides that described for the axial ribs, is in the strength of the siphonal fasciole. This is well marked in perhaps a dozen large specimens. It would appear to be a function of maturity, possi- bly a gerontic phenomenon, however other indi- viduals of equal size do not show equivalent en- largement. A few of the specimens with fascio- lar enlargement show a less well-developed plait and a slightly "fatter" shell outline, but there is no clear cut correlation between any grouping of shell characters suggesting distinct forms. About 60% of specimens are drilled, the hole be- ing typical of that made by naticids. Discussion Borsonia brasiliana is considered a member of the subfamily Borsoniinae on the basis of the presence of a columellar plait, which is the primary feature of the group with respect shell morphology. Placement in the genus Borsonia is based on similarity of shell structure to other members of the genus, all dextral, especially B. prima from the Italian Miocene. (Bellardi, 1839, p. 30). There are adequate specific differences to warrant the conclusion that Borsonia brasiliana is a distinct species and not a sinistral mutation of another. Similarity to Borsoyiia prima is evi- dent on comparison with that species. (Bellardi, 1847, pi. 4, fig. 13; Powell, 1966, pi. 8, figs. 9, 10). The two are reasonable "mirror images", however Borsonia prima has spiral sculpture, less well-developed axials, and is larger. A fur- ther modification of the axial ribbing in their reduction to peripheral nodules is seen in the Recent species of the genus from the Western Atlantic: Watson's Borsonia ceroplasta from off Puerto Rico, and B. silicea of Brazilian waters. (Watson, 1886, pi. 18, fig. 2, and pi. 21, fig. 8 respectively). Other differences are also pre- sent. Interestingly, a species widely separated geographically, Borsonia Jaffa, is the only other form lacking spiral sculpture. (Cotton, 1947, p. 14 and accompanying plate). Borsonia Jaffa shows peripheral nodules only and has an elon- gated anterior canal. A new sinistral form invites comparison with the other known sinistral turrids, although there is little similarity besides left-handed coil- ing. None are borsoniids. The genus Antiplayies Dall (1902, p. 513), a Recent and fossil group from the American west coast, has the sinus nearly on the periphery, no axial ribs, and a very different shell outline. It is a member of the sub- family Turrinae according to Powell (1966, p. 52). The genus contains a number of species 138 THE NAUTILUS October 28, 1983 Vol. 97(4) names many of which are undoubtedly syno- nyms and needs review. The gemmate Sinistrel- la Meyer (1887, p. 18), with two species, from the S. E. United States Eocene is also turrinine, having the sinus on the periphery. A left-handed species known only as Pleurotoma sinistraUs (Petit, 1839, pi. 1) from the Senegal coast has a rather pupoid shell outline, the sinus near the periphery, and no axials. It was figured by Reeve (1843, pi. 10, sp. 81) and Tryon repeated Reeve's illustration (1884, pi. 13, fig. 64). The figure is poor but nevertheless differs from Petit's in showing what appears to be a beaded subsutural cord. Perhaps two species are in- volved. The species should be investigated. Reeve's illustration of Hind's Conopleura striata (1846, pi. 36, sp. 330a), unnecessarily renamed partita, is sinistral but in error. The species is dextral. Reeve makes no mention of the shell be- ing sinistral and his other figure (sp. 330b) is dextral. Tryon (1884, pi. 8, fig. 7) continues the error, commenting that "one of Reeve's figures shows a reversed shell, a rarity in this genus". He recognizes Reeve's name as unnecessary. ACKNOWLEDGMENTS The author wishes to thank the NMNH for the opportunity of working with its magnificent col- lection, and particularly to express his apprecia- tion to Dr. Joseph Rosewater for his kind sup- port and assistance. Virginia 0. Maes was most helpful in her review and critique of the paper. Also Drs. A. W. Baden Powell and R. Tucker Abbott made valuable suggestions. LITERATURE CITED Bellardi, L. 1839. Bull. Soc. Geol. de France, vol. 10, 517 pp. 1847. Monografia delle Pleurotome Fossili del Piemonte. Mem. Roy. Accad. Sci. di Torino, 2nd ser., 9:3-122, 4 pis. Cotton, B. C. 1947. Some Southern Australian Turridae. S. Australian Nat. 24:13-16, 1 pi. Dall, W. H. 1902. Illustrations and Descriptions of New, Unfigured, or Imperfectly Known Shells, Chiefly Ameri- can, in the U. S. National Museum. Proc. USNM 24(no. 1264):499-566, pis. 27-40. Meyer, 0. 1887. Beitrag zur Kenntnis der Fauna des Altter- tiars von Mississippie und Alabama. Ber. Senck. natur. GeselL. 22 pp., 2 pis. Petit de la Saussaye, S. 1839. Guerin's Mag. de Zool, 2nd ser., vol. 1, pi. 1, accompanying text. Reeve, L. A. 1843-1846. Conchologia Iconica, vol. 1, Pleuro- toma, 40 pis., accompanying text. London. Tryon, G. W. 1884. Manual ofConchology. parts 23, 24, pp. 151-413. 34 pis. Philadelphia. Watson, R. B. 1886. Challenger Reports, Scaphopoda and Gasteropoda, pt. 42, vol. 15, 756 pp., 53 pis. A NEW SPECIES OF CATINELLA (SUCCINEIDAE): PULMONATA FROM SOUTHERN MICHIGAN Dorothea S. Franzen Illinois Wesleyan University Bloomington, Illinois 61702 ABSTRACT A neu> species o/Catinella (Succineidae) with its shell characteristics, reproduc- tive organs, pigmentation and, habitat is described. It is knovm only from the type locality. Long Lake, Cass County, Michigan. In the course of field studies on succineid gas- tropods in the midwestern states I found a small, slender, hitherto undescribed species of Catinella. Vol. 97(4) October 28, 1983 THE NAUTILUS 139 Catinella protracta n. sp. (Figs. 1 and 2) Description of holotype: Shell: (Fig. 1, A, B) Amber-colored, translucent, shining, imperfor- ate, slender, elongate, composed of S'A inflated, tightly twisted whorls separated by a sharply in- cised suture; height 7.8 mm, width 4.2 mm. A knoblike nuclear whorl tops the turreted spire; whorls increase rapidly in size resulting in a tumid ultimate whorl. Ovate aperture equals about six-tenths of height of shell (Table 1). Sharply edged peristome very fragile. Very slender, amber columella follows inner border of peristome, curves as it disappears into the ultimate whorl (Fig. 1 B). Nuclear whorl finely wrinkled and pitted. Remainder of shell surface finely striated resulting in a shining appearance. Body and Mantle Surfaces: Surface of head and body white, irregularly tuberculate. Supe- rior (posterior) tentacles finely tuberculate. Pig- mentation lacking from tentacles, dorsal and lateral surfaces of body. Genital aperture cres- cent-shaped, about 0.7 mm in length, situated on anterior right-hand side of body. On either side of body a pedal groove, continuous from labial palp to posterior tip of body, separates foot from lateral body wall; a suprapedal groove parallels the pedal groove. Shallow, vertical grooves in- cise the pedal and suprapedal grooves. These vertical grooves produce shallow scallops along the margin of foot and along the body wall especially when the animal is in a somewhat con- tracted state. Sole of foot white and unpig- B FIG. 1 . A, B, Holotype o/Catinella protracta n. sp. (Height 7.8 mm); Paratypes o/ Catinella protracta n. sp; C, D (Height, 7.5 mm); E (Height, 9.1 mm); F (Height. 7.2 vim); G, H (Height. 7.0 mm). 140 THE NAUTILUS October 28, 1983 Vol. 97(4) s^ ^.g- n p. < < 2 X f— 1 v^ .—1 0) D.J=I < U) 2 3 >— ( ■>^ I — 1 CD CXXI < CO X X t4H 0) O ^1 3 s: ■M +-> ^1 -o 0 ■ H n S < ^ O OJ M +-> 3 c. 4-> M h ■ r4 (I) (1< p. X < ■P J= £ 4-' M •n •H ■H C) 2 X X. 4-" •o •H 3 ■»-» r: M • H <1) X 1« in n .— < U n o z rOL0OJLnv£»000000O O— I lD'-hO'— (O^~lOC0^- ■^sDOC0O^0Ln^0O CTiLnfNLnLOfN^oor-- OOfNOOOOO'— <0'-HCTi rjoo t-O t^ rj ro ri r-o t-O Kl rj rj rO O'-OLOoof^iLoi^Lorj LOO LOLOLOt-OLOsOLTJI-OLn I— tOOOOI^^DLOLT)^ '^rsirsj'^'^'ct'-a-'"*-'^- rOPOtOKitOtotOtOt^ to j= >, • o LO ♦J u. ■H t-~ •H • >; CT) ^ in T-* n! • 0) M 0) O Q J< n & o * n! o LO _1 ~1 I J ■(-» -0 o (D ^ at ul X Q. 01 r. i/^ o >-. -H o rt 3 X H u. -J u "O o o LO O ■-HrjO'TfOCnOf"!'— t LO'^^'^'^rtrt-rj-^ LO CTt ^c fN.— (■— <\OOOr--rOOOLO^ \D^OlO\D«O^^OnD''0^ ■-H^LOooora-r^or^L/i oo^o co<— iCTiLOtOf-HOiLOLorg -^-i- LO^UOOO—i.— I.— , T) (U c in ^ a> o oJ 3 s U- _1 U -3 •<:r CM rg to LO LO to LO Oi \J~i to -^ Tj- O LO 00 0) bO o: •H 1) Vol. 97(4) October 28, 1983 THE NAUTILUS 141 mented. Mantle collar and transparent mantle are unpigmented. Colors of some of the viscera seen through the mantle and translucent shell: the elongate cres- cent-shaped, light golden nephridium; the slight- ly darker golden digestive gland; an irregularly- shaped, netted, brown band located on the body along the anterior and posterior borders of the second body whorl. Holofype: Catalogue No. FMNH 205821. Paratypes No. FMNH 205822, Molluscan Col- lection, Field Museum of Natural History, Chicago, Illinois. Additional paratypes in the private collection of the author. Description of Paratypes: Shell; (Fig. 1, C, D, E, F, G, H) Shells of mature snails attaining a height of 9.1 mm are comprised of 3 - 3V2 in- flated whorls. Dimensions of the ten largest shells of the two series included in this study, number of shells measured and the median of each series measured are recorded in Table 1. Of the ten largest shells of the two series the range of height is 7.0 - 9.1 mm; the range of the width is 3.8 - 5.1 mm; the largest apertures occupy from 54.5 to 70.5 percent of entire height of shell; the median of widtli/height are: 1967, .565, of 1975, .552. Other dimensions and rela- tive dimensions are also to be noted. A SEM photograph of a nuclear whorl (Fig. 2) shows the surface to be finely wrinkled and pit- ted. Granules of sand and/or soil are frequently adherent to the shell. Body, mantle and viscera: The white body, mantle collar and mantle are usually unpig- mented. On some individuals scattered black flecks occur on body wall, mantle collar, mantle and sole of foot. Occasionally part of edge of sole of foot is tinted with black pigment. There is an absence of patterns of pigmentation as observed in other species of Catinella: C. parallela Franzen (Franzen, 1979, p. 64); C. vagans (Pilsbry) and C. waccamawensis Franzen (Franzen, 1981, pp. 118, 121). The transparent membrane of the floor of the mantle cavity is sometimes pigmented with small brown flecks. A broad, irregularly shaped, netted, brown band is located on the body along the posterior and anterior borders of the second body whorl; the anterior (lower) band is frequently the larger of FIG. 2. Scanning-electron-photomicrograph of the nuclear whorl o/Catinella protracta n. sp. the two. Color of viscera (seen through mantle and shell): the elongate, crescent-shaped, cream or light to darker golden colored nephridium is not outlined by a band of pigment characteristic of other species of Catinella; the digestive gland varies from a cream color to a golden brown; the gut twines around lobes of the digestive gland. Reproductive System: (Fig. 3) The albumin gland (AG) triangular, composed of fine acini, enclosed within a thin, transparent, unpig- mented sheath, seminal vesicle (SV) elongate, subequally bilobed, enclosed within a thin sheath FIG. 3. Reproductive organs o/ Catinella protracta n. sp. P, penis: EP, epiphalhis: PA, penial appendage: PRM, penial retractor muscle: SPD, spermathecal duct: SP. spermatheca: HD. hermaphroditic duct: AG, albumin gland: GA, genital atrium: VA, vagina: OD, oviduct: VD, vas deferens: PG, pro- state gland: FS. fertilization scu;: SV, seminal vesicle. 142 THE NAUTILUS October 28, 1983 Vol. 97(4) sparsely pigmented with brown flecks. The her- maphroditic duct (HD), sparsely pigmented with brown flecks, and the seminal vesicle join to form the fertilization sac (FS) from which a duct divides to form the convoluted oviduct (OD) which leads to the vagina (VA) and the sperm duct (SD) which leads into the prostate gland (PG). The oval prostate gland (PG), enclosed within a thin, transparent, unpigmented sheath, is composed of small acini slightly larger than those of the albumin gland. As the vas deferens approaches the penis (P) from the prostate gland it follows the penis along its dorsal surface and enlarges to form the epiphallus (EP) which enters the distal end of the unsheathed penis. The penis enlarges immediately into an elongate cylindrical form. The penial appendage (PA) is spherical; its base less than half the length of the penis; the vertical dimension is almost twice the body of the penis. Penial retractor muscle (PRM) is broad; its fibers insert onto the epiphallus, penis and penial appendage. The globular spermatheca (SP) is connected to the vagina by an elongate, slender, spermathe- cal duct (SPD). The short vagina (VA) expands as it enters into the genital atrium (GA). Radula and Jaw: Radulae of three paratypes were mounted and stained. The number of rows of teeth occurring on the radulae examined range from 93 to 95. There are few teeth on the anterior-most rows; the number increases rapid- ly posteriorly. The number of marginals and laterals of representative rows of those radulae are recorded in Table 2. The characteristics of the teeth of Catinella protract a n. sp. are described as follows and illustrated in Fig. 4A. The central tooth (C) bears a broad, long basal plate having a pos- terior serrated margin flanked on either side by a rounded boss. The pointed mesocone extends downward to about the lower fourth or beyond the basal plate. A short, pointed ectocone flanks the mesocone on either side. The laterals (2-L-L, 5-L-L) have a pointed mesocone varying in length from a short structure (probably worn) to many extending downward to the posterior margin of the basal plate. The mesocone is flanked medially by a small, pointed endocone and laterally by a larger, undivided, pointed ec- TABLE 2. Formulae of representative rows of teeth of Catinella protracta n. sp. from two s'pecimens, field no. 359, Long Lake, Cass County, Michigan. Slide No. of Rows of Teeth Row M L C L M A 95 20 7 - 9 - 1 - 7 - 6 30 8 - 9 - 1 - 8 - 7 50 9 - 8 - 1 - 8 - 8 55 8 - 9 - 1 - 8 - 8 60 9 - 8 - 1 - 8 - 8 66 9 - 8 - 1 - 8 - 8 B 93 30 9 - 9 - 1 - 8 - 10 31 10 - 8 - 1 - 9 - 9 35 iO - 8 - 1 - 8 - 10 40 9 - 9 - 1 - 9 - 9 45 9 - 8 - 1 - 9 - 9 50 9 - 9 - 1 - 9 - 9 59 9 - 9 - 1 - 9 - 9 70 10 _ 8 - 1 - 9 - 9 tocone. The basal plate of the outermost laterals is shorter than of the more medial laterals. The marginals (1-L-M, 4-L-M, 6-L-M) smaller than the laterals have a short basal plate which is broader than long, especially true of the outer- most marginals. The small endocone is pointed. The larger, pointed mesocone extends to or beyond the posterior margin of the basal plate. The ectocone of the inner-most marginals is divided into two, that of the outermost into L-L-5 FIG. 4. A: Representative radula teeth o/Catinella protracta n. sp. C. central tooth; L-L-2, 2nd left lateral: L-L-5, 5th left lateral: L-M-1, 1st left marginal: L-M-i, J,th left marginal: L-M-6, 6th left marginal. B: A jaw o/ Catinella protracta n. sp. Vol. 97(4) October 28, 1983 THE NAUTILUS 143 three, four or five cusps. The distinction be- tween the laterals and marginals is not always clear because a tooth with a relatively short basal plate whose ectocone is divided into two cusps may be flanked on either side by one whose ectocone is undivided, or a tooth with a relatively long basal plate may have an ectocone divided into two cusps. The structural details of the teeth of Catinella protracta n. sp. compare with those of the species of the genus, namely (1) the ratio of the laterals to the marginals approximates 1:1 and (2) the short basal plate of the marginals is broader than long. Such features of the genus were noted by Quick to be true of Catinella (Suc- cinea) armaria ("B.-Ch.") (Quick, 1933, Fig. 4, p. 296) and by Franzen of C. parallela Franzen (Franzen, 1979, p. 66, Table 2; p. 67, Fig. 3A); of C. vagans (Pilsbry) Franzen, 1981, p. 120, Table 2 and Fig. 3; of C. waccam.awensis Franzen (ibid, p. 122, Table 4; p. 123, Fig. 6A). The jaw (Fig. 4B) is amber colored. Anteriorly the collar has a median, bluntly pointed fold. The median indentation of posterior edge of col- lar is flanked on either side by a lesser indenta- tion. Geographic Distribution and Habitat: Cati- nella protracta n. sp. known only from type locality. Type locahty: Field No. D.S.F. 359; Long Lake, Cass County, Michigan, Sunset Boulevard, 0.4 mi S of U.S. Hwy 12, west shore of north end of lake. The locality, an unimproved portion of the shore. Just beyond the collecting site a point of land extended into the lake. July 14, 1967: C. protracta n. sp. found living on and under boards of a broken dock and on wet ground at base of and among roots of sedges and reeds. The area, unshaded. July 15, 1975: C. protracta n. sp. was living on wet (soggy) ground under matting of dead sedges as was, also, Oxyloma retusa (Lea). Distinctive Features: Shell: Comprised of 3 -3V2 inflated, tightly coiled, sharply incised whorls, attaining a height of 9.1 mm; spire elon- gate. As shown in Table 3 comparisons of ratios of height of aperture to height of shell of three species of Catinella verify that the shell of C. protracta n. sp. is comparatively narrower and the spire relatively longer. Comparative data TABLE 3. Compaynsons of ratios of shell dimensions of four species o/ Catinella. Width Height H. of Aperture H. of shell Range Median C. waccamawensis Franzen C. vagans {PilsbryJ .50 .492 .58 .567 .585 .56 .53 .49 .474 .518 515 552 565 68 .647 663 .619 679 .632 62 .59 643 .597 645 .60 Range 701 705 75 797 819 72 711 70 528 - 658 606 - .711 .563 .655 .726 .720 Median .611 .611 .69 .766 .777 .67 .67 .64 .577 .653 taken from Franzen, 1979, p. 65; 1981, pp. 118, 121. The specific name protracta refers to the com- paratively narrower and relatively longer or protracted spire as noted above. Pigmentation: A pattern of pigmentation on the mantle, head, dorsal and lateral body sur- face lacking which is in contrast to characteris- tic patterns of other species of Catinella noted by Franzen: C. parallela Franzen (Franzen, 1979, p. 64); C. vagans (Pilsbry) and C. wacca- mawensis Franzen (Franzen, 1981, pp. 118, 121). Sometimes scattered flecks of black pig- ment are present on head, body and sole of foot. The nephridium not outlined by a black band such as noted to be present in the above listed species. Pigmentation of the sheath covering seminal vesicles, fertilization sac, and oviduct are lightly pigmented with brown flecks. A broad, irregu- larly-shaped, netted brown band is located on the body along the posterior and anterior bor- ders of the second body whorl. The conspicuous band is seen through the shell. Reproductive System: Prostate gland equal in size to, or larger than, albumin gland. Seminal vesicles elongate, bilobed, subequal in length. Penial appendage large, inflated, spherical. ACKNOWLEDGMENTS National Science Foundation Grants-in-Aid No's NSF G18000 and NSF GB2715 provided laboratory equipment. Dr. A. Byron Leonard 144 THE NAUTILUS October 28, 1983 Vol. 97(4) read the manuscript and offered helpful sugges- tions. LITERATURE CITED Franzen, Dorothea S. 1979. Catinelh parailela, a New Succineidae (PuJmonata) from Midwestern United States. The Nautilus 93(2-3):63-69. Tables 1-2. Figs, 1-3. 1981. Catinella vagans (Pilsbry) and a New Species of Catinella (Succineidae Pulmonata) from the Shore of Lake Waccamaw, North Carolina. The Nautilus 95(3): 116-124. Tables 1-4, Figs. 1-5. Quick, H. E. 1933. The Anatomy of British Succineae. Proc. Mai. Soc. London 20(6):29.5-318, pi. 23-25, tables 1-5, figs. 1-18. NEW RECORDS OF INDO-PACIFIC MOLLUSCA FROM COCOS ISLAND, COSTA RICA Donald R. Shasky 834 W. Highland Avenue Redlands, California 92373 In April of this year. Captain Richard Calla- way, of Balboa, Panama, and I spent 6V2 days SCUBA diving for mollusks at Cocos Island, ap- proximately 300 miles south by southwest of Puntarenas, Costa Rica. Dives were made from the Victoria, an 82 foot motor-schooner based at Puntarenas, Costa Rica. On my return to the Costa Rican mainland, I met Dr. Michel Montoya, who has a paper in press titled, "Los Moluscos de la Isla del Coco, Costa Rica. Lista Anotada de E species." His paper is a complete literature review list- ing 16 species of bivalves, 89 gastropods, 4 chitons, and 9 cephalopods. No scaphopods or nudibranchs have been reported from the island. This is a total of only 118 species. Dr. Montoya, who also spent 6V2 days diving at Cocos Island, in June, and I, are now prepar- ing our own check-list which will add approxi- mately 100 additional molluscan species to the known Cocos Island marine fauna. The new Indo-Pacific records that we found at Cocos Island are: Viriola abboffi (Baker and Spicer, 1935) ScaienoHtoma subulata (Broderip, 1832) Cypraea (n. sp.) Burgess, 1983- in press (Venus) Ckaronia tritonis (Linnaeus, 1758) Favartia garretti (Pease, 1868) Persicula pulcheila (Kiener, 1834) Spondylus nicobaricus Schreiber, 1793. (Syn: S. histrix Roding, 1798) Vi7-iola abbotti was described from Samoa, and has recently been reported living in Hawaii. A single dead specimen was found. Scalenostoma subulata has, according to Waren, 18 synonyms. It has been reported in all tropical seas except for the eastern Pacific. A single live specimen of Charonia tritonis was taken at 40 meters. A previously unre- ported Charonia tritonis from the Galapagos is cited in a letter dated August 26, 1965, from Mrs. Carmen Angermeyer to William Old at the American Museum of Natural History. Mrs. Angermeyer purchased this shell from Jorge Pincay, who collected it in 2 meters of water just north of Punta Mangle, Fernandina Island. Mr. Pincay was a crew member of the Charles Dar- win Research Station's vessel, Beagle. Favartia garretti, has up until now, been known only from the Hawaiian Islands. Numer- ous specimens were taken at Cocos Island, under dead coral at depths of 13-26 meters. I have had an unidentified Favartia in my collec- tion from La Cruz de Huantecoxtle, which is ap- proximately 30 miles north of Puerto Vallarta, Mexico. It appears to be this species. The Persicula pulcheila was a single dead specimen. Vol. 97(4) October 28, 1983 THE NAUTILUS 145 The Cypraea (n. sp.) Burgess, 1983, has been examined by Dr. Burgess. He examined my specimen after this new species was already in press. He has informed me that it is a fairly widespread species being found as far west as Australia. Dr. William Emerson has cited my specimens of Cypraea talpa Linnaeus, 1758, in a compan- ion paper in this issue of the Nautilus. Captain Callaway and I took 7 specimens in depths of 7-14 meters. Although I have not had comparative mate- rial, the specimens of Spondylus nicobaricus from Cocos Island, seem to match the size, col- or, and hinge serrations as outlined by Dr. Kay, for S. histr-ix. I wish to thank Dr. William Emerson for allowing me to read the correspondence be- tween Mrs. Angermeyer and William Old and Dr. Emerson. I also wish to thank Dr. Montoya for sharing the results of his trip with me and to the crew of the Victoria and to Mary Crowley of Ocean Voyages, Sausalito, California, for mak- ing our trips possible. I also wish to thank Dr. R. Tucker Abbott for pointing out that Spondylv^ histrix is a junior synonym of S. nicobaricus. LITERATURE CITED Baker, F. and V. D. P. Spicer. 1935. New species of mollusks of the genus Triphora. Trans. San Diego Soc. Nat. Hist. 8(7):35-46, 5 pi. Burgess, C. M. 1970. The Liviyig Courries. pp. 381, pi. 44. A. S. Barnes and Company. Emerson, W. K. and William E. Old, Jr. 1964. Additional records from Cocos Island. The Nautilus 77{3):90-92. Hertlein, L. G. 1937. A note on some species of marine mol- lusks occurring in both Polynesia and the western Amer- icas. Proc. Amer. Phil Soc. 78:303-312, 1 pi. 1963. Contribution to the biogeography of Cocos Island, including a bibliography. Proc. California Acad. Set., ser. 4. 32(8):219, 4 figs. Kay, E. A. 1979. Hawaiia7i Marine Shells, pp. 653, fig. 195. Hawaii: Bishop Museum Press. Keen, A. M. 1971. Sea Shells of Tr-opical West America (2nd ed.) California: Stanford University Press. Pease, W. H. 1868. Synonyms of marine gasteropodae in- habiting Polynesia. Aynerican J. Conchology 4:103-132. Sowerby, G. B. (2nd of the name) 1842-1887. Thesaurus conchyliorum: or monograph of genera of shells. 5 vols. London. Waren, A. 1980. Revision of the genera Thyca, Stilifer, Scalenostoma. Mucronalia. and Echineulima (Mollusca, Prosobranchia, Eulimidae). Zoologica Scripta 9:187-210, fig. 107. OCCURRENCE OF THE UNIONID, ANODONTA IMPLICATA SAY, IN NORTH CAROLINA Rowland M. Shelley North Carolina State Museum of Natural History P.O. Box 27647 Raleigh, North Carolina 27611 ABSTRACT Authentic valves of the naiad, Anodonta implicata Sa^, have been collected along the Chowan River in North Carolina approximately 5 miles south of the Virginia state line, a range extension of some 175 miles from the Potomac River. The mollu^k is considered a "threatened species" in Noi'th Carolina. Previous publications have reported the range of Anodonta implicata Say as being from Nova Scotia and New Brunswick, Canada, to the Potomac River of Maryland and Virginia (Johnson 1946, 1970; Burch 1975). Records by Athearn and Clarke (1962) and Fuller (1977) of material from South Carolina and the upper Cape Fear River basin of North Carolina, 146 THE NAUTILUS October 28, 1983 Vol. 97(4) FIG. 1. Anodonta implicata Say J'rimi. the Chownn River, North Carolina. respectively, are erroneous and were based on heavy specimens of A. cataracta Say with distinct pallial lines, the latter also having discolored nacre similar to that of implicata (Johnson 1970, personal communication). How- ever, I can now positively report the occurrence of implicata in North Carolina from four valves I collected on the east bank of the Chowan River in Gates County, 5.5 miles NW of Eure, on 16 September 1971. The valves agree with John- son's descriptions (1946, 1970) in the color of the nacre and the thickening of the anterior margin below the pallial line, and match known speci- mens of implicata from Massachusetts. They are deposited in the North Carolina State Museum invertebrate research collection under catalog numbers 1151 and 1621. This site, which is about 5 miles south of the Virginia state line, represents a range extension of around 175 miles and indicates that the mollusk can be ex- pected in Coastal Plain parts of the intervening river systems of Virginia. Fuller's report (1977) was part of the North Carolina Endangered Species Symposium, where implicata was assigned to the "undeter- mined" category because of insufficient data to assess its statewide conservation status. Since so little of the Chowan drainage is in North Carolina and this is its known southern limit, implicata should properly be regarded as "threatened" in the state, based on the criteria of the symposium's mollusk committee. This designation is substantiated by the occasional eutrophication-algal bloom problems in the Chowan River not far from the sample site, as these could have deleterious effects on the molluscan fauna. Extensive collections in pied- mont sections of the Cape Fear, Neuse, and Tar- Pamlico drainages have produced many speci- mens of cataracta but none of implicata, and if the latter is in these systems, it probably is in the Coastal Plain. It is also a potential inhabi- tant of the Coastal Plain part of the Roanoke drainage, located between the Chowan and Tar- Pamlico. ACKNOWLEDGMENTS I am grateful to Richard I. Johnson, Museum of Comparative Zoology, for confirming my identification of the Chowan specimens of im- plicata, for comparative material from Massa- chusetts, and for advice on Fuller's record. The accompanying figure was kindly prepared by Mary Kay Clark of the North Carolina State Museum. LITERATURE CITED Athearn, Herbert D. and Arthur H. Clarke. Jr. 1962. The freshwater mus.sels of Nova Scotia. Natl. Mus. Canada, Bull. No. 183, Cont. Zool., 1960-61:11-41. Burch, John B. 1975. Freshwater Unionacean Clams (Mol- lusca: Pelecypoda) of North Am£rica. Rev. ed.. Malcologi- c