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THE NAUTILUS

Volume 102 1988 AUTHOR INDEX

154 99 Lee. H. G. 78, Absalao, R. S. Mackie. L 73 Adams, W. F 125 G McKiNSTRY. D M 127 Askew, T. A. 89, 92 McLean, H 99 AUFFENBERG, K 40. 154 J. W. 129 AUFFENBERC, T 40 \1endl. 1 1 MlKKELSEN, P M. BlELER, R. 131 115 Moore, D. R. Boss, K. J. W. 102 p 149 Myers, B. bouchet, 47 o'sullivan, j. B. Collins, T. M 102 Pawlik. R. 47 D'AsARO, C. N 1.34 J. Petit, R. E. 130, 164 DAtiilio, A 106 E. 65 H. 30 Pip, DuShane, 129 36, 110 POLACO, O. J Emerson, VV. K. 92 125 POMPONI, S. .\. Gerberich, a. G. 131 123 RoMER, N. S. Gordon, M. E. III 36, 110 Sage, W E , Haimovicl M. 82 Santos Cruz, R. L. 99 , 92 DOS Harasewych, M. G. 56 Smith, DC 155 Hayek. L. C 54 56 Snyder, MA Hershleb, R. 78 164 Thompson, F. G. Kabat, a. R. 102 73 Vermeij, G. J KiLGOUR, B V\'.

NEW TAXA PROPOSED IN VOLUME 102 (1988)

GASTROPODA 89 charlestonensis Askew, new species (Pleurotomariidae) Perotrochus 100 McLean. Absalao and Santos Cruz, new species (Turbinidae) Macrarcnc difiilala 102 Wrmeij and Collins, new species (Neritidae) Nerita fortidentala 131 Romer and Moore, new species (Rissoidae) Alvania colombiana 106 shaskyi DAttilio and Myers, new species (Muricidae) Faiartia (Murexiella) 150 Bouchet, new species (Buccinidae) Metula crosnieri 151 new species (Buccinidae) Metula africana Bouchet, 54 Sn\der, new species (Fasciolariidae) Latirus martini 36 Emerson and Sage, new species (Turbinellidae) Vasum Stephana 130 new name (Cancellariidae) Axi'lella Petit 154 Auffenberg and Lee, new species (Terebridae) Terebra imitatrix 78 Hypselostoma holimanae Thompson and Lee, new species (Pupillidae) CEPHALOPODA 82 Eledone gaucha Haimovici, new species (Octopodidae) rHE NAUTILUS

Volume 102, Number 1 February 16, 1988 ISSN 0028-1344

A quarterly devoted to malacology.

Marine Biological Laboratory LIBRARY

FEB 2 4 1988

Woods Hole, Mass. EDITOR-IN-CHIEF Dr. Aurele La Rocque Dr. Geerat J. V'ermeij Dr. M. G. Harasewych Department of Geology Department of Biology Division of Mollusks The Ohio State L'niversitv University of Mar\ land National Museum of Columbus, OH 43210 College Park, MD 20740 Natural History Smithsonian institution Dr. James H. McLean Dr. Gilbert L. Voss Washington, DC 20560 Department of Malacology Division of Biology and Living Los Angeles County Musetim of Resources ASSOCIATE EDITOR Natural History Rosenstiel School of Marine and 900 E.xposition Boulevard Dr. R. Tucker Abbott Atmospheric Science Los Angeles, CA 90007 Universit) of American Maiacoiogists, Inc. Miami P.O. Bo.x 2255 4600 Rickenbacker Causewav Melbourne, FL 32902 Dr. Arthur S. Merrill Miami, FL 33149 % Department of Mollusks CONSULTING EDITORS Museum of Comparative Zoology SUBSCRIPTION INFORMATION Harvard University suKscription rate Dr William K. Emerson The per volimie Cambridge, MA 02138 is for individuals Department of US $20.00 and Living Invertebrates US $30.00 for institutions. Postage Dr. Donald R. Moore outside is The American Museum of the United States an Division of Marine Geologv Natural Historv additional US $2.00 for surface and Geophysics New York, NY 10024 and US $10.00 for air mail. All Rosenstiel School of Marine and orders should be accompanied by Atmospheric Science pavment and sent to: THE Mr. Samuel L. H. Fuller University of Miami 1053 Mapleton .Avenue NAUTILUS, P.O. Box 3430, Silver 4600 Rickenbacker Causeway Spring, 20901. Suffiekl. CT 06078 MD Miami, FL 33149

Dr. Robert Hershler Change of address: Please inform Mr Richard E Petit the publisher of address Division of Mollusks \our new P.O Box 30 at least 6 weeks in advance. All National Museum of North Myrtle Beach, SC 29582 Natural History communications should include both old new addresses (with Smithsonian Institution and Dr. Edward Petuch Washington, DC: 20560 J. zip codes) and state the effective Department of Geology date. P"lorida .Atlantic L'niversitv Dr. Richard S. Houbrick I?6ca Raton,' "Fl' 33431 0028- Division of Mollusks THE NAUTILUS (ISSN is published quarterly National Museum of 1344) by Dr. G. Alan Soleiri Natural History Trophon Corporation, 363 Department of Invertebrates Silver Spring, Smithsonian Institution Crescendo Way, MD Field Museurri o£ Natural Historv Washington, DC 20560 20901. Chicago, IL 60605

Second Class postage paid at Silver Mr. Richard I. Johnson Dr. David H. Stansber\ additional mailing Department of Mollusks Spring, MD and Museum of Zoology Museum of Comparative Zoology offices. The Ohio State Universitv Harvard I'niversity Columbus, OH 43210 Cambridge MA 02138 POSTMASTER; Send address changes to: Dr. Ruth D Turner THE NAUTILUS Department of Mollusks P.O. Box 3430 Museum of Comparative Zoolog\ Silver Spring, MD 20901 Harvard Universit\ Cambridge, MA 02138 THEt^NAUTILUS

Volume 102, Number I February 16, 1988 ISSN 0028-1344 CONTENTS

Rudiger Bieler Aiiatoni) and reproductive biology of two western Atlantic Paula \1. \likkelsen species of N'itrinellidae, with a case of protandrous hermaphroditism in the Rissoacea

Helen DuShane Geographical distribution of some Epitoniidae (Mollusca: Gastropoda) associated with fungiid corals 30

\^ illiam K. Emerson A new species of Vastim (Gastropoda: Turbinellidae) from Salter E. Sage, HI off Somalia 36

Kurt Auffenberg Density, spatial distribution, activity patterns, and biomass Troy Auffenberg of the land snail, Geophoriis bothropoma Moellendorff (Prosobranchia: Helicinidae) 40

Notice 46

Marine Biological Laboratory LIBRARY

FEB 24 1988

Woods Hole, Mass.

THE NAUTILUS 102(l):l-29, 1988 Page 1

Anatomy and Reproductive Biology of Two Western Atlantic Species of Vitrinellidae, With a Case of Protandrous Hermaphroditism in the Rissoacea

Kiidiger Bieler Paula M. Mikkel^en Smithsonian Marine Station Harbor Branch Oceanographic at Link Port Institution 5612 Old Dixie Highway 5600 Old Dixie Highway Ft Pierce, FL 3-1946, USA Ft. Pierce, FL 34946, USA

ABSTRACT and studies on the gross morphology of Cochliolepis parasitica Stimpson, 1858 (by Moore, 1972) and C. Two western Atlantic vitrinellids, Cyclostremiscus beauii albicerata Ponder, 1966 (by Ponder, 1966), published (Fischer, 1857) and Circulus texanus (Moore, 1965) new com- information on anatomy is available for only one species bination are redescribed based on a study of live snails from of this family, Circulus striatus (Philippi, 1836) from the burrows of the stomatopod crustacean Lysiosquilla scabricauda (Lamarck, 1818). Anatoms and reproductive biology are the eastern Atlantic. Data on Ci. striatus, extensively emphasized, with the first recorded description of spawn and presented by Fretter (1956) and later summarized by larval development in the famiK'. Synon\mies are given, and Fretterand Graham (1962, 1978), were based on material a lectotype is selected for Cy. beauii. Literature data concerning which Fretter and Graham (1978:228) described as "the anatom) are reviewed for the marine near-planispiral risso- only live specimens . . . obtained by Fretter (1956) from aceans; Vitrinellidae (including Cyclostremiscus and Circulus, the stomach of the starfish Astropecten . . . dredged on with Circulidae as a synonym) is considered distinct from Tor- sandy bottoms 28-30 m deep in the Gulf of Gascony." nidae. Unusual morphological aspects of Cy. beauii are dis- Thus, the present concept of vitrinellid anatomv' is cussed, including external ciliation patterns, pallial tentacles based mainly on Fretter 's description of a single species, (which are functionalK and morphologically different from striatus, the species of the name-bearing each other), and stomach morpholog>' (with a large posterior Circulus tvpe chamber). Protandrous sequential hermaphroditism in Cy. beauii genus of the nominal family Circulidae (see Discussion, is inferred (for the first time in the Rissoacea) from morpho- below). Discussions of phylogenetic relationships (or s\n- logical evidence of sex change correlated with size. onymies) between families such as Vitrinellidae and Tor- nidae have demonstrated the need for anatomical and Key words: Cyclostremiscus; Circulus; Vitrinellidae; Torni- for Fretter, dae; Rissoacea; systematics; anatomy; hermaphroditism. reproductive data these groups (e.g., 1956; Tavlor & Sohl, 1962; Moore, 1965; Golikov & Starobo- gatov, 1975; Boss, 1982; Graham, 1982; Ponder, in press). INTRODUCTION Studies on two species of western Atlantic N'itrinellidae are reported herein. Populations of Cyclostremiscus Despite the great number of nominal species assigned to beauii (Fischer, 1857) and Circulus texanus (Moore, 1965) the poorK defined, cosmopolitan, marine rissoacean fam- were discovered in Florida in sand-flat burrows of the ily \ itrinellidae Bush, 1897, little is known about the stomatopod crustacean Lysiosquilla scabricauda (La- biology of its members. Their small size, mostly unknown marck, 1818). These burrows are U-shaped and extend habitats, poor representation in collections, and frequent up to 1.5 m into the sediment, with horizontal distances confusion with other small-shelled members of groups of 6-7 m between the two openings; a pair of stomatopods such as Cyclostrematidae Fischer, 1885, Skeneidae Thiele, inhabits each burrow system, maintaining it over long 1929, Turbinidae Rafinesque, 1815, and Tornidae Sacco. periods of time (Serene, 1954; R. B. Manning, personal 1896, may account for the lack of attention paid to this communication). Cyclostremiscus beauii, with a shell family, and why authors who have attempted revisions diameter of 6-8 mm, is one of the largest vitrinellids, a (e.g., Trvon, 1888; Bush, 1897; Melvill, 1906; Pilsbry & fact that facilitated detailed studv of its morphology and

Olsson, 1945; Pilsbry & McGinty, 1945a,b, 1946a,b, 1950; anatomy and allowed a test of Fretter s hypothesis that Laseron, 1958; Moore, 1964; Adam & Knudsen, 1969) some of the characters found in the much smaller (2 mm) relied almost entireK' on shell characters to distinguish Circulus striatus are size-related (Fretter, 1956:380). ta.xa on all taxonomic levels. Special emphasis was placed upon characters of the

Aside from Pilsbry and McGinty 's (1945a, 1946b) reproductive svstem, for which Moore (1964:18) noted sketches of crawling animals of several nominal genera. "Nothing is known of the reproduction of the famiK b

Page 2 THE NAUTILUS, Vol. 102, No. 1

except that the animals are dioecious, and that the male shells, radulae, jaws, and opercula were coated with gold/ is provided with a penis." Data presented herein suggest palladium, and scanned using a Zeiss Novascan-30. Fig- that Cy. beauii is a protaiidrous sequential hermaphro- ures 1-3 were photographed using a Hitachi S-570 scan- dite. Only incomplete data are available for Circulus ning electron microscope. Radular terminology is after texanus, as animals of this species were collected only Bandel (1984:3). twice, both times before the actual beginning of this Protoconch and teleoconch diameters were recorded study. However, since gross anatomy, spawn, and de- as the greatest dimension perpendicular to the columellar veloping eggs were observed for this species, available axis. Teleoconch height was the greatest dimension par- data are presented here. allel to the columellar axis, measured from the apex to the base of the aperture. Umbilical diameter was the greatest distance between the columellar lip and the most MATERIAL AND METHODS prominent portion of the umbilical wall, measured in

ventral \ iew . Teleoconch w horls were counted from the Samples were taken from stomatopod burrows in shal- protoconch Il-teleoconch line to the farthest extent of low-water sand flats in the Indian River lagoon just inside the periphery (= the point on the outer lip used for the Ft. Pierce Inlet, St. Lucie County, eastern Florida greatest shell diameter). The number of protoconch whorls (27°28.3'N, 80°17.9'W) using a stainless steel bait pump was determined b\ the method of Ta\ lor (1975:10; sum- ("yabby pump") in conjunction with sieves of 1-2 mm marized by Jablonski & Lutz, 1980:332, fig. 4). mesh. Depths at low tide ranged from less than 0.5 m Cited repositories are (* indicates location of voucher to supratidal, wherein the water level lay several centi- material): meters below the level of the sand. Living snails were maintained in Bnger bowls of sea- ANSP—Academy of Natural Sciences of Philadelphia, water at room temperature (24 °C). Carmine and fluo- PA. rescein sodium particles were used to observe ciliary CAS— California Academy of Sciences, San Francisco. action and currents produced by the animals. For gross *IRCZM— Indian River Coastal Zone Museum, Harbor dissections, shells were cracked and animals subsequently Branch Oceanographic Institution (HBOI), Ft. Pierce, relaxed using magnesium sulfate crystals ("epsom salts"). FL. Other anaesthetic chemicals (7% magnesium chloride in MCZ—Museum of Comparative Zoology, Harvard Uni- distilled water, menthol crystals) were tried but pro- versity, Cambridge, M.A. duced little or no effect with gradual addition, or too MNHN—Museum National d'Histoire Naturelle, Paris. strong an effect resulting in retraction. Methylene-blue/ *RSMAS—Rosenstiel School of Marine and Atmospheric basic-fuchsin and neutral red were used to better delin- Sciences, University of Miami, FL. eate tissues and organs in gross dissections. Structures UNC-IMS— Institute of Marine Sciences, University of and organs were measured following in part the outline North Carolina, Morehead City, NC. given by Davis and Carney (1973: fig. 4A). Terminology *USNM— National Museum of Natural History, Smith- of the nervous system is after Davis et al. (1976). sonian Institution, Washington, DC. For histological sectioning, animals were relaxed as above and fixed in either glutaraldehyde-formalin solution (4% formalin, 2.5% glutaraldeh\de in 0.1 M phos- RESULTS phate buffer, pH 7.2) or 5% buffered formalin (Humason, Rissoacea Gray, 1847 1962:14). Shells were either broken and removed, or dissolved in a 1% solution of ethylene diamine tetraacetic = Truncateilacea Gray, 1840, "submission to be made to ICZN acid (EDTA, adjusted to pH 7.2). Specimens were to suppress this name" (Ponder, 1985:15). embedded in paraffin, sectioned at 5-7 ^m and stained with Alcian Blue/ Periodic- Acid-Schiff (PAS), counter- Vitrinellidae Rush, 1897 stained with Harris' Hematoxylin/Eosin (Humason, 1962: 125, 269, 298). Staining reactions described in the text Cyclostremiscus Pilsbry & Olsson, 1945:266. refer to this method unless otherwise rioted. Photomi- Type species: VitrineUa panamensis C B. Adams, 1852 crographs of sections were taken with a Zeiss Photomi- (by original designation). croscope-3. Radulae and jaws were extracted by dissolving the surrouiitling soft tissue in a solution of lO'^ sodium hy- Cyclostremiscus beauii (Fischer, 1857) droxide. Spermatozoa were prepared for SEM by placing (figures 1-52; tables 1, 2) a drop of concentrated sperm in seawater onto a coverslip Adeorbis Beauii Fischer, 1857a:173 [nomen nudum]. placed in a covered petri dish containing droplets of 25% Adcorlns Reauii Fischer, 1857b:286, pi. 10, fig. 12 [Guade- glutaraldeliyde, and passing the coverslip through an loupe].— Bu.sh, 1897:104. in etlianol series ending acetone, and then critical-point Cyclostrema Beam [sic].— A. Adams, 1866:251, pi. 255, fig. 25 drying the sample. Whole animals were fixed, passed [after Fischer, 1857b]. through an ethanol series, transferred to amyl acetate, Cyclostrema bicarinata Guppy, 1866:291, pi. 17, figs. 5a, and critical-point dried. These, together with air-dried ["Miocene" (Lower Pliocene), Jamaica]. R. Bieler and P. M. Mikkelsen, 1988 Page 3

Skenea sulcata "Bush" Simpson, 1887:61 [nomen nudum, see Table 1 . .Anatomical characters and character states, and hab- Moore, 1964:131], itat types of species in the vitrinellid-toriiid complex Adeorlris Beaui.—DaW, 1889:150; 1892:345. Adeorbis Beam var bicarinata —DaW. 1903:1595 ["Oligo- 1) Projecting snout bilobed, lateral extensions: (a) absent; (b) cene," Jamaica]. present. "C.irculus" Incarinatus. — Woodring, 1928:439, pi. 37, figs. 10- 2) Cilia along cephalic tentacles: (a) absent; (b) present 12 [neotype designation]. 3) Terminal stiff setae on cephalic tentacles: (a) absent; (b) "Adeorhis" heaui. — Woodring, 1928:440. present. C.irculus stirophorus M. Smith, 1937:67, pi 6, figs. 2a, b [Pho- 4) Number of pallial tentacles. cene, Florida]. 5) Pallial tentacles: (a) all finger-shaped; {b) upper finger- Cyclostrema angulata. — Hertlein & Strong, 1951:110 [West shaped and lower paddle-shaped. Indies]. 6) Upper pallial tentacle: (a) naked; (b) with motile cilia; (c) Cyclostremiscus {Ponocyclus) heaui Incarinatus. — Pilsbry, 1953: with stiff setae; (d) with distinct motile cilia and/or stiff 427, pi. 55, figs. 1-le. setae. Vitrinella iSolariorbis) fceaui.— Abbott, 1954:138. 7) Lower pallial tentacle: (a) naked; (b) with motile cilia; (c) Vitrinella beaui. — Wells et ai. 1961:267. with stiff setae; (d) with distinct motile cilia and/or stiff Cyclostremiscus beaui. — Moore, 1964:131-135. — Morris, 1973: setae. 138, pi. 40, fig. 19.—Porter, 1974:143.—Emerson & Ja- 8) Gill filaments: (a) projecting from aperture in crawling cobson, 1976:64, pi. 18, fig. 21 —Abbott & Dance, 1982: position; (b) not projecting from aperture in crawling po- 58, text-fig. sition. Cyclostremiscus {Ponocyclus) beaui. —Warmke & Abbott, 1961: 9) Anterior foot margin: (a) straight or only weakly indented; 60, pi. 11, fig. b.—Humfrey. 1975:76, pi. 3, figs. U-lla. (b) cleft. Cyclostremiscus (Ponocyclus) beauii. — Abbott, 1974:85, text- 10) Posterior foot margin: (a) simple and rounded or weakly fig. 786.— Rios, 1975:38, pi. 10. fig. 131; 1985:41, pi. 16, indented; (b) cleft.

fig. 177.— Yokes & Yokes, 1983:15, pi. 25, figs. 4-4a. 11) Operculum nucleus: (a) concentric; (b) subcentral. 12) Number of whorls on operculum. Material examined: Lectotype (designated herein): 10.5 13) Eyes: (a) distinctly developed; (b) lack nerve supply. (Guadeloupe). of mm, MNHN unnumbered Neotype 14) Osphradium: (a) small, ciliated groove; (b) distinctly de- Cyclostrema bicarinata. 7.4 mm, USNM 115621 (Plio- veloped, paralleling ctenidium. cene, Jamaica); other material: 60 specimens: NORTH 15) Penis: (a) simple, without glandular processes, recurved;

CAROLINA: 1 specimen with dried animal, BEVER- (b) with glandular area, directed straight back; (c) with

IDGE Sta. I (ex Astropecten). UNC-IMS. FLORIDA: Ft. several, finger-like processes. Pierce Inlet: 10 March 1987, 1 male, 3 unsexed; 2-3 May 16) Habitat: (a) under rocks; (b) under scales of annelid Poly- odontes lupina (Stimpson); (c) ? from stomach of starfish; 1987, 1 male; 24 June 1987, 3 males; 3 August 1987, 6 (d) in burrows of stomatopod Lysiosquilla scabricauda (La- males; 31 August 1987, 7 females, 1 1 males; 1 4 September marck); (e) sandy mud bottom; (f) under large boulders 1987, I female; 27 September 1987, 3 females, 2 males. on well-ox\genated sand\ mud. Peanut Island, Lake Worth Inlet: 11 August 1987, 1 female, 1 male. Boynton Beach: 1 shell, ANSP 277740.

Miami: 1 shell, EOLIS Sta. 311, USNM 449192. Fowey Light: 2 shells, EOLIS Sta. 187, USNM 449193; 1 shell', 5-8) on first whorl, becoming more dense and regularly EOLIS Sta. 129, USNM 449194; 1 shell, EOLIS Sta. 142, spaced on second whorl. Fields of pustules on body whorl USNM 449195; 1 shell, EOLIS Sta. 170, USNM 449196; intersected by smaller concentric ridges, 4-6 between 1 shell, EOLIS Sta. 355, USNM 449198. Turtle Harbor: major ribs, added first between suture and first major 2 shells, EOLIS Sta. 61, USNM 449197. Sand Key: 1 shell, rib, subsequently between peripheral ribs. Strong cords EOLIS Sta. 162, USNM 449199. Kev West: 1 shell, EO- forming sharp peripheral and basal keels, separated by LIS Sta. 63, USNM 449200. Drv Tortugas: 1 shell, USNM wide flat area, inclined ventrally and marked by growth 61114; 1 shell, USNM 271949. Cape San Bias; 1 shell, lines, spiral cords, and pustules. Base widely umbilicate, USBF Sta. 2402, USNM 323914. CARIBBEAN: St. Mar- with 0-5 concentric ribs between basal keel and umbi- tin; 1 shell, ANSP 20621. Jamaica: 1 shell, USNM 426872; licus; larger specimens showing decreasing number of 1 shell, USNM 442372. SOUTH AMERICA: CoveSas, ribs with increasing size. Umbilical wall often with 2-4 Colombia: 1 shell, USNM 364409. narrow ribs. Outer lip sinuous, with shallow sutural sinus. Microstructure (figure 4) of 3 layers: 2 thick cross-la-

Description mellar layers and 1 thin homogenous outer layer thickened to form spiral ribs. Teleoconch (figures 1-3): Shell large for family [usually 2-3 whorls, 6-8 mm diameter (x = 6.9, n = 56); Protoconch (figures 9-11): Diameter 0.40-0.48 mm

height: X = 3.8, n = 33; umbilical diameter: x = 1.5, n = (f = 0.45, n = 32). Protoconch I (prior to hatching) 30; largest specimen (female): diameter 11.5 mm, height smooth, of about 1 whorl (diameter ~ 0.23 mm). Pro- 6.2 mm, umbilicus diameter 2.4 mm, teleoconch whorls toconch II (after hatching, before settling) of an addi- 3%], nearly planispiral. Opaque-whitish, with 5-8 strong, tional whorl, sculptured with irregular, more-or-less con- concentric ribs on apical side, above peripheral keel. centric markings (figure 11). Total protoconch of 2 whorls, Widely-spaced irregular pustules between ribs (figures rather high-spired (spire angle = 50°). Page 4 THE NAUTILUS, Vol, 102, No. 1

Table 2. Summary of anatomical and habitat data for species in the vitrinellid-tornid complex. Characters and character states ** are listed in table 1. Sources of data listed under each species. = type species of genus; * = synonym of type S[)ecics of genus. *' = c according to VUmre. 1964:22; d according to Moore. 1964:159 W] biological data from Florida, except for Cochliolepis parasitica (also South Carolituil. Co. alhiccrata (-\e« Zfalaiid ' Cirrulus sirialiis (iucrnseN. English Channel

Characters

Species 12 3 4 5 6 8 9 10 11 12 13 14 15 16

Croup 1 **Vitrinella helicoidea C. B .\dunis, 1850 Pilsbry & McCinty, 194.5a: pi. 2, fig. 5; 1946b: 13. a b b 2 adaaaa? ?a??a *Vitrinella praecox Pil.sbr\ & McGinty, 1946 Pilsbry & McGinty, 19'45a: pi. 2, fig. 4; 1946b:14. a b b 2 aadaaa? ?a??? Teinostoma carinacallus Pilsbr\ & McGinty, 1946 Pilsbry & McGinty, 1946b:17, pi. 2, fig. 6b. b 2 adaaaa? ????a Teinostoma lerema Piisbrv & McGintv, 1945 Pilsbry & McGintv, 194.5a:6, pi. 2, fig. la. 1, 2 a^d'aa?? ????a Teinostoma parvicallum Piisbrv 6< McGint>, 1945 2.' Pilsbry & McGinty, 194.5a:4. pi 2, fig. addaaa? ????a **Pleuromalaxis halcsi (Piisbrv & McGinty, 1945) Pilsbry & McGinty, 1945a: 10, pi. 2, fig. 8. a b adaaa?? ????a **Cochliolepis parasitica Stirnpson, 1858 Stimpson, 1858:307ff.; Moore, 1972:100ff. a a b 2 aabaaaa ?a?ab Cochliolepis allncerata Ponder, 1966 a Ponder, 1966.38, pi 5 a b b 1 a — — baaa 6a??a *Circultts striatus (Philippi, 1836)

Fretter, 1956:369ff ; Fretter & Graham, 1978:227£f. a a b 2 accbaaal2abac Circtilus texanus (Moore. 1965) This paper a b b 2 acbaaaa 8abad Cyclostremiscus beaitii (Fischer, 1857) This paper b 2 bcbaaaa 7abad Cyclostremiscus pentagonus (Gabb, 1873) Bush, 1897:127, pi. 22, figs. 6, 12a-g [as Skenea trilix]. ? 9 ?????? ao o a 7 ?

Group 2

**Tomura hicaudata (Piisbrv & McGintv, 1946)

Pilsbry & McGinty, 1945a: pi. 2, fig.' 9; 1946b:15. b ? ? ? ? *Parviturhoides interruptus (C. B. .\dams, 1850)

Moore, 1962:695ff., fig. IB; 1964:21, 156ff.; 1972:106ff., figs. 5, 6. a b b 2 a dbaba ?a?b Group 3 **Tornus subcarinatus (Montagu, 1803) Woodward, 1898:140ff., pi. 8, figs. 1-3, 5-7; Fretter & Graham. 1978:229ff; Graham, 1982:144ff. bb2aaaaaab 'b

External anatomy and organs of the mantle cavity (fig- rated Irom remainder ot snout bv strongiv ciliated groove. ures 12- 18, 20-25, 27, 34-37): Living animal trans- Serial sections revealed 2 narrow longitudinal bands of lucent vellovvish-orange, with buccal mass, anteriormost cilia, situated in grooves on each side, beginning shortly gill tilaments, and tip of penis rose-pink; digestive gland behind tip of snout. Two slender, De.xible, cephalic ten- brownish-orange. Base of ctenidial lamellae on osphra- tacles reaching appro.ximately twice length of snout when (iial side pigmented white; central area of osphradium fullv extended. Left tentacle fitting into shallow notch vvliitf Hanked bv brown lines on each side, resulting in formed bv ventral shell keel (figure 14). iMack eves on pattern of parallel lines on left body side (figure 12). outer expanded bases of cephalic tentacles, each equipped Long snout terminating in pair of muscular "lips," sep- with spherical, transparent lens. arated by vertical slit with mouth opening. Lips sepa- Ciephalic tentacles exhibiting elaborate pattern of mo-

Figures 1-11. Cyclostremiscus heauii. specimens from Ft. Pierce Inlet, Florida (SEM) (figures 1-3, USNM 846323). 1. Shell, apical view (8.0 mm diameter). 2. Shell, umbilical view (5.0 mm diameter) 3. Shell, apertural view (4.1 mm diameter) 4. Microstructure of lateral body wall; fracture surface parallel to growing edge 5. Teleoconch sculpture, apical view. Circled numbers indicate location of sculptural details in figures 6-8. Stars indicate location of sutures. 6. Detail of teleoconch sculpture, first whorl. 7. Detail of teleoconch sculpture, second whorl. 8. Detail of teleoconch sculpture, third (= body) whorl. 9. Protoconch, apical view.

,\rro\v indicates sculptural line between protoconchs I and II. 10. Protoconch, lateral view. 11. Sculpture of protoconch I (left, smooth) and protoconch II (right, sculptured) Scale bars: 4, 5, 8-10 = 0.1 mm; 6, 7, 11 = 10 fim. Page 5 R. Bieler and P. M. Mikkelsen, 1988

11^ Page 6 THE NAUTILUS, Vol. 102, No. 1 R, Bieler and P. M. Mikkelsen, 1988 Page 7

lilc cilia and stiff bristles. Ventral tip of each tentacle ctenidium, attached along entire length to inner surface with L-shaped, heaviK -ciliated groove surrounding of mantle, originating on posterior left, curving over smooth area, forming tactile pad (figure 25, tp). Nu- dorsum, terminating just above right eye where last few merous stiff bristles (lost during fixation, therefore not gill filaments protrude from aperture. Numerous (< 180) evident in histological or SEM preparations) distal to and gill filaments (figures 16, 21) flattened, elongated leaflets, just behind pad; additional bristles sparsely distributed almost finger-like when contracted, forming tapered over distal quarter of tentacle. T\\ o longitudinal ciliated blades when extended. Filaments longest in central part tracts, situated in grooves, extending from region of pad of gill, decreasing somewhat in size toward both ends. on ventral side. Near tentacle base, innermost groove "Supporting rods" lacking; filaments well supplied with sloping dorsally toward dorsal midline; outermost groove blood spaces and muscles (transverse muscle bands giving ending proximal to expanded tentacle base (figure 20). extended filaments ladder-like appearance; figure 16). [•fattened base of tentacle facing snout surface covered Both sides of blade carrying wide band of cilia off-center, b\ additional, irregular, parallel tracts of cilia (figure 20). closer to left (osphradial) side. Along right side in same Third longitudinal ciliated groove on dorsal side of ten- relative position, each filament with longitudinal row of tacle extending from flattened, triangular area just below small embedded crystals (figure 16, cr). Narrow rim clear, eye to near tentacle tip (figure 18). Ciliated grooves usu- somewhat thinner on "crystal" side, bearing continuous alK lined b\ narrow bands of brown pigment. Cilia fulK' band of cilia. Filaments draining into large efferent bran- retractable into grooves. C^ilia in grooves with distinct chial vessel leading to heart (figure 12). Filaments hardly movement pattern (right tentacle: dorsal row—toward reacted to direct physical stimuli, contracting rapidly tentacle base, inner and outer ventral rows—toward tip; when either cephalic tentacle or finger-shaped pallial left tentacle: opposite directions). (These movements are tentacle (see below) touched. easily mistaken for direction of ciliary beating, and there- Whitish hypobranchial gland (figures 12, 13) paral- fore current flow. However, further microscopic obser- leling entire length of rectum, most conspicuous poste- vation with the aid of carmine and fluorescein sodium riorly. Osphradium (figures 12, 22, 23) paralleling almost particles revealed lateral beating of the individual cilia entire left side of gill, comprised of wide central area across the tentacle, away from the snout, indicating that and two strongK developed, heavily ciliated, lateral zones. the apparent ciliary movement actualK reflects the con- Central area w ith irregular chevron-like pattern of tracts duction of nervous impulses along the tentacle.) In cross- of shorter cilia. Osphradial ganglion (figure 12, osg) very section, each tentacle usually with 3 nerves, 1 larger conspicuous at point about 'A of total length from mantle blood sinus, and several smaller blood spaces in central edge. Mantle cavity ending immediately behind poste- area, especially around nerve cords (figure 24). rior end of ctenidium. Foot (figures 12-14) elongate (just reaching posterior Two pallial tentacles (figures 13, 27) arising from just shell margin in crawling animal), flattened, densely and inside right mantle edge; upper (= most dorsal) tentacle finely ciliated, with anterolaterally recurved corners; finger-shaped, unciliated, somewhat closer to mantle edge, broadh rounded posteriorly and slightK indented at pos- curling into shallow sutural sinus, directed dorsally. Low- terior terminus. Anterior pedal mucous gland (figure 14, er tentacle paddle-shaped, ciliated at edges (except on amg) present, opening at center of transverse slit across narrow stalk) and also across its broad surface; narrow anterior, leading edge of foot. No posterior mucous gland band at tentacle edge unciliated. Single stiff bristles some- or metapodial tentacles. Locomotion by ciliary action. times occurring at tip. Lower tentacle originating at point Operculum (figures 17, 34-37) corneous, circular, mul- on mantle edge ventral and slightly more interior to tispiral, with about 7 whorls and small internal knob, upper finger-shaped tentacle; directed anterolaterally. supported by finely ciliated opercular lobes (figure 17, Stimulation with forceps or needle caused immediate ol) on dorsal side of foot. Lobes simple, unpigmented, contraction of upper tentacle (followed by contraction without tentacles. Lemon-shaped muscle scar on internal of cephalic tentacles and exposed ctenidial filaments). surface of operculum (figure 35). Lower tentacle showed little response to touch and can- Mantle edge (figures 12, 13) somewhat scalloped, cor- not markedly contract (compare tentacles in figure 27). responding to spiral ribs of shell. Large monopectinate Lower tentacle observed to regulate and enhance water

Figures 12-17. Cyclostremiscus beauii. 12. Male, left side, in crawling position (shell removed). Penis reflected anteriorly, out

of mantle cavitv . 13. Female, right side, in crawling position (shell removed). Mantle slightly reflected to show relative insertion points of pallial tentacles. 14. Crawling animal, ventral view. 15. Diagrammatic cross-section through mantle cavitv of male, at level of osphradial ganglion. 16. Gill filament. 17. Diagrammatic view of closed operculum, as seen when animal is retracted, showing position of opercular lobes. Scale bars: 12-14 = 1.0 mm; 15 = 0.5 mm; 16 = 1 mm. ag, albumen gland; amg, anterior mucous gland; br, immobile bristles; cit, ciliary tract; com, columellar muscle; cont. connective tissue; cr, crvstals; ct, ctenidium; dg, digestive gland; ebv, efferent branchial vessel; es, esophagus; he, heart; hg, hypobranchial gland; in, intestine; ki, kidney; Ipt, lower pallial tentacle; mc, mantle cavity; me, mantle edge; ol, opercular lobe; op, operculum;

OS, osphradium; osg, osphradial ganglion; ov, ovary; pe, penis; pvd, pallial vas deferens; re, rectum; sh, shell; sn, snout; st, stomach, ten, cephalic tentacle; upt, upper pallial tentacle. Page 8 THE NAUTILUS, Vol. 102, No. 1 H. Bieler and P. M. Mikkelsen, 1988 Page 9

longitudinal, dorsal folds, their lower exten- tlow , by paddle positioning and by strong ciliary action, muscular, it'spccti\el\-, in or out of right side of mantle cavity. sions coiling upwards to form .semi-isolated compart- ments, .interior esophagus thus divided into strongly cil- Alimentary system: Mouth opening between pair of iated dorsal food channel (figure 26, dfc), larger ventral muscular lips into large buccal mass (figure 42; length channel, and two small lateral pockets. Posterior to nerve

1..') mm in specimen 8.5 mm shell diameter). Radula ring (figure 42), ventral projections into main channel protrmling from short radular sac (figure 42, ras) ex- increasing gradualK in size, but without clear distinction tending somewhat behind and to left of buccal mass. between anterior and mid-esophageal regions. Food Paired jaws (figure 41) each crescent-shaped, appro.x- channel remaining dorsal. Dorsal folds and ventral pro- imately 0.5 x 0.16 mm (in specimen 7.8 mm shell di- jections gradually decreasing in size, with posterior ameter), composed of interlocking diamond-shaped ele- esophagus as a simple muscular tube. ments 14 /im in length. \ar\ ing slightK in shape across Stomach (figures 12, 38-40) amber in color, slightly surface of jaw. translucent in li\'ing animal, approximately 4 x 1.5 mm Radula (figures 28-33) taenioglossate, with about 100 (in largest specimen, 11.5 mm shell diameter), lying on rows (max. 138; n = 7), length == 1.3 mm, width ~ 0.2 exterior surface of visceral coil, appressed to the surface mm. Rachidian tooth (figure 31) wider than long (0.40 of digestive gland just posterior to heart and kidney, /im w ide. n = 8), with acute posterior corners projecting encompassing approximately Vi total length of digestive laterally, and concave front. Main cusp narrowly-trian- gland. Stomach consisting of two continuous chambers gular, unserrated, with 4-5 flanking cusps on each side (figure 39) differing in function: anterior chamber ('/a of (number of flanking cusps var\ing within single radula, total stomach length) containing gastric shield, working apparently by splitting and fusion), decreasing laterally end of cr\stalline style, and all openings into stomach; in size; base with 1 strong basal denticle per side, midway remaining -/3 forming large sorting and storage chamber. between posterior corners and central ridge. Lateral tooth Esophagus entering stomach on left side at junction of ifigures 32, 33) with asymmetrical cutting edge, strongly anterior and posterior chambers. From this point, series indented at front edge, bearing large, narrowK -trian- of folds extending transversely across stomach, poste- gular, unserrated main cusp and highly variable number riori) into posterior chamber, and anteriorU toward in- of flanking cusps (3-6 inner, 7-8 outer) decreasing in testinal opening. Opening to digestive gland lying to size laterally. Base of lateral tooth with broad central right of esophageal opening, between it and gastric shield. ridge; basal platform long, blade-like. Apex of inner mar- Gastric shield (figures 39, 40), with cup-like lateral wing ginal (figures 30, 33) with short, stout main cusp, and upon which crystalline style rotates, protruding into an- 12-14 subequal, inner and outer flanking cusps. Stalk terior chamber, and central longitudinal portion with long, blade-like, with robust supporting ridge. Apex of flattened lateral expansion that cradles style, positioning outer marginal (figures 28, 30) with sharp, undulating it against cup-like grinding surface. inner margin (sometimes finely incised into closely-ad- Style sac and intestine, at anterior end of stomach, hering flanking cusps), and smooth, rounded outer mar- usualK partially obscured b\ connective tissue and kid- gin. ney. Style sac (figure 39, ss) narrow, finger-shaped, ap-

Esophagus opening widely into buccal cavity, without proximately '/3 length of stomach, not communicating esophageal pouches. Salivary glands (figures 19, 26, 42) directly with intestine [Johansson's (1940:1) group 3, re- narrow, tube-like, empt\ ing into buccal mass at its lateral vised after Mackintosh (1925)]. Style completely trans- mid-point, extending posteriorly along esophagus. Posi- parent, rod-shaped (length 1.8 mm, diameter 0.35 mm), tion of salivar\ ducts relative to circumesophageal nerve rotating within style sac by action of denseK packed cilia ring highly variable: some or all passing through ring on style sac walls, protruding into anterior chamber of

(n = 3; figure 49), stopping just anterior to ring (n = 4), stomach through flesh\ tube-like structure above gastric or extending past without going through ring (n = 2). shield. Channel extending betw een digestive gland open- Relati\ e lengths of sali\ ary glands vary. Anterior part of ing and intestinal opening at anterior terminus of anterior esophagus, in cross-section (figures 26, 43), bearing 2 chamber.

Figures 18-27. Cyclostremiscus beauii, specimens from Ft. Pierce Inlet. Florida (light micrographs or critical-point dried SEM preparations). 18. Head-foot (male), with mantle edge reflected posteriorly, dorsal view (SEM). 19. Sagittal section of male through buccal mass and esophageal region. 20. .\nterior view of snout and left cephalic tentacle, showing pattern of ciliated tracts on ventral surface (SEM). 21. Tip of gill filament ^SEMl. 22. Osphradium, anterior to osphradial ganglion (SEM). 23. Osphradium, cross-section through osphradial ganglion 24. Cephalic tentacle, cross-section. 25. X'entral tip of cephalic tentacle, showing tactile pad (SEM). 26. Esophagus, cross-section through anterior section, with salivary glands. 27. Pallial tentacles: upper, finger-shaped tentacle (retracted), and lower, paddle-shaped tentacle (SEM). Scale bars; 18 = 0.5 mm; 22 = 50 ^m; 19, 20, 23, 24, 26, 27 = 0.1 mm; 21, 25 = 20 ^m. bm, buccal mass; bs, blood space; car, cartilage; cit. ciliar\ tract; cont. connective tissue; df, dorsal folds of esophagus; dfc, dorsal food channel of esophagus; es, esophagus; leg, left cerebral ganglion; Ipt, lower pallial tentacle; me, mantle edge; ne, nerve; os, osphradium; osg. osphradial ganglion; pe, penis; ra, radula; rs, receptaculum seminis; sag, salivary gland; sn, snout; tp, tactile pad; upt, upper pallial tentacle. Page 10 THE NAUTILUS, Vol. 102, No. 1 R. Bieler and P. M. Mikkelsen, 1988 Page 11

Posterior chamber (figures 39, 47, pch) with irregular Kidney opening at posteriormost end of mantle cavity, longitudinal folds along its left side leading from esoph- without conspicuous ciliated tract associated with open- ageal opening, and well-defined longitudinal groove on ing. No gonopericardial or renogonadial ducts observed. its right side leading toward gastric shield. Central area Nervous system: (iircumesophageal ganglia (figures 42, \\ ith series of ciliated transverse folds. 43, 49) moderately concentrated. RPG ratio [Davis et Proximal portion of intestine (figure 39, in) consisting ai, 1976:263; defined as length of pleurosupraesophageal of 3 histologicalK separate sections: (1) slightly bulging connective/(length of connective + length of right pleu- section immediately adjacent to stomach, containing ral ganglion + length of supraesophageal ganglion)] av- large typlilosole, leading into second section via small eraged 0.49 (n = 7). Cerebral ganglia connected by nar- loop; (2) very muscular, ball-shaped section passing un- row commissure, each separated from pleural ganglia by der style sac to join third section; and (3) moderately constriction. Tentacular nerves with distinct swellings at ridged intestine proper, initially quite narrow, dilating their bases. Pedal ganglia each with paired connectives, slightly as it passes, without further loops or undulations, connecting anteriorly to cerebral ganglia, posteriorly to toward rectum in mantle cavity. Anus set back from cerebropleural junctions. Pedal commissure very short, mantle edge. No special ciliated tracts from anus to man- narrow. Subesophageal ganglion somewhat smaller than tle edge and exterior. supraesophageal ganglion; connective to its pleural gan- Observed flow of particles {figure 39, smalt arrows): glion much shorter. Statocysts (figure 44, stc) about 110 Large and small food particles enter stomach through )xm diameter. Buccal ganglia small, conspicuous, joined esophagus. Counter-clockwise whirlpool at esophageal by commissure passing beneath esophagus at posterior opening preliminariK' sorts particles according to size. end of buccal mass. Smaller particles pass laterally to right in groove toward Highly-vacuoled connective tissue surrounding nerve gastric shield and style. Large particles move into pos- ring, as well as other organs and areas throughout body. terior chamber along left series of longitudinal folds, passing to far posterior terminus of stomach. From there, Reproductive system: .Animals in male phase (for dis- particles pass into central area where peristaltic action cussion of sex change, see below) distinguished from manipulates and returns large particles to anterior cham- functional females by smaller size and by dark-orange, ber. ConcurrentK . smaller particles separate and/or break rather than creamy-orange or beige, gonadial coloration. oft, and follow transverse folds toward right longitudinal Penis large (size at rest: length 2.5-3.0 mm, width at groove, and then anteriorly within groove toward gastric midlength 0.4 mm), muscular, somewhat flattened (fig- shield. Large particles in central area of sorting chamber ures 12, 18, 42, 44) arising just behind and slightly right continue moving anteriorly to region of esophageal open- of bases of cephalic tentacles, coiling counter-clockwise ing, where they pass rapidly by ciliary action directly to back into mantle cavity. Fully extended penis may be intestinal opening. Small particles entering area of gastric longer than cephalic tentacles {e.g., 4.4 mm vs. 3.0 mm). shield are manipulated by clockwise-rotating style against Subcentral penial duct terminating in opening on slightly cup-shaped flange of gastric shield. Resultant particles hooded tip, which, unlike remainder of penis, is uncil- move directly left into opening of digestive gland, guided iated. Long, closed pallial vas deferens extending from by cuticularized lateral folds near style sac opening. Un- penis, along right side of body, to prostate in posterior acceptable particles and material returning from diges- part of mantle cavity (figure 44). Vas deferens relatively tive gland are shunted via longitudinal groove toward wide (diameter ~ 0.2 mm), tubular, forming prominent, intestinal opening. Ball-shaped proximal section of in- somewhat undulating ridge that differs from surrounding testine probably serves as a pellet compressor. tissue by its shiny, unciliated surface and by its white Live specimens fed on single-celled algae and detritus appearance caused by heavy internal ciliary action. Pros- scraped from laboratory- aquaria walls. Fecal pellets oval, tate orange, egg-shaped, 0.65-0.90 x 0.35-0.48 mm (n = 0.33 X 0.19 mm (n = 5), round in cross-section, with 5), lying ventral to rectum, connected to right pallial rounded ends. vvall, its lumen communicating with mantle cavity by slit (~ 0.35 mm length) at its base (figures 44, 48). Vis- Renopericardial system: Two-chambered heart and ceral vas deferens much narrower, passing through pos- surrounding kidney visible on left surface of visceral coil, terior mantle wall, leading along inner coil of visceral posterior to ctenidium and hypobranchial gland (figure mass, where widened, extensively-coiled portion func- 12). Kidney large, with nephridial gland on its outer wall. tions as vesicular seminalis (figure 47, vs) before reaching

Figures 28-37. Cyclostremiscus beauii. radula and operculum, specimens from Ft. Pierce Inlet, Florida (SEM). 28. Outer marginal teeth. 29. Radula, whole mount 30. Inner marginal teeth. 31. Rachidian teeth. 32. Lateral teeth. 33. Tips of lateral (left) and

inner marginal (right) teeth 34. Operculum, outer surface (2 1 mm diameter). 35. Operculum, inner surface (1.7 mm diameter). 36. Operculum, oblique lateral view of inner side (2 2 mm diameter). 37. Opercular peg, oblique lateral view. Scale bars: 28, 30- 33 = 10 Mm: 29, .37 = .50 fim.

rac, rachidian teeth. Vol. No. 1 Page 12 THE NAUTILUS, 102,

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pispec mpg es PP9 R. Bieler and P. M. Mikkelsen, 1988 Page 13

testis. Testis simple, elongated sac, yellow to bright or- In ripe females, unoriented sperm, oltcn in large quan- ange, along inner right side of visceral coil, totalling 50- tities, found throughout length of sperm channel and in 60% of its length (figure 44). anterior sperm pouch (functional bursa copulatrix). Spermatozoon (56-62 nm total length, n = 3) with Packed oriented sperm, with heads embedded in walls, slightly twisted head comprising elongated nucleus and found in the two posterior pouches (receptacula seminis). pointed acrosome (~ 14% of total length), very long, Visceral oviduct very thin-walled, leading from coiled narrow midpiece (« 58%) and long tail (* 28%). Atyp- oviduct to ovary. Ovar\ situated at right side of visceral ical sperm not found. mass, structure not grape-like, extending over central Female reproductive tract (figures 13, 46) adjacent to 80% of digestive gland, covering half to all ot right side rectum along right side of mantle cavity. Female open- of coil (figure 46). ing, about level with anus, situated on muscular papilla (During copulation, sperm are apparently deposited hanging freel\ in mantle ca\it\ at distal end of capsule in the \agina, temporariK stored in the bursa copulatrix gland. From there, closed sperm duct leading posteriorly, and then stored in the two posterior receptacula. Fertil- initially forming very thick-walled muscular vagina (fig- ization occurs in the anterior part of the coiled oviduct, ure 52), with lumen almost filled with large gland cells. after which fertilized eggs pass through the ciliated lu- Sperm duct giving rise to small, ball-shaped, dorsally- mina of the albumen and capsule glands where they are situated sperm pouch w ith weakK muscular walls. Thin- surrounded by capsular and mucous material. Eggs ap- walled ducts leading from sperm pouch and muscular parently enter the mantle cavity through the vagina, as vagina joining shortly before opening into slit-like lumen no other female opening was found.) of massive capsule gland. \'agina-t>pe muscular wall gradualK disappearing, while thin-walled part forms Sex and size relationships (figure 53): Specimens less sperm channel ("sperm groo\e " in "ventral channel of than 7.6 mm shell diameter (n = 25) were all males. = authors), in communication with gland but partially sep- Specimens of shell diameter greater than 8.6 mm (n arated by lateral fold (figure 46, cross-section B, If). An- 6) were all fully functional females without male repro- terior part of capsule gland w ith large, turquoise-staining ductive structures. Specimens 7.8-8.5 mm shell diameter gland cells; remaining capsule gland staining dark blue (n = 6) were "transitional" in appearance (figure 45); in sections. [M least in the anterior part, the gland cells visceral and pallial reproductive organs were clearly fe- are arranged in a comple.\ pattern (see Ponder, in press). male, however, with visible remnant of pallial vas def- A detailed histological description was hampered by the erens and "wound" (= penis scar) at attachment site of extremely strong staining reaction of the glands.] Pos- penis. Pallial vas deferens of smallest "transitional" spec- teriorly, glandular mass continuing, with communicating imen present as unciliated duct, partialK opened at prox- lumina, as albumen gland, .\lbumen gland massive, thick- imal end, extending between penis scar and distal end walled, with narrow, slit-like lumen, pinkish-orange in of capsule gland. No positive connections observed. This living animals, staining turquoise in sections; folded as specimen also with nearly transparent gonad containing S-shaped loop on right side of animal, partK in parallel small amount of whitish ovarian tissue in early part of with posterior part of capsule gland. Posteriormost part coil, and with no sperm in the female system (it appar- of albumen gland pressing against, or, in large females, ently had not yet mated in its female phase). Largest extending beyond posterior mantle wall. Sperm channel "transitional" specimen with fully-developed ovary, faint separating at junction of capsule and albumen glands, longitudinal marking on integument in position of vas forming closed o\iduct. penetrating posterior mantle wall, deferens, and weak penis scar. Simultaneous possession and forming large, widened, non-glandular coil, con- of both ovarian and testicular tissues not observed. taining (from sections and dissected specimens) both "Transition" somewhat correlated with date of col- sperm and eggs. At its posterior end, coiled oviduct giving lection (figure 53). Collections made between March and rise to 2 proximal sperm pouches, as inconspicuous, sub- mid-August exclusively male. Fully developed females, equal, ball-shaped sacs, closeK' adjacent, and partialK' "transitional" individuals, and males encountered in each hidden under la\ers of connective tissue and kidnev. of four collections in later part of August and September.

Figures 38-43. Cyclostremiscus beauii. 38. Left lateral view of visceral coil, showing position of stomach. Dashed line indicates location of dorsal incision. 39. Stomach, dorsal view, opened at incision shown in figure .38. .\rrows inside stomach indicate flow^ of particles, 40. Gastric shield. 41. Right jaw, inner side, posterior end up. showing details of jaw elements. 42. Head with mid- dorsal incision show ing buccal mass, circumesophageal nerve ring, and base of penis. Connective tissue and minor nerves removed. 43. Central nervous s\stem, left side, oblique lateral view. Scale bars: 39, 42 = 0.5 mm; 41, 43 = 0.1 mm

bg. buccal ganglion; bm, buccal mass; cpc, cerebropedal connective; cs, crystalline style; ct, ctenidium; dg, digestive gland; dgo, opening of digestive gland; es, esophagus; eso. opening of esophagus; gon, gonad; gs, gastric shield; in. intestine; ino. opening of

intestine; ki. kidney; leg, left cerebral ganglion; Iplg. left pleural ganglion; Ipg. left pedal ganglion; mo, mouth; mpg. metapodial ganglion; on, optic nerve; os, osphradium; pch. posterior chamber; peb, base of penis; pipe, pleuropedal connective; plspec, pleurosupraesophageal connective; ppg, propodial ganglion; pvd, pallial vas deferens; ra, radula; ras, radular sac; rcg, right cerebral ganglion; rplg, right pleural ganglion; sag, salivary gland; sbeg, subesophageal ganglion; speg, supraesophageal ganglion; ss, style

sac; st, stomach; stc, statocyst; stl, statolith; tns, swelling of tentacular nerve; ty, typhlosole. Page 14 THE NAUTILUS, Vol. 102, No. 1

spc'^ spc

Figures 44-46. Cyclostremiscus beauii. reproductive system (semi-diagrammatic) 44. Male phase 45. "Transitional female organs phase. 46. Female phase, with three cross-sections (A, B, C) through female glands. Small arrows indicate reflected

ct, ctenidium; hg, ag, albumen gland; an, anus; be. bursa copulatrix; eg, capsule gland; co, coiled oviduct; cont. connective tissue; opening of prostate gland into mantle cavity; hypobranchial gland, If. lateral fold, ov, ovary; pe, penis; pr, prostate gland; prom, .

R. Bieler and P. M. Mikkelsen, 1988 Page 15

Mating and larval development unknown. Protoconch 1857a;173), did not give an indication of the number of morphology suggests planktonic veliger stage (see Dis- specimens in the original lot. Moore (1964T32) men- cussion). Sex change within single individual not ob- tioned a "holotype" in the Paris Museum. However, the served. single A. fceauii-specimen in that type collection (MNHN

unnumbered, vidi) is much smaller than the dimensions Habits and habitat: To our knowledge, this is the first given by Fischer (teleoconch diameter 6.3 mm, height record of habitat type for this species, i.e., within the 3.5 0.42 umbilical di- burrows of thestoniatopod L(/sio.sq(n7/a scabricaiida, oth- mm, protoconch diameter mm, ameter 1.1 mm; protoconch whorls 2, teleoconch whorls er published records (,see synonym) ) refer only to empty 2%; vs. 10.5 mm maximum shell diameter in the original shells. Wells et al. ( 1961 ) recorded Cyciostremiscus beatiii description). is therefore considered the from the stomachs of Astropecten articutatus (Say, 1825), The specimen only remaining syntype of a formerly larger lot, and is however it is unclear whether the material was alive here selected lectotype. when swallowed b> the starfish. Another specimen from as the clo.se gut contents of A. articutatus, in the UNC-IMS collec- Woodring (1928:440) pointed out resemblance between Guppy's (1866) Cyctostrerna bicarinata tions, from 24 m depth off North Carolina, contained and Adeorbis beauii Fischer, but separated the two be- dried animal tissue and is the only other verified live- collected specimen known to the authors. Although bur- cause the latter "has a more strongly sculptured base." in turn separated his species CAr- rows of other local species, e.g., callianassid shrimps, Smith (1937:67) new polychaetes, hemichordates, and sipunculans, were also culus stirophorus from both "Circuhis bicarinatus Gup- py" ' sampled, these vitrinellids were not collected in associ- and "Circuhis beauii Fischer, by differing numbers of spiral ribs. re-examination ation with an\- burrower but Lijsiosquilla during this upper and basal A of the stud\ neotype (Woodring, 1928) of Cyclostrema bicarinata and the original description and figure The snails probabK feed on algae, bacterial films and (USNM 115621) of Circuhis stirophorus revealed that fall detritus in the burrows. They are capable of handling Smith both within the range of variation (partly caused ontoge- larger items in their alimentary tract, as evidenced by by netic of basal sculpture) displayed the Ft. various shell pieces and foraminiferan shells found in the change by Pierce are stomach. From the absence of glandular esophageal population, and both here synonymized (see pouches and the presence of a crystalline style in the also Moore, 1964:131). The holotype of Circuhis stirophorus could not be located. Although the original author stomach, it can be inferred (Yonge, 1930) that free pro- indicated the in it teolytic enzymes, capable of digesting animal matter, type to be the MCZ collection, was never received by that institution (K. Boss, personal are not present in this species and would not be expected J. it also at in this group. communication); was not located the Florida Individuals were almost invariably collected in groups State Museum, Gainesville (F. G. Thompson, personal of more than one animal per burrow sample; the max- communication). Cyciostremiscus beauii is the largest vitrinellid imum number encountered in one burrow sample was in the seven. Cyciostremiscus heauii was twice found crawling western Atlantic. Two Recent Panamic species are very similar in size, sculpture: openly on sand or seagrass in the vicinity of Lysiosquilla shape and burrow openings. In captivity, the animals were active Cyciostremiscus major Olsson & Smith (1951:46, pi. figs. la,b) differs in crawlers and were not distracted by light; dark, sheltered 3, from Panama from Cy. beauii areas in the tank (provided by black plastic film con- having more numerous spiral ribs throughout and a more rounded periphery (holotype figured tainers) were not preferred. During resting periods, all ANSP 187199; by specimens attached themselves to the wall of their con- Olsson & Smith, 1951). Cyclostrema gordana Hertlein Strong pi. tainer, just above the water level, by means of highly & (1951:110, 9, figs. 3, 4, 7; holotype CAS 064803, vidi) from the \ iscous mucus produced by the anterior pedal gland. This behavior was not correlated with food availability or Gulf of California, differs conchologically from Cycio- stremiscus beauii in having a prominent double spiral water qualitv'. It might reflect tidal rhythm, but material rib on the base between the basal keel the umbilicus was not sufficient to test that hypothesis. and (see Pilsbry, 1953: pi. 55). Hertlein and Strong (1951: Geographical distribution: Western Atlantic, from 110) gave the following measurements for the "unique North Carolina to Brazil (Ceara and Alagoas; teste Rios, type": maximum diameter 9.7 mm, minimum diameter 1985:41). Recorded from the Pliocene of Jamaica and 7.0 mm, and height 3.3 mm; our examination of the Florida (see synonymy). holotype yielded, respectively, 8.6, 6.8, and 3.9 mm. Taxonomic remarks: Fischer (1857b;286), in the orig- Hertlein and Strong (1951:110) compared C. gordana to inal description of Adeorbis heauii (in his earlier pub- "Cyclostrema angulata A. Adams [1850] from the West

lication, the name occurred only as a nude list name; Indies ' [following "Pilsbry" (= error for Tryon), 1888:

ps, penis scar; pvd, pallia! vas deferens; pw, posterior pallial wall; re, rectum; rs, receptaculum seminis; sc, sperm channel; spc,

sperm channel; te. testis; va, vagina; vo, visceral oviduct; vs, part of visceral vas deferens serving as vesicula seminalis; vvd, visceral vas deferens. Page 16 THE NAUTILUS, Vol. 102, No. 1

Figures 47-52. Cycloslrcmiscus heauii, specinu-iis from Ft. Vwvcv Inlet, Florida (light micrographs or critical-point dried SEM preparations). 47. Cross-section through \isceral ma.ss (male), at level of ve.siciila seminalis. 48. Section through prostate, showing R. Bieler and P. M. Mikkelsen, 1988 Page 17

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D o o o o o n15

2 3 4 5 6 7 8 9 10 11 12 Imml

Figure 53. Cyclostremiscus beauii. Plot of maximum shell diameter vs. number of teleoconch whorls (TW). Inset: Histogram summarizing sex distribution of collections, Ft. Pierce Inlet population. A = collecting period March-July, B = August 1-15, C = August 16-31, D = September, 1987. Open circles = empty shells and unsexed specimens. Open squares = functional males. Cross- hatched open squares = "transitional" females with pallial vas deferens and/or penis scar. Solid squares = females without male structures. Star indicates lectot>pe specimen (MNHN unnumbered).

92, they considered C. angulata a senior synonym of Beaui, Fischer (fig. 63), a West Indian species" (Tryon,

Cijclostremiscus beauii —see below] and stated that it 1888:92-93). Tryon 's figures (1888: pi. 32, figs. 64, 65) differed "principally in the smaller size and more de- may represent Cyclostremiscus beauii. However, as sim- pressed form." However, a re-examination of the C. gor- ilar species are known from the Indo-Pacific, we feel that dana t%pe showed it to differ principaJK- b\' a strong, the two nominal species should not be synony mized with- second basal keel surrounding the umbilicus. It definitely out a re-examination of Cyclostrema angulata type ma- belongs in the genus Cyclostremiscus, as advocated by terial, which could not be located in the British Museum Keen (1971), (Natural History) (London) nor in the Redpath Museum Tryon (1888), followed by Hertlein and Strong (1951), (Montreal). considered Cyclostremiscus beauii a synonym of Cy- The type species of Cyclostremiscus, Vitrinella pan- clostrema angulata A. Adams, 1850: "C. angulata, A. amensis C. B. Adams, 1852, and several other, similar Ad. was described as from the Philippines on the au- species (see, e.g., Pilsbry & Olsson, 1945) have distinct thority of Cuming, but as that great collector sometimes axial sculpture which is lacking in Cy. beauii, Cy. major, made mistakes, the localit> needs confirmation. There Cy. gordana, and a number of other nominal species. can be no doubt of the identity with this species of C. Pilsbry (1953) described the subgenus Ponocyclus in Cy-

lumen and opening into mantle cavity. 49. Section through circumesophageal nerve ring, with saUvary glands. 50. Spermatozoon

(SEM). Arrow indicates junction of niidpiece and tail. 51. Same, detail of two heads (SEM). Arrow indicates junction between head and midpiece. 52. Cross-section through vagina (left) and rectum (right). Scale bars: 47-49, 52 = 0.1 mm; 50 = 4 ^m; 51 = 2 ^m.

cc, cerebral commissure; cont, connective tissue; ope, cerebropedal connective; dg, digestive gland; es, esophagus; hg, hypobranchial gland; leg, left cerebral ganglion; Ipg, left pedal ganglion; mc, mantle cavity; mw, mantle wall; pc, pedal commissure; pch, posterior chamber of stomach; pr, prostate gland; prom, opening of prostate gland into mantle cavity; rcg, right cerebral ganglion; re, rectum; rpg, right pedal ganglion; sag, salivary gland; te, testis; tm, transverse muscle band; va, vagina; vs, part of visceral vas deferens serving as vesicula seminalis; vvd, visceral vas deferens. Page 18 THE NAUTILUS, Vol. 102, No. 1 R. Bieler and P. M. Mikkelsen, 1988 Page 19

clostremiscus \\ itli Adeorbis beauii as the type species, lateral midline. Suture impressed. Ventral surface below and distinguished it from Cyclostreniiscu.s by its lack of keel less convex, often with 30-40 widely-spaced, low axial sculpture. As Pilsbr\ alread\ pointed out in his axial ribs which are primarily evident from inside of description of Ponocijclus (1953:426). there are a number body whorl (figure 57). L'mbilicus wide (~ 25% of width). of species showing intermediate conditions ("weak traces Outer lip ver>- slightly reflected; some specimens with of axial sculpture") and the name is not used here. one former varix. Aperture at oblique angle to dorso- Due to the small number of species studied in the ventral axis. Sutural sinus shallow. Periostracum thin, \ itrinellidae, little can he said about anatomical char- transparent, with spiral grooves more numerous than on acters on the generic level. Aside from the large and shell surface. elaborately-sculptured shell, the extensive ciliation of the head-foot, and the paddle-shaped pallial tentacle, Cy- Protoeonch (figure 58): Smooth, 0.5 mm diameter, clostremiscus beauii differs anatomically from Circulus about 2 whorls. No sculptural demarcation separating striatus (see Fretter, 1956), Ci. texaniis (see below), and protoeonch I and protoeonch II. Cochliolepis parasitica (see Moore, 1972) in ha\ing the pallial portion of the intestine straight and uncoiled. External anatomy and organs of the mantle cavity (figures 64-66): Living animal translucent white with buccal mass (visible through proboscis) and protrusible Circulus Jeffreys, 1865 (page 315) portion of ctenidium rose-pink; digestive gland orange; Type species by monotypy: Delphinula duminiji Re- ner\e ring area opaque yellow, visible through integu- quien. 1848 (page 64; not "Trochus duminyi Requien" ment between eyes. Head with long, extensible snout, as usually cited) [= Yalvata? striata Philippi. 1836; see rounded and terminally notched at mouth. Ventral side Jeffre\s, 1865:317, where he used the junior synon> m D. of snout ciliated behind mouth opening, with cilia beat- duminyi only to a\oid secondary homonymy with Tro- ing toward mouth. Two long cephalic tentacles w ith im- chus striatus Linne, 1767]. mobile bristles on slight]} enlarged tip; motile cilia at least at tip and along side facing snout. Left cephalic tentacle fitting into shallow notch formed by shell keel Circulus texanus (Moore, 1965) new combination (figure 65). Eyes black, on slight bulges at base of cephalic (figures 54-68; tables 1, 2) tentacles. Neck region very long, slender. Foot elongate,

Vitrinella texana Moore, 1964:66 [unpublished dissertation). flattened, with anterolaterally recurved corners; fineK

Vitrinella texana Moore, 1965:76, pi. 7. figs. 4-6 [Gulf of ciliated sole broadly rounded posteriorly, extending well Mexico]. beyond shell (figure 65). Transverse crease on sole about 1964" ? Vitrinella texana "Moore, —.Andrews, 1971:73-74, text- '/^ of total length from anterior edge. Anterior pedal "1965"], fig. —Andrews, 1977:887 [citing text-fig. [poor fig- mucous gland (figure 65, amg) opening at slit across = ure; Vitrinella sp., teste Ode, 1987.35]. entire leading edge; mid- ventral fold evident on posterior Vitrinella (Vitrinella) feiana,— Abbott, 1974:83. quarter of sole, but presence of posterior mucous gland Vitridomus texana. —Ode, 1987:37. not confirmed. Locomotion by ci]iar\ action. Operculum Material examined: Ho]ot\pe: empt\ shell, 1.8 mm, (figures 61-63) corneous, circular, multispiral {ca. 8 USNM 636311 (Texas). Other material: 15 specimens. whorls), with small central peg and rounded-triangular FLORIDA: Ft. Pierce Inlet: 2-3 May 1987, 1 unsexed; muscle scar on inner surface. Operculum supported by 24 June 1987, 14 specimens (4 males, 7 females, 2 un- opercular lobes (figure 65, ol) on dorsal side of foot. Lobes sexed, 1 empty shell). simple, unpigmented, without tentacles. Epipodial tentacles absent. Two finger-shaped pallial tentacles (figure 64, Ipt, upt) Description arising separately but adjacently from inner mantle edge, Teleoconch (figures 54-56): Shell small (1.7-1.8 mm at right side of animal. Upper (= most dorsal) tentacle diameter, 0.55-0.65 mm height), with 1V2-1% teleoconch unciliated, with immobile terminal bristles, curling into w horls; almost planispiral, sculptured dorsalK' and ven- shallow sutural sinus, directed dorsalK during crawling. trally with about 18 fine spiral ribs; transparent when Lower tentacle bearing motile cilia, directed anterolat- alive, opaque after death. Ribs slightly stronger, more erally. Ctenidium (figures 64, 65, ct) with numerous fin- widely spaced just below suture on dorsum and at pe- ger-shaped filaments, attached for most of its length to riphery, where about 3 ribs form rounded keel below internal surface of mantle; anteriormost filaments darker

Figures 54-63. Circulus texanus, specimens from Ft. Pierce Inlet, Florida (SEM) (figures 54-56, USNM 846324). 54. Shell, apical view (2.7 mm diameter). 55. Shell, umbilical view (2.7 mm diameter). 56. Shell, apertural view (2.2 mm diameter). 57. Inside surface of base of bod> whorl, showing shallow radial grooves. 58. Protoeonch. 59. Larval shell, left lateral view (167 /im max. diameter! 60. Radula. 61. Operculum, inner surface (0.7 mm diameter) 62. Operculum, outer surface (0.8 mm diameter). 63. Operculum, oblique view of inner surface with opercular peg. Scale bars: 57, 58 = 0.2 mm: 60 = 5 ^m; 63 = 50 ^m.

rac, rachidian teeth. Page 20 THE NAUTILUS, Vol. 102, No. 1

Figures 64-68. Circulus texanus. 64. Crawling female, dorsal view, drawn as with transparent shell. 65. Crawling animal, ventral view, drawn as with transparent shell. 66. Head of male, left lateral view, showing penis. 67. Egg mass 68. Two egg capsules, each with veliger larva shortly before hatching. Scale bars: 64, 65 = 0.5 mm; 66, 67 = 0.25 mm. ag, albumen gland; amg, anterior mucous gland; br, immobile bristles; eg, capsule gland; ct, ctenidium; dg. digestive gland, ebv, efferent branchial vessel; ey, eye; he, heart; in, intestine; ki, kidney; Ipt, lower pallial tentacle; ol, opercular lobe; op, operculum; ov, ovary; pe, penis; re. rectum; sh, shell; sn, snout; ss, style sac; st. stomach; ten, cephalic tentacle; umb, umbilicus; upt, upper pallial tentacle. in color, not attaclu'd to tnantle skirt, extending out ot main cusp and subequal flanking cusps (6-7 inner, 9 aperture at right side of head. E.xtent of osphradium not outer). Apex of inner marginal tooth with numerous long, ascertained. thin, flanking cusps on either side of similarly-sized main cusp. Apex of outer marginal tooth with long, thin inner Alimentary system: Riiduia (figure 60) taenioglossate. flanking cusps, slightly more robust than tho.se of inner Rachidian tooth wider than long, with acute posterior marginal tooth; cusps at termiiuis subequal in size; outer corners projecting laterally and concave front. Main cusp margin with 2-3 cusps decreasing in size, remainder of narrow, unserrated, with 5-6 slightly smaller flanking outer margin smoothly rounded. Jaws composed of ele- cusps on each side (number of flanking cusps var\ ing ments approximately 4 ^m in length. within single radular rihhon) decreasing laterally in size; Stomach traii.spinciit, with rotating style often clearly base with 1 weak elevation per side (in position of basal visible through shell. Posterior part of stomach not mark- denticle), midway between posterior corners and central edly elongated. Intestine (figures 64, 65, in) forming wide ridge. Lateral tooth with asymmetrical cutting edge, loop in posterior part of mantle cavity. Anus just above

strongly indented at front edge, with narrow , unserrated anterior end ol capsule gland in females. Fecal material R. Bieler and P. M. Mikkelsen, 1988 Page 21

as continuous rods of irregular length, rather than pellets, ming or resting on bottom of bowl, with strong ciliary with no obvious surface sculpture. action on extended vela. No further development noted. Added cultures of mixed single-celled green algae ap- Reproductive system: Penis ifigure 66, pe) a posteriorly parently not excepted; all larvae died within next 2 days. directed, counterclockw ise, double coil; base arising pos- [From the size difterence between these larvae (170 nm) terior to eyes, somew hat right of dorsal midline. Testis and the protoconchs of adult shells (500 nm), a longer orange. lar\al stage, w ith extensive food intake can be predicted.] Ovary (figure 64, ov) filling inner side of early whorls habitat: Habitat as described for Cyclo- with grape-like lobes, lighter in color than digestive gland. Habits and stremiscu.s hcaiiii. In the laboratory, animals of Circiilns Capsule gland and much darker-staining (in metln lene very active fed on normally-occurring blue) albumen gland (figure 64, eg, ag) forming compact texanus were and bacterial surface films. mass on right wall of mantle cavity, below rectum, ter- algal and minating anteriorly at level of anus. Position and number Geographical distribution: Circuhis texanus (type lo- of sperm pouches not ascertained. cality: Mustang Island, near Port Aransas) was previously No evidence of se.xual dimorphism associated with reported to be endemic to the northv\estern Gulf of Mex- possible sequential hermaphroditism. Males both consid- ico, and has never been reported alive (Moore, 1964:41; erably smaller (1.7 mm diameter at 1% whorls) or larger .\ndrews, 1977:87; Ode, 1987:37). No fossil records are (1.8 mm diameter at 1': whorls) than females in same know n. sample (1. 74-1. 78 mm at I'/s whorls). One copulation briefly observed, wherein male paused while crawling Taxonomic remarks: Moore (1965:77) placed this species over dorsal surface of female, with heads adjacent (male in the genus Vitrinclla C. B. Adams, 1850, and pointed at left) and at angle of about 30°. Position of penis not out the differences in shell shape and sculpture in com- ascertained. parison to other species. However, members of Vithnella (based on the type species V. helicoidea C. B. Adams, Eggs and larval development: Three weeks after col- 1850; see tables 1, 2) are generally smooth-shelled or lecting, 7 females 4 males were placed into glass and weakly sculptured and are usually furnished with a ca- bowl with seawater (24 °C). Overnight, 15 egg masses rina bordering the umbilicus (e.g.. Keen. 1971:377; Ab- were laid; each mass contained 1-5 eggs, each in clear, bott, 1974:82). We feel that this species is better placed flattened, circular capsule (egg diameter 1 10 fim: capsule in Circuhis sensu lato because it agrees in shell shape diameter 170 ^m; n = 20). All eggs found in same de- and sculpture with the type species, Circuhis striatus velopmental stage, uncleaved, completely round, finely from the eastern Atlantic. Ode's (1987:37) placement of granular. Soft capsule walls of eggs laid in groups touch- this species in VHridomus Pilsbry & Olsson, 1945, cannot ing each other; each "mass," even if consisting of single be accepted: members of Vitridomus [a "rather feebly egg, b\ sticky, clear, gelatinous layer (figure 67). coated defined genus (Pilsbry & Olsson, 1952:36), very similar Eighteen additional egg masses were found in glass jar to Teinostoma H. & A. Adams, 1853] ha\ e the umbilicus in which specimens had been kept since collecting. Total partly covered by a callus. number of masses (with respective numbers of single-

egg-capsules) was: 4 masses with 1 capsule, 4 masses with 2, 13 masses with 3, 9 masses with 4, and 3 masses with DISCUSSION 5 capsules. One mass with 5 eggs followed through development: Comparison of the anatomical information on Cyclo- From beginning of equal, holoblastic 2-celled stage, 60 stremiscus beauii and Circuhis texanus with literature

minutes elapsed to 4-celled stage, 205 minutes to 8-celled data revealed that published information is scarce or stage, 385 minutes to 16-celled stage, respectively. Gas- misleading, and that genus- as well as family-level taxa

trula-stage discernible after 1 1 hours; embryos rotating of marine near-planispiral Rissoacea are poorly defined. after 17 hours. Three nominal families, Vitrinellidae Bush, 1897, Tor- X'eliger larvae (figure 68) fully developed at 104 hours, nidae Sacco, 1896, and Circulidae (for authorship, see intermittently or constantly rotating, with transparent below), are currently in use for members of this group, shell, 2 short veliger lobes, black eye spots (not situated and the lack of comparative data led Ponder (in press) on tentacles at this point), statocysts and operculum [much to treat all three families as a single group in a phylo- as described for Caecum glabrum by Gotze (1938:108, genetic analysis of the Rissoacea. In the follow ing dis- text-fig. 33), but with smaller vela]. Periphery of velar cussion, we (a) review the available data on species of lobes bearing long cilia, underlain by row of shorter cilia. this complex, (b) compare Cy. beauii and Ci. texanus Capsule size now approximately 150 x 120 fim, greatest w ith these data and with other members of the Rissoacea,

shell diameter 107 /xm, longest dimension of veliger with (c) suggest a preliminary grouping for the species for extended \ela 133 ^m. During entire process, 3 of 5 w hich anatomical data are available, (d) demonstrate the embryos developed completely synchronously; other 2 differences between Vitrinellidae and Tornidae, (e) eval- initially about 45 minutes, later several hours, behind. uate the taxonomic status of the nominal family Circu-

First veligers hatched after 130 hours and swam actively. lidae, and (f ) discuss some unusual aspects of the anatomy For follow ing 36 hours, veligers observed either swim- and reproductive biology of Cy. beauii. namely ciliation. Page 22 THE NAUTILUS, Vol. 102, No. 1

tentacle shape, stomach morphology, and hermaphro- type " peg in other families of Rissoacea (character con- ditism. sidered plesiomorphic in this superfamily), while the cir-

cular shape of the vitrinellid operculum is considered (a) Available literature data (see tables I, 2). derived (Ponder, 1985:5). Except for Fretter's (1956) study of Circulus striatus, The osphradium of Cyclostremiscus beauii is ver) sim- and Woodward's (189S) and Graham's (1982) work on ilar to those described for Circulus striatus (see Fretter, Tornus suhcarinatus, most of the available data on the 1956:372) and for species of the Rissoidae (Johansson, anatom\ of marine near-planispiral Rissoacea are re- 1939:319, pi. 3, figs. 3, 4, text-fig. 5; Haszprunar. 1985: stricted to descriptions or illustrations of external mor- 476, figs. 7k, 16). phology, especially Pilsbr\ and McGinty's (1945a, 1946b) The radulae of the two species studied here ha\ e only sketches of western .\tlantic forms. Some of the few one pair of basal denticles (or elevations) on the rachidian published observations on the radular structure and gross tooth, a feature known from some other rissoacean fam- morphology of vitrinellid softbodies are misleading (see ilies, such as Hydrobiidae (see Bandel, 1984:29, text-fig. also Moore, 1964, 1972). Based on the misconception that 47) and Rissoidae (see Ponder, 1985:10). This character these species are archaeogastropods. Rush (1897:127, 142, was hvpothesized to be primitive in the Rissoacea by pi. 22, figs. 12a-g) erroneously constructed a rhipido- Ponder (1985:119). glossate radula for Circulus trilix (Bush, 1885) [= Cy- The position of the salivary glands relative to the cir- clostremiscus pentagonus (Gahh, \8S7), fide Moore, 1964; cumcsophageal nerve ring is often used in family- and 138]. The "supplementar\ plicated gill" (Stimpson, 1858: superfamiK -level discussions (e.g., Ponder, 1983:236, 258; 308) of Cochliolepis parasitica uas not found in that in press). However, as a highly variable character in the species by Moore (1972:101). Pilsbry (1953:427) men- populations of Cyclostremiscus beauii studied herein, tioned a taenioglossate radula for Cyclostremiscus beauii, further data on additional species are necessarv before and added "This will be figured in a paper now in prep- it can be reliabl) utilized to infer phylogenetic relation- aration b\ T.L. McGinty and the author " This work was ships of the Vitrinellidae. apparentl) never published. The extensive anterior section of the esophagus of Cy- (b) Comparison of Cyclostremiscus beauii and Circulus clostremiscus beauii, with its long, coiled, dorsal folds, texanus with other \itrinellids and rissoaceans. is verv similar to that of Hydrococcus brazieri (see Pon- Both Cyclostremiscus beauii and Circulus texanus dis- der, 1982:77). play "typical" rissoacean characters (as outlined by Fret- The conspicuous ciliated tract leading from the renal ter & Graham, 1978:153; Boss, 1982:984; Ponder, 1983, opening to the head of Circulus striatus (see Fretter, in press) and agree to a large extent with the previous 1956:372) is lacking in Cyclostremiscus beauii. concept of Vitrinellidae (Fretter, 1956; Moore, 1972; see The presence of markedly vacuoled connective tissue

tables 1, 2, Group 1). As in most other vitrinellid species, in Cyclostremiscus beauii is a character shared by Cir- the cephalic and pallial tentacles bear immobile bristles culus striatus and Tornus subcarinatus (see Graham, and/or motile cilia. Tracts of motile cilia on the cephalic 1982:147). tentacles occur in numerous groups in the Rissoacea; for The nervous system of vitrinellid species is tvpically

lack of comparative data, it is not yet clear whether the rissoacean and shows moderate (Cyclostremiscus beauii) arrangement of these cilia in grooves, as in Cy. beauii, to high (Circulus striatus; Fretter, 1956:377) concentra-

is unusual. Photographs of critical-point dried tentacles tion. The RPG ratio of Cy. beauii, averaging 0.49. lies of a freshwater rissoacean, the hydrobiid Tryonia ctath- within the range of those of members of the rissoacean rata Stimpson, 1865 (see Hershler & Thompson, 1987: family Pomatiopsidae (see, e.g., Davis ii Mazurkiewicz,

27, figs. 13-17), show no grooves present in that species. 1985:45, table 8). Cyclostremiscus beauii and Circulus texanus have two The structure of the male and female reproductive closely-spaced pallia! tentacles on the right side. This systems encountered during this stud\ is ris,soacean (e.g.,

arrangement is present in most studied vitrinellids (the Johansson, 1956). As in most other rissoaceans (e.g.. Pon- exception being Cochliolepis albiceratus Ponder, 1966), der, 1985:6), the albumen and capsule glands form a in the type species of Tornus (Tornidae, see below), in single, continuous mass [a character not recognizable from Hydrococcus brazieri (T. Woods, 1876) (monotypic Hy- Fretter's (1956) description and figures of Circulus stria- drococcidae, see Ponder, 1982), and in the rissoid genus tus]. The N'itrinellidae differ from the Rissoidae (e.g.. Rissoina Orbigny, 1840, where some species have either Ponder, 1985: text-fig. 2) wherein a non-homologous the anterior (right) or posterior (left) pallial tentacle bi- structure, an expanded part of the upper oviduct, pos- lobed (Ponder, 1985:78). As described for Cy. beauii, the terior to the bursa copulatrix and/or receptacula seminis, function of the upper tentacle, which usually bears im- functions as an albumen gland. This "upper oviduct

" mobile bristles, and is held upright and bent around the gland of the Rissoidae is homologous with the coiled

aperture, is clearly sensory, while the lower one, often part of the visceral oviducts of the Hvdrobiidae (see distinctly ciliated and sometimes broadened, controls and/ Johansson, 1956) and N'itrinellidae [Fretter's (1956) "ren- or enhances water flow. al oviduct ']. Both species have a small central opercular peg which Most members of the Rissoacea have one bursa cop-

is probably not homologous with the lateral "neritid- ulatrix and one receptaculum seminis near the posterior R. Bieler and P. M. Mikkelsen, 1988 Page 23

to pallial \\ all, ifquii iiig the sperm to travel the entire length "most, if not all, species appear have a planktonic of the sperm ehannel (vaginal lumen, sperm groove of veliger stage of some duration." the ventral channel of authors) immediately after cop- The sperm cells of Cyclostremiscus beauii are of the ulation. Others have develeoped additional, ilistal sperm general type known tor other members of Rissoacea (see pouches (anterior sperm-storage structure, distal l)lind Gotze, 1938; Franzen, 1955). The twisted acrosome and sac, sac-hke vestibule, spermatheca, pseudo-bursa of au- relative lengths of head, midpiece and tail are similar to thors), eitlier in addition to [e.g., Ptisillina incon.^pictia those described by Franzen (1955) for Caecum glabrum (Alder, 1844); see Johansson, 1939:337. text-fig. 22 (as (Montagu, 1803). The head is much shorter than that of Rissoa); Rissoidae], or instead of the pro.ximal bursa cop- the two rissoids previously studied [Pusillina incon.spicua ulatrix [e.g., Htjala vitrea (Montagu, 1803); see Johans- (Alder, 1844) (as Rissoa) and Onuba striata (J. Adams, son, 1949: text-fig. 1; Iravadiidae]. These distal sperm 1797); Franzen, 1955:406-409], and the relative length pouches are not necessarily homologous with each other of the midpiece much greater than in Hydrolna ulvae and certainly not with the proximal bursa (see, e.g., Sla- (Pennant, 1777). voshevskaya, 1978). Vitrinellids for which such data are Direct communication of the coiled ("renal") oviduct available (CArciilus striatus and Cyclostremiscus beauii) with the kidney (Fretter, 1956), previously considered differ from most other rissoaceans in the presence of two unique for Circulus or the Vitrinellidae, is now also known more-or-less eciually developed, proximal receptacula for other families of the Rissoacea (Tornidae, Graham, seminis which, in position and size, look much like the 1982; Truncatellidae, Fretter & Graham, 1962). two sperm sacs {i.e., bursa copulatrix and receptaculum Anatomical characters of Circulus striatus, which seminis) of other Rissoacea {e.g., in the rissoid Lucidestea Fretter (1956:380) discussed as probably "associated with Laseron, 1956; see Ponder, 1985:67, text-fig. 3). Whether small size and body form," are equally expressed in the the presence of two proximal receptacula seminis is a much larger Cyclostremiscus beauii, suggesting that distinguishing character for Vitrinellidae cannot yet be Fretter's hypothesis was incorrect. determined, as accessory receptacula have been demonstrated for members of the genus Alvania in the Ris- (c) Preliminary grouping of "vitrinellid-like" species for soidae (Johansson, 1956; Ponder, 1985). In both Circiilus which anatomical data are available. striatus (see Fretter, 1956:377) and Cyclostremiscus The species for which sufficient gross morphological beauii, the distal end of the coiled oviduct serves as the data are available appear to fall into three groups, two fertilization area, and not as an additional functional of which are here considered of familial rank and one receptaculum as has been reported for some other ris- comprising species of incertae sedis (tables 1, 2). Mem- soaceans [e.g., Alvania suhsoluta (Aradas, 1847), where bers of Group 1, currently placed in the genera Vitri- the distal end contained oriented sperm; Johansson, 1956: nella, Teinostoma, Pieurornalaxis, Cochliolepis, Circu- 380]. lus and Cyclostremiscus, are here considered to belong The spawn mass of members of the Rissoidae usually to the Vitrinellidae (the placement of the species de- has numerous eggs per capsule in planktonic forms, while scribed as Cochliolepis albicerata Ponder, 1966, is some-

' one-egg-per-capsule is characteristic of "direct devel- what doubtful as it has only one pallial tentacle, and the opment (see Lebour, 1937; Thorson, 1946). This does not morphologies of its osphradium and penis are unknown). hold true for the spawn of Circiilus texanus, which gen- Some of the characters attributed to the family Vitri- erally resembles that of the freshwater rissoacean Bithijn- nellidae in Moore's dissertation (1964) were based on ia tentaculata (Linne, 1758), as described and illustrated Parviturboides interruptus (C. B. Adams, 1850) (tables by Ankel (1936:164, text-fig. 142B). [The number of eggs 1, 2, Group 2), later considered a species of uncertain there, however, is larger (4-24; Lilly, 1953:104), and systematic position (Moore, 1972:107). This species dif- hatching occurs at the crawling stage]. Jablonski and Lutz fers from the other forms here grouped in Vitrinellidae (1980:336; after Taylor, 1975) stated that the Rissoacea by the following anatomical characters: posterior foot ". with immobile cilia, left cephalic tentacle with . . follow one of two developmental pathways: those margin that hatch as crawling juveniles from relatively large eggs four low swellings on proximal posterior border, penis (140-320 nm) and those that hatch from relatively small with glandular area and directed straight back. Another eggs (60-130 jxm) and spend 2-3 weeks as planktonic species, Tomura bicaudata (Pilsbry & McGinty, 1946) veligers. Both species studied here definitely fall into differs in many head-foot characters (tables 1, 2, Group the latter category: veliger shells of Circulus texanus 2) and is here also considered as incertae sedis. White (figure 59) were one full whorl smaller than the final (1942:92) advocated the inclusion of "Cyclostrema" bushi larval shells as seen in the adult protoconch (figure 58); Dautzenberg & Fischer, 1907, in the Vitrinellidae/Tor- the protoconch of Cyclostremiscus beauii (figures 9-11) nidae complex and published anatomical descriptions shows a distinct line and change of sculpture between and illustrations of that species. While some morpho- protoconch I and protoconch II, with almost another full logical characters [head with well-developed eyes and whorl of growth before metamorphosis, suggesting sev- terminally ciliated cephalic tentacles, the well-developed eral weeks (K. Bandel, personal communication) of osphradium (1942: text-fig. 6), and the circular, multi- planktonic life. In his unpublished revision of western spiral operculum (1942: pi. 2, fig. 3)] agree well with the Atlantic Vitrinellidae, Moore (1964:18) inferred that species listed here as vitrinellids, the glandular pouches Page 24 THE NAUTILUS, Vol. 102, No. 1

in the esophagus and, most of all, the pair of epipodial that complex (metapodial tentacle present, esophageal (not pallial) tentacles on the right side of the animal pouches and glands present, penial glands present, os-

(White, 1942:90, text-fig. 5), prevent inclusion in \itri- phradium short, posterior end ot foot not simple) do not nellidae. Tornm siibcarinatus (tables 1, 2, Group 3) is occur in N'itrinellidae as understood here, and must refer here considered a member of a separate famiK (see be- to Tornus and other tornid genera studied b> Ponder low). (Pseudoliotia Tate, 1898, Scrupus Finlay, 1927; unpublished).

(d) Family relationships: Vitrinellidae-Tornidae. Vitrinellidae and Tornidae fit well into the Rissoacea, Several attempts have been made to define the key and an independent grouping of these famiilies as Tor- characters of the family Vitrinellidae [e.g., Moore, 1965: nacea (e.g., Kuroda et al., 1971; Golikov & Starobogatov, 74, 1969:170, 1972:107ff.; Boss, 1982:991). However, be- 1975) is not justified. The family Adeorbidae Montero- cause of the small number of species fulK studied ana- sato, 1884, used b\- some authors for members of this tomicall), no single synapomorphy defining the famiK complex, is a s\non> m of Tornidae, since Adeorbis S. is clear at this time. The most obvious features of the Wood, 1842, is an objective synonym of Tornus Turton Vitrinellidae are: a low-spired, translucent white shell; & Kingston, 1830 (see Iredale, 1914:172, 1915:344). long cephalic tentacles equipped with terminal bristles (e) The nominal family Circulidae. and, in most cases, motile cilia; two closely-spaced pallial The taxonomic status of the nominal family Circulidae tentacles on the right side; a large monopectinate gill remains problematic. The family name "Circulidae" was often projecting to the right of the head; a large linear first used b> Fretter and Graham (1962:642. ".\ppendix osphradium; a foot with simple or only slightly indented I"), in a list of taxa treated in that publication. The text anterior and posterior margins; a horn\ concentric, mul- ' reads merely "Circulidae: Circulus striatus (Philippi). tispiral operculum; and the possession of a non-glandular Fretter and Graham (1962) did not state that they in- curved penis in the male. Some of these features (shell tended to create a new family, did not mention the name shape, pallial tentacles, and projecting gill) have led au- in the main bod\ of the text, even u hen the taxonomic thors (e.g., Ta\ior & Sohl, 1962) to s\nonymize Vitri- position of the genus (1962:550, 618) and the composition nellidae and Tornidae, based on Woodward s (1898) work of the superfamiK (1962: 622-623) were discussed, and on Adeorbis [= Tormts] .siibcarinatus (Montagu, 1903). in fact ne\er used any famiK name but N'itrinellidae Others (e.g.. .\dam & Knudsen, 1969; Moore, 1972) have when the\ referred to Ci. striatus in subsequent publi- pointed out differences between vitrinellids and Tornus/ cations (Fretter bi Graham, 1978:227; Graham, 1982: toniids in features of the e\ es (functional eyes are lacking 147). Lacking an\ description, definition, or bibliograph- in T. .'iubcarinatiis ) and, or the operculum (oval and pau- ic reference, a name thus introduced is not taxonomically cispiral in T. siibcarinatus) and have separated the two available (ICZN, 1985: Art. 13). "Circulidae" could families. In a redescription of the anatomy of T. sub- therefore be regarded as a nomen nudum. carinatus, Graham (1982:147) saw the opercular shape However, Golikov and Starobogatov (1975:211) ac- as the "single difference between the two nominal fam- cepted "Circulidae Fretter & Graham, 1962" as a valid ilies and again advocated synonymy. family and stated (1975:218) that "the characteristics of Additional anatomical characters of Tornidae that the latter famiK are found in Fretter (1956:381)," there- warrant separation from Vitrinellidae are: (1) the con- by referring to the summar\ of Fretter s description of spicuous, elongate osphradium of the latter is not present Ci. striatus. This fulfills the requirement of ICZN (1985) in Tornus (Graham, 1982:144, found only "a small cil- Art. 13(a)ii (Bibliographic Reference to Published State- iated groove, which may be a reduced osphradium"), (2) ment) and, unless there are earlier such statements that the attachment of the ctenidial axis to the mantle is short have escaped us, this makes Goliko\ and Starobogatov and the axis hardly supplied with blood vessels, and (3) (1975) the authors of Circulidae, with Circulus as the the penis of 7". siibcarinatus bears several finger-like name-bearing t\ pe. The nominal family Circulidae is, processes which are not known in vitrinellids as delimited with anatomical descriptions available for Ci. striatus here. [Fretter & Graham (1978:231) described the cte- and Ci. texanus, much better detined than the X'itrinel- nidium of T. siibcarinatus as "partly bipectinate." This lidae, where our knowledge ot Vitrinella is based onK is apparentK in error, as both Woodward (1898) and on shell characters and Pilsbry and McGinty's (1945a, Graham (1982) described and illustrated that gill with 1946b) sketches of crawling animals (tables 1. 2). .\11 only a single row of lamellae.] As T. siibcarinatus lives available data suggest s>non\in\ of N'itrinellidae and in a comparable habitat (under boulders on well-oxy- Circulidae (tables 1, 2; Boss, 1982:991 ). In any case, the genated sand or iiuid) and is of about the same size as placement of N'itrinellidae and Circulidae in separate Circulits spp.. the drastic difference in osphradial type superfamilies, or even separate superorders (Golikov & indicates phylogenetic difference rather than specializa- Starobogatov, 1975), is highly exaggerated. tion on a low taxonomic level. Ponder (in press) combined the "tornicl-\ itrinellid-cir- (f) Unusual features of Cyclostremiscus beauii. culid complex" as lamiK 'i'ornidae tor the purpose of The stud) of ('yclustrentisciis beauii revealed a num- his pin logenetic analysis of the Rissoacea. However, most ber of features that differ from other members of the of the characters and character states he attributed to N'itrinellidae (and, in part, from the Rissoacea). Most of R. Bieler and P. M. Mikkelsen, 1988 Page 25

these features may be related to either its unusually large and lemales. However, a later in-depth study (Thiriot-

size (for a \ itrinellid) or its unusual habitat in stoniatopod Quievreux, 1982:]67ff.) of these species through annual burrows. cycles shov\ed a seasonal (and geographical) pattern of penis-size distribution in females, but no evidence of sex Ciliation and tentacle shape: The extensive ciliation of change. The survey did "not support the hypothesis of the gills, cephalic tentacles and lower pallial tentacle provides eflective respirator) and excretory currents. On a successive hermaphroditism" (1982:167). In Cyclo- stremiscus heauii, remnants of the male are the right side of the head, where in- and outgoing cur- apparatus only evident in the smallest functional ieniales (figure rents are present, water flow is controlled by the paddle- with a gradual decrease of male structures with size. shaped tentacle. Low tides and high water temperatures 53), As outlined above, we take this as evidence for a sex are likely tc create anoxic conditions in the burrows, and change in this species rather than a simple sexual di- a large vitrinellid such as Cyclontremiscus heauii may morphism in shell size. During ontogeny, the male re- be more strongly affected by oxygen deficiencies than, productive system apparently disappears completely and for instance, the smaller Circuhis texanus. and thus would is replaced organs. anteriormost part of benefit from an elaborate s\stem to produce and direct by female The the albumen gland is found in the currents. The onl\ other known commensal \itrinellid, pinkish-orange female same relative position as the prostate (of similar color Cochliolepis parasitica, is much smaller, and lives di- and reaction to staining) in the male phase. As no animal rectly on its host [under the scales of the giant scale worm Pohjodontcs lupina (Stimpson, 1856); see Stimpson, 1858; was found with a very early stage of development of the it whether these or- llartman, 1945:10]. Moore (1972:104) did not find cilia albumen gland, cannot be decided gans, or parts of them, are homologous. [Reid (1986), in on the gill filaments of Cochliolepis parasitica, suggesting a study of Mainwaringia Nevill, 1885, the first reported tliat currents produced by the annelid are sufficient to case of protandrous sequential hermaphroditism in the supply the snail. All other species studied live under rocks Littorinacea, found both a closed pallial oviduct and an (table 2). open prostate in the intersexual and female stages, "sug- Stomach: The rissoacean stomach does not normally gesting that these structures are not strictly homologous" have a caecum or an elongated posterior chamber, as (1986:225). In that littorinid group, small penes are re- occurs in various other prosobranchs ("it is not possible tained in the female phase (1986:237), and the pallial

to see any trace of it in the . . . Rissoacea"; Fretter & oviduct opens into the also-retained pallial vas deferens, Graham, 1962:225). OnK a few exceptions are known: then serving to carry egg capsules (1986:238).] With this Ponder (1985:78) described the stomach of Rissoina (Ris- admittedly small data set for Cy. beauii from only two soidae) as "very long due to elongation of posterior cham- localities, and the absence of data on individual devel- ber," and Ponder (in press) listed the character state opment and longevity, settlement cues and adult mo- "posterior gastric chamber not small" for the families bility, several scenarios could be constructed. It is, for Emblandidae, Truncatellidae and Stenothyridae. The example, theoretically possible that functional males ar- presence of a large posterior chamber in Cijclostreiniscus rest growth to prolong the male phase, or that the change beauii, similar in relative shape and organization to that into the female phase occurs only after successful mating of Pomatias elegans (Miiller, 1774) (Pomatiasidae), as as a male. However, no evidence was found to consider described by Graham (1939:90, fig. 6D), is therefore sur- this sex change as labile, i.e., environmentally mediated, prising. It may allow Cij. beauii to maintain a regular as is known for other caenogastropod families such as supply of food particles by regulating fluctuations in the Calyptraeidae and Stiliferidae (see Hoagland, 1978, for rate of food intake (as suggested by Graham, 1939:93, examples and discussion). The data are further insuffi- for Pomatias). especially since feeding must be strongly cient to demonstrate either seasonal or partner-induced affected b> tidal, and therefore water-level, changes change. The sex change in Cy. beauii appears to occur within the stomatopod burrows. A relatively large animal only once, at a predetermined size (figure 53; about 8 such as Cy. beauii would be more strongly affected than mm shell diameter, 3 teleoconch whorls), and it appar- smaller vitrinellids, e.g., Circulus texanus, which were entlv affects most, if not all, individuals (as the general observed to feed in very small pockets of water (personal shift in gender over time in the Ft. Pierce population observation), thus maintaining a more-or-less continuous suggests). feeding activity, as has been described, e.g., for a mem- Thus Cyclostremiscus beauii is interpreted as a pro- ber of the genus Caecum (Morton, 1975:14). tandrous sequential hermaphrodite. The term sequential

Hermaphroditism: The most striking result of this study (= consecutive, successive) is important here. In a review was the discovery of small functional females in Cy- of the terms protandry, protogyny and hermaphroditism, clostremiscus beauii with apparently functionless rem- Hoagland (1984:86) defined protandry as "the function- nants of the male reproductive system. Penial structures ing of an organism first as male, then as female, with no are known to occur in various female rissoaceans (par- further sex change. The two sexual phases are separated ticularly hydrobiids and rissoids). Thiriot-Quievreux by a phase in which male primary and secondary sex (1977:779ff.) based her hvpothesis of sequential her- characters disappear, and the animal re-differentiates as maphroditism in four species of Ris.soa on the presence a female." This however omits those species which, after of more-or-less developed penes in immature specimens an initial male-only phase, have both male and female Page 26 THE NAUTILUS, Vol. 102, No. 1

reproductive systems developed and tiiiictional (e.g., micrographs for figures 1-3. Dr Richard S. Houbrick members of Rissoellidac and Omal(>i;\ ridae; see below), provided valuable field assistance (he found the largest i.e., protandrous simultaneous hermaphrodites. [Simul- temale, without his glasses). Drs. Klaus Bandel (Univer- taneous hermaphrodites, in our understanding, do not sitiit Hamburg, West Germany), Donald R. Moore necessariiK use "the same gonad to produce both eggs (KSM.\S) and Winston F. Ponder (Australian Museum, and sperm" as defined b% Hoagland (1984:85).] S\dne\ ) offered comments on species identifications and Although, at this point, we do not know whether pro- larval shells Dr Ra\mond B. Manning (USNM) was first tandrous sequential hermaphroditism is the exception, to bring the molluscan associates of Lysiosquilla to our rather than the rule, in this family, the occurrence of attention and provided information about the stomato- hermaphroditism in Cyclostremiscus beauii could be in- pods. Dr. Robert Robertson (ANSP), Dr. Patrick Nuttall terpreted as an adaptation to its peculiar habitat char- and Ms. Kathie Way [British Museum (Natural History)], acterized b\ low densit\ and relative isolation. Protan- Dr. Terrence M. Gosliner (CAS), Dr. Fred Thompson drous sequential hermaphroditism has an ad\antage for (Florida State Museum, Gainesville), Dr. Kenneth J. Boss

Cy. beauii, as each individual thereby minimizes the age (MCZ), Dr. Philippe Bouchet (MNHN), Hugh J. Porter at which it first reproduces and increases the likelihood (UNC-IMS), Dr. M. G. Harasewych (USNM), and Fred- of finding a compatible mate in a small group. Inbreeding erick J. C'ollier (USNM) provided comments on and or between siblings would also be reduced (see Ghiselin, loans of specimens entrusted to their care. We also thank 1969, for discussion). the following: Julianne Piraino (SMSLP) for SEM assis- Members of the approximately 25 families grouped tance; Patricia Linley (HBOI) for advice on histological under Rissoacea are general!) described as gonochoristic techniques; Tom Smoyer for help with photographic (see, e.g.. Boss, 1982:984), although possibly derived from printing; Dr. Robert Hershler (USNM), Dr. M. G. Har- ancestors that were sequential hermaphrodites (Slavo- asewych (USNM), Richard E. Petit (North Myrtle Beach, shevskaya, 1984). Reported cases of hermaphroditism in SO, kristen Metzger (HBOI), and Carol Browder (HBOI) prosobranchs (e.g., Webber, 1977:10; Fretter, 1984:15) for help with literature acquisition. Drs. Kevin J. Eck- include onl\ two genera that have classicalK been as- elbarger (HBOI), M. G. Harasewych (USNM), Robert signed to this superfamily, Omalogyra (Omalogyridae) Hershler (USNM), and Richard S. Houbrick (USNM) are and Rissoella (Rissoellidac). Fretter (1948) described in thanked for valuable comments on the manuscript. The detail the anatomy and reproductive biology of O. ato- senior author gratefully acknowledges financial support muilPhilippi. 184'l)andfi. Jiap/!a(!a(Alder,'l848). Both provided by a NATO postdoctoral fellowship, adminis- species were found to be protandrous simultaneous her- tered b\ the German .Academic Exchange Service maphrodites, with some likelihood of self-fertilization in (D.\AD), and research facilities provided by the Smith- O. atomus (1948:612, 621, 630; see also Fretter & Gra- sonian Marine Station at Link Port, Ft. Pierce, FL. ham, 1962:381, and 1978:218, 223). [Fretter & Graham, This is Smithsonian Marine Station Contribution No. 1964:134, refer (erroneously:') to the .same two species as 217 and Harbor Branch Oceanographic Institution Con- being "protandrous con.secutive hermaphrodites. '] Both tribution No. 622. families have been subsequently removed from the Ris- soacea and have been recently placed outside the Cae- LITERATURE CITED nogastropoda, near (Salvini-Plawen & Haszprunar, 1987) or in (Ponder & Waren, in press) the Allogastropoda Abbott, R. T. 1954. .American seashells. D. Van Nostrand (= Heterostropha of authors). Thus, Cyclostremiscus Company, Inc., Princeton, NJ, xiv + 541 p., 40 pis seashells: Mollusca beauii is to our knowledge the only species in the su- Abbott, R. T. 1974. .American the marine coasts .America, 2nd perfamily Rissoacea for which protandrous sequential of the Atlantic and Pacific of North ed. Van Nostrand Reinhold Company. New York, 663 p., hermaphroditism has been demonstrated. However, as 24 pis. sequential hermaphroditism is not as easily recognized Abbott, R. T. and S P Dance. 1982 Compendium of sea- as simultaneous hermaphroditism, it might be more shells: a color guide to more than 4,200 of the world's widely distributed in the N'itrinellidae and/or other fam- marine shells. E. P Dutton, New York, x -I- 411 [+ 1] p., ilies of this group. The mismatch in descriptions of the illus. male reproductive system ol Tornus subcarinatus, where genres de Mol- Adam, W. and J. Knudsen. 1969 Quelques a penis was lacking according to Woodward (1898) and lusques prosobranches marins inconnus ou peu connus de was found by Graham (1982), might find an explanation r.Afrique occidentale. Bulletin Institut Royal des Sciences 1-69. after all. Naturelles de Belgique 44(27):

Adams, .A. 1850. Monographs of Cyc/o«(rf'ma, Marryat, and

Sejmratista, Gra\ . two genera of gasteropodous mollusks. ACKNOWLEDGEMENTS Proceedings of the Zoological Society of London 185041- 45. For invaluable help in the field and laboratory, we es- Adams, A. 1866. Monographs of the genera Cyclostrema, pecially thank William D ("Woody") Lee (SMSLP), Adeorbis, and Teinostoma. Thesaurus Conchyliorum. w hose enthusiasm, expert technique and keen eye added Sowerby, G. B. (ed.) 3:249-262, pis. 255-256. of this Dr. M. G. shells of the Texas coast. University of significantly to the progress study. Andrews, J. 1971. Sea Harasewych (USNM) furnished the .scanning electron Texas Press, .Austin, xvii -I- 298 p.. illus. R. Bieler and P. M. Mikkelsen, 1988 Page 27

Shells and shores of Texas. University of Marine Biological Association of the United Kingdom 27: Andrews, J. 1977. Texas Press, Austin, xx + 365 p., illus. 597-632. Ankel, W. E. 1936. Prosobranchia. In: Grimpe, G. and E. Fretter, V. 1956. The anatomy of the prosobranch Circulus of its position. Wagler (eds. ) Die Tierwelt der Nord- und Ostsee, Vol. striatus (Philippi) and a review systematic I\b. .-Kkademische Verlagsgeselischaft, Leipzig, p. 1-240. Proceedings of the Zoological Society of London 126:369- Bandel, K 1984. The radulae of Caribbean and other Me- 381. sogastropoda and Neogastropoda, Zoologische Verhande- Fretter, V. 1984, Prosobranchs. In: Wilbur et al. (eds.). The

lingen No. 214, 188 p., 22 pis. Mollusca, Vol. 7, Reproduction Academic Press, New York

In: Parker, S. P. (ed. in chief). and London, p. 1-45. Boss, K. J. 1982. Mollusca. Synopsis and classification of living organisms. Vol. 1. Fretter, V. and A. Graham. 1962. British prosobranch mol- McGraw-Hill Book Company, New York, p. 945-1166. luscs: their functional anatomy and ecology. Ray Society,

xvi -I- Bush, K. J. 1897. Revision of the marine gastropods referred London, 755 p. to Cyclostrema, Adeorbis. Vitrinella, and related genera; Fretter, V. and A. Graham. 1964. Reproduction. In: Wilbur, with descriptions of some new genera and species belong- K. M. and C. M. Yonge (eds). Physiology of Mollusca, ing to the .\tlantic fauna ot .America. Transactions of the Vol. 1. Academic Press, New York and London, p. 127- Connecticut .^cadeni) 10:97-144, pis. 22, 23. 164. Dall. W H 1889. \ preliminary catalogue of the shell-bear- Fretter, V and A. Graham. 1978. The prosobranch molluscs ing marine mollusks and brachiopods of the south-eastern of Britain and Denmark. Part 4— Marine Rissoacea. Jour- coast of the United States, with illustrations of many of nal of Molluscan Studies, Supplement 6:153-241. Plio- the species. Bulletin of the United States National Museum Gardner, J. 1948, Mollusca from the Miocene and lower

37:221 p., pis. 1-74. cene of Virginia and North Carolina. Part 2. Scaphopoda Dall, W. H. 1892. Contributions to the Tertiary fauna of and Gastropoda. U.S. Geological Survey Professional Paper Florida, with especial reference to the Miocene Silex-beds 199-B, i-iii, 179-310, pis. 24-38. of Tampa and the Pliocene beds of the Caloosahatchie Ghiselin, M. T. 1969. The evolution of hermaphroditism among River, 11. —Streptodont and other gastropods, concluded. animals. Quarterly Review of Biology 44:189-208. Transactions of the Wagner Free Institute of Science 3(2): Gotze, E 1938. Bau und Leben von Caecum glabrum (Mon-

201-448, 1 map, pis. 13-22. p. 459-473. tagu). Zoologisches Jahrbuch (Systematik), Jena 71(1-2): Dall, VV. H. 1903. Contributions to the Tertiary fauna of 1-190. Florida with especial reference to the Silex Beds of Tampa Golikov, A. N. and Y. I. Starabogatov. 1975. Systematics of and the Pliocene Beds of the Caloosahatchie River in- prosobranch gastropods. Malacologia 15(l):185-232. cluding in many cases a complete revision of the generic Graham, A. 1939. On the structure of the alimentary canal groups treated of and their American Tertiary species. of stvle-bearing prosobranchs. Proceedings of the Zoolog- Part VI. Concluding the work. Transactions of the Wagner ical Society of London (B) 109(1):75-112. Free Institute of Science of Philadelphia 3(4):i-xiv, 1219- Graham, A. 1982. Tornus subcarinatus (Prosobranchia, Ris- 1654, pis. 48-60. soacea), anatomy and relationships. Journal of Molluscan Studies 48:144-147. Davis, G. M. and W, P. Carney. 1973. Description of On- Guppy, R. L. 1866. On the Tertiary Mollusca of Jamaica. comelania lindoensis, first intermediate host of Schisto- J. Quarterly Journal of the Geological Society of London soma japonicum in Sulawesi (Celebes). Proceedings of the 16-18. Academy of Natural Sciences of Philadelphia 125(1 ):1- 22(l):28i-295, pis. 1945. of North Carolina. 34. Hartman, O. The marine annelids Bulletin of the Duke University Marine Station 2:1-54, 10 Davis, G. M., V. Kitikoon, and P. Temcharoen 1976. Mono- pis., 2 charts. graph on "Lithoglyphopsis" aperta, the snail host of Me- Haszprunar, G. 1985. The fine morphology of the osphradial kong River schistosomiasis. Malacologia 15(2):241-287. sense organs of the Mollusca. I. Gastropoda, Prosobranchia. Davis, G. M. and M. Mazurkiewicz. 1985. Systematics of Philosophical Transactions of the Royal Society of London, Cincinnatia winkleyi (Gastropoda: Hydrobiidae). Pro- B 307:457-496. ceedings of the Academy of Natural Sciences of Phila- Hershler, R. and F. G. Thompson. 1987. North .'American delphia 137:28-47. Hydrobiidae (Gastropoda: Rissoacea): redescription and Emerson, W. K. and M K. Jacobson 1976 The American systematic relationships of Tryonia Stimpson, 1865 and Museum of Natural History guide to shells: land, fresh- Pyrgulopsis Call and Pilsbry, 1886. The Nautilus 101(1): water, and marine, from Nova Scotia to Florida. Alfred 25-32. .\. Knopf, New York, 482 p. text -I- xviii p. index, 47 pis Hertlein, L. G. and A. M. Strong. 1951 Eastern Pacific ex- coquilles de la Fischer, P. 1857a. Etudes sur un groupe de peditions of the New Y'ork Zoological Society. XLIII. Mol- (suite). Conchyliologie famille des Trochidae Journal de lusks from the west coast of Mexico and Central .America. 6:168-176. Part X. Zoologica, Scientific Contributions of the New York Fischer, P. 1857b. Etudes sur un groupe de coquilles de la Zoological Society 36(2):67-120, pis, 1-11. famille des Trochidae (fin) Journal de Conchyliologie 6: Hoagland. K. E. 1978. Protandrv and the evolution of en- 284-288, pi. 10. vironmentally-mediated sex change: a study of the Mol- Franzen, A. 1955. Comparative morphological investigations lusca. Malacologia 17(2):365-391. into the spermiogenesis among Mollusca. Zoologiska Bi- Hoagland, K. E. 1984. Sententia. Use of the terms protandry, drag fran Uppsala 30:399-456, pis. 1, 2. protogyny, and hermaphroditism in malacology. Ameri- Fretter, V. 1948. The structure and life history of some minute can Malacological Bulletin 3(l):85-88. prosobranchs of rock pools: Skeneopsis planorbis (Fabri- Humason, G. L. 1962. Animal tissue techniques. W. H. Free- cius), Omalogyra atomus (Philippi), Rissoella diaphana man and Company, San Francisco and London, xv -f

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Page 28 THE NAUTILUS. \'ol. 102, No. 1

Humfrey, M. 1975. Sea shells of the West Indies; a guide to and the Gulf of Mexico. Ph.D. dissertation. University of the marine molluscs of the Caribbean. Taplinger PubMsh- .Miami, Coral Gables, FL, xi + 235 (+ 2) p. ing Company, New York, 351 p., 32 pis. Moore, D. R. 1965. New species of \itrinellidae from Gulf International Code of Zoological Nomenclature [ICZN]. 1985. of Mexico and adjacent waters. The Nautilus 78(3):73-79, Third edition. Ride, VV. D. L.,C. W. Sabrosky. G. Bernardi, pis. 7, 8. and R. \' Mel\ille(eds.). International Trust for Zoological Moore, D. R. 1969. Cyclostrema miranda Bartsch, a synonym Nomenclature, London, .xx + 338 p. of Tornus subcarinatus Montagu. The N'eliger 12(2):169- Iredale, T 1914. On some invalid niolluscan generic names. 170, Proceedings of the Malacological Societv ol London 11(2): Moore, D. R. 1972. Cochliolepis parasitica, a nonparasitic 170-178. marine gastropod, and its place in the Vitrinellidae. Bul- Iredale, T. 1915. The nomenclature of British marine Mol- letin of Marine Science 22(1):100-112. lusca. Journal of Conchology 14(ll):341-346. Morris, P. A. 1973. A field guide to shells of the .Atlantic and Jablonski, D and R. A Lutz. 1980. Molluscan larval shell Gulf Coasts and the West Indies, 3rd ed. Houghton Mifflin morphology Ecological and paleontological applications. Company, Boston, xxviii -I- 330 p., 76 pis. Morton, 1975. Form and habit in some small gastropods of In: Rhoads', D. C. and R. A Lutz (eds. ) Skeletal growth J. New Zealand boulder beaches. The \'eliger 18(1): 1-1 5. of aquatic organisms. Topics in geobiolog) , Vol. 1. Plenum

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Tliiriot-Quie\reux, C. 1977. Particularite de I appareil genital THE NAUTILUS 102(1 ):30-.35, 1988 Page 30

Geographical Distribution of Some Epitoniidae (Mollusca: Gastropoda) Associated with Fungiid Corals

Helen DuShane Research Associate mailing address: Natural History Museum 15012 El Soneto Drive Los Angeles Count) Whittier, CA 90605, USA Los Angeles, CA 90007, USA

ABSTRACT to the tollowing species of epitoniids: Epitonium costulalum (Kiener, 1839), originally described from an The known ranges of three coral associated epitoniids, Epito- unknown locality; Epitonium ulu Pilsbr\. 1921, origi- nium costulalum (Kiener, IS39), Epitonium ulu Pilsbry, 1921, nalK described from Hawaii; and Epitonium hullatum and Epitonium hulldtttm (Snwcrln, 1844) are reviewed (Sowerby, 1844), originalK described from the Philip- pines. An unidentified Epitonium sp. associated with Fungia was also reported by Sabelli and Taviani INTRODUCTION (1984) from the Red Sea. following abbreviations appearing in the text are The family Epitoniidae has had a complicated nomen- The as follows: clatural history, with more than 3,000 Recent and fossils defined

genus Epitonium is an overwhelm- species named. The ANSP—Academy of Natural Sciences, Philadelphia. P.\. ingly large group about which itiucIi has been published, GENEVA—Museum d Histoire Naturelle, Geneva, Sw it- but information on habitats, ecology, growth patterns, zerland. and reproduction is known for relatively few species. NHM, L.A. —Natural History Museum. Los Angeles Some epitoniids are perhaps the only gastropods e.xcept County, Los Angeles, CA. with certain coraiiiophiiidae to have feeding associations Bratcher (Collection —Twila Bratcher, Los .Angeles, C.\. iuiigiid corals. Chane)' Collection —Henry Chaney, Redondo Beach, CA. The stony corals comprise a large group of animals DuShane Collection — Helen DuShane, Whittier, CA. listed showing a great variety of form. Wells ( 1956:F388) Kaiser Collection — Kirstie Kaiser, Park Citv, UT.

1 1 living genera in the family Fungiidae Dana, 1846, and further stated. "Within the family Fungiidae some animals are attached as juveniles, but as adults become RECORDS rubble substrates. Others re- detached and live free on Epitonium costulatum (Kiener, 1839) main attached throughout their lives." There are 11-12 (figures 1, 2) genera, within w liich are probably 49 species. Fungiids live in warm seas in water temperatures 22 °-33 °C. The Discussion: Epitonium costulatum (Kiener, 1839) as

corallum (skeleton) is rounded or elongate, varying in far as presentK know n lives in warm seas from 10°N to length from about 5 cm to 35 cm, and in width from 5 SCN, from the Red Sea, Bay of Bengal (India and Thai- " cm to 20 cm Known commonly as "hat' or "mushroom land), and the Philippine Islands. This is the first report corals, living FuiifJia sp|). arc pink, green, off-white, or of the egg capsules oi E. costulatum found on the un-

pale tan. Distribution of these corals is extensive, from derside ol Fungia sp. from the Red Sea. Each egg capsule the east coast of Africa, across the Arabian Sea, the Bay was encrusted w ilh bits of white coral-sand and attached of Bengal, Western Pacific Ocean and including the to another by two slender threads (figure 2). The number Southern and Flastern Pacific oceans. of egg capsules per mass is approximately 100. Previous reports ol epitoniids associated with Fiingia Hoot (1958) was the first to report this species as an are those of Root (1958:8), Robertson (1963:57, 1965:7, unidentified eiJitoniid li\ing under Fungia, in the Sulu 1970:45), Bosch (1965:267), Taylor (1977:254), Kay (1979: Archipelago, Philippines. His six specimens are at the

152), Bell (1982:508, 1985:161 ). Sabelli and Taviani (1984: Academy of Natural Sciences of Philadelphia (ANSP 92). These papers report at least three species of epito- 230639) and were studied b\ Robertson (1963) who later niids associated w ith species ol Fungia. (1970) compared them with the holot\pe in CJeneva A collection of Epitonium spp. made by SCUBA divers (original locality unknown) and concluded that Root's

in the northern Red Sea verified that Fungia is "home" specimens are indeed £. costulatum. The ANSP speci- H. DuShane, 1988 Page 31

Figure 1. Apertural view of Epitoniitm costulatum (Kiener, 1839). Length 28 mm, width 13 mm, DuShane Collection. Straits of Tiraii, Red Sea Figure 2. Coral-sand encrusted egg mass of £. costulatum. Chaney Collection. Straits of Tiran, Red Sea. Figure 3. Epitonium ulu Pilsbr\, 1921. Length 13.5 mm, width 5.5 mm, Bratcher Collection. Saudi .Arabia, Red Sea. Figure 4. £. ulu with egg mass. Length of shell 16 mm, width 11 mm, Chaney Collection. Tiran Island, Straits of Tiran, Red Sea. Figure 5. .Apertural view of Epitonium bullatum (Sowerby, 1844). Length 13 mm, width 9 mm, Kaiser Collection. Thomas Reef, Sinai, Red Sea. Figure 6. Dorsal view of E. htllatum. Kaiser Collection. Page 32 THE NAUTILUS, Vol. 102, No. 1

mens have 21-26 costae and range in length from 12 4 are laid on the concave underside of Fungia spp. attached to 35.3 mm (Robertson, 1963, 1970). to both the coral and to adult Epitonium. Robertson (1963:60) stated, "There are some wentle- Bosch (1965) reported that Fungia .scutaria Lamarck, traps which seemingly live throughout most ot their post- 1816, from Kaneoke Ba\, Oahu. Hawaii, was infested larval lives with (relatively) large sea anemones (or cor- with both eggs and adults of Epitonium ulu. The snails ..." large amounts of pink tissue containing symbiotic als). Such species are . . . £. aff. costulatum "This had F.pitonitim. the oiiK wentletrap so far found w ith a coe- algal cells and nematoc\sts the\ had ingested. leiiterate other than a sea anemone, presurnabK feeds Taylor (1977) studied the growth rate of Epitonium on Fungia." We now know that £. costulatum lays its ulu feeding on the sea anemone Aiptaaia sp. and found gelatinous egg masses under Fungia on a sandy, rubble that the intracapsular development time for this species substrate, in depths of 2-30 m. Feeding observations are varied from five days to several weeks. She found that lacking. after hatching the ju\ eniles added approximately 0.2 mm Subsequent to 1958, other specimens have been col- of shell length per day. lected from various Indo-Pacific localities. Bell (1982) observed that £. ulu produced a "mean of 32 capsules per day, each capsule containing 500-600 eggs." She later stated (Bell, 1985) that "embryos com- Recent records: Ba\ of Bengal, India. One specimen, plete intracapsular de\elopment in six da\s at 26-27''C length 31 mm, with 17 costae and 12 whorls. DuShane ' and hatch as planktotrophic veligers. Collection. Epitonium ulu has been collected recentK from five Raya Island, Bay of Bengal, Thailand. One specimen, localities.

length 34.5 mm, with 23 costae and 1 1 \\ horls. DuShane Collection. Recent records: Maldive Islands. Robertson (1965) re- Phuket Island. S\V Thailand, Collected by fishermen ported an Epitonium sp. living in association with Fun- under "hat" coral, 15-20 m, 5/22/85. Three specimens, gia. He now identifies the four specimens as Epitonium lengths 23.5, 24.5, 25 mm, with 19, 20, 26 costae and 7, ulu (personal communication, 1986). Guinea. Collected by Tw ila Bratcher, 1980, 9, 12 whorls. DuShane Collection. Papua New Phuket Island, S\V Thailand. Collected by fishermen from under Fungia sp. (figure 3), 11m, water temper- under "hat" coral, 15-20 m, 5/22/85. One specimen, ature 28 °(]. length 16 mm, w ith 30 costae and 11 w-horls. length 23.5 mm, with 20 costae and 10 whorls. NHM, Bratcher Collection, and one specimen, length 12 mm L.A. 124505. with 30 costae and 12 whorls. DuShane (Collection. Tiran Island (SW), Straits of Tiran, Red Sea (27°57'N, Tiran Island (SW), Straits of Tiran, Red Sea (27°57'N, 34°32'E). Collected by Henry Chaney, 10/31/85, SCU- 34°32'E). Collected b\' Henry Chaney. 10/31/85. SCU- BA, 2-5 m in rubble under Fungia sp., four specimens, BA, 2-5 m, with egg mass, under detached Fungia on three specimens, Chaney Collection, lengths 13, 20, 29 rubble substrate. Five live-taken specimens, all exuding specimens, Chanc\ Collec- mm, with 18, 23, 28 costae and 8, 9, 13 whorls; one mucus, were collected. Four specimen, DuShane Collection, length 28 mm, with 24 tion, lengths 3, 4, 5, 16 mm, with 10, 11, 24, 33 costae specimen, DuShane Col- costae and 12 whorls. .\\\ C^haney collected specimens and 9, 10, 12, 12 whorls; one were live-taken, two were pink in color, with gelatinous lection, length 6.5 mm, with 21 costae and 11 whorls, (Collected b\ egg masses as described above. Sinafir Island, Saudi Arabia, Red Sea. Thomas Reef, Sinai, Red Sea (27°59'N, 34°27'E). Col- Twila Bratcher, 10/30/85, SCUBA, 9 m, with eggs under lected by Kirstie Kaiser, 10/30/85, SCUBA, 6.1 m in detached Fungia. Two live-taken specimens, lengths 9, 9. whorls. Bratcher sand pockets under live, detached Fungia sp., one spec- 13.5 mm, with 17, 24 costae and 10 imen, length 23 mm, witli 21 costae and 11 whorls, water Collection. temperature 25 °C, live shell attached to coral. Kaiser Thomas Reef, Sinai, Red Sea (27°59'N, 34°29'E). Col- Collection. lected by Kirstie Kaiser, 10/30/85, SCUBA, 6 m, water of Little Hiva, Maldive Islands. Collected by Henry Cha- temperature 25 °C. .\ttached with eggs to underside detached Fungia. 1\ ing in small sand pockets with sand ney, 8/31/86, SCUBA, 1 m reef sand, under Fungia repanda Dana, 1846, one specimen, live-taken, length and light rubble, l-'our live-taken specimens, lengths 6, 14 mm, with 20 costae and 11 whorls. DuShane Collec- 8, 15, 19 mm, with 17, 20, 23, 32 costae and 10, 10, 13, collected. Kaiser tion. 14 whorls, all exuding mucus when Collection. All three S(CUB.'\ divers (Bratcher. Chaney, Kai.ser) Epitonium ulu Pilsbry, 1921 collected egg masses (figures 4, 8-10) associated with the Sea. The eggs (figures 3, 4, 7-10) specimens of Epitonium ulu from Red were off-w liite w ith light purple spots. One egg sac con- Discussion: Epiloniuiit ulu l'ilsbr>, 1921, a widely dis- tained 4()()-()()0 embr\()s in different stages of de\elop- tributed species, lives in warm seas from 30°N to 5°S, ment. Each translucent egg sac w ithin the cluster is pa- from the Red Sea, to the Maldive Islands, New Guinea pillose over the entire external surface. The papillae are transparent and Haw aii. It is specific to certain species of the stony softly rounded. From each egg sac three coral genus Fungia. Masses of beige colored egg capsules twisted threads are attached at each end of the oval egg H. DuShane, 1988 Page 33

.113 mm

Figure 7. Epitonium ulu Pilsbry, 1921 with egg masses on concave underside of Fungia sp. Figure 8. Egg capsules of £. ulu that appear to be empty and transparent. Figure 9. Developing, opaque embryos of £. ulu located on the inner surface of the transparent sac Figure 10. Prehatching veligers of £. ulu. Diameter of a single snail 0.113 mm.

sac, and eventually twist together to form a single knotted shell the animal appeared as a uniform gelatinous mass mass. Each connecting thread appears to be many times with two dark spots. the length of the sac itself. The egg capsules e.xamined under a microscope (300 x ) Epitonium (Globiscala) bullatuni (Sowerby, 1844) showed the young capsules in three different stages of (figures 5, 6) development. Within each capsule development was uniform. The first stage capsules appeared to be empty and Discussion: Epitonium {Globiscala) bullatum (Sower- transparent (figure 8). The second stage showed devel- by, 1844) lives in warm seas from 35°N to 30°S, from the oping, opaque, white embryos that appeared to be lo- Red Sea, East Africa, New Guinea, Australia, the Phil- cated on the inner surface of the sac, but not filling the ippines, and Japan. Kilburn (1985:330-331) reported this entire compartment (figure 9). The third stage consisted variable species from off Southern Mozambique, and in of well-developed veligers, thus crowding the capsule addition to living under species of Fungia, "under rocks (figure 10). One capsule in the third stage of development in low tide pools, associated with the actinian Fseudac- contained 400-600 free swimming veligers of equal size tinia flagellifera (Hertw.) on w hich it feeds; on occasion and development. The diameter of a single veliger was it ma\ be partly covered by the basal disk of the anem- 0.113 mm. The gibbous shell is transparent e.xcept for one. Juvenile bullatum shelter among coralline algae, the darkened area of the columella. In each embrvonic probably feeding on the small anemones that are at- Page 34 THE NAUTILUS, Vol. 102, No. 1

tached to the fronds." He synonymized Scala (Globis- tions; to William K. Emerson and James H. McLean for cala) papyracea do Boiiry, 1912 from Natal, Epitonium reading the manuscript and offering advice; to the divers,

(Glolmcala) woolacuttae Kersiake, 1958 from South Twila Ikatcher, Henry (^hane\ , and Kirstie Kaiser, who Queensland and GlobUcala kashiuajimensis .\zuma, 1962 returned (rom the Red Sea with epitoniids that formed from Japan with £. bullatum (Sosserby, 1844). the backbone of this paper; to the personnel of the .\llan Hancock Foundation Library, Universit>- of Southern Recent records: Thomas Reef, Sinai, Red Sea (27°59'N, California, for help in locating several rare volumes; to .34°27'E), collected by Kirstie Kaiser, 10/30/85, SCUBA, Da\id Nhilliner and Bertram Draper for the excellent 6. 1 m, water temperature 25 °C. one live specimen from photographs; and to Kirstie Kaiser for the drawings. under live, detached Fungia. length 13 mm, with 40 almost imperceptible costae, 7 whorls, protoconch whorls lost, spiral striations weak when viewed under a micro- LITERATURE CITED scope (10 X ), umbilicate, color white. A second live spec- Azuma, M, 1962. Descriptions of five new species of Japanese imens was eaten by a wrasse (Labridae), before it could Epitoniidae. Venus 22(2);130-136, text figs. 1-6. be collected. An egg mass was observed and photo- Bell, J. L. 1982. Larval development and metamorphosis of graphed with this species but not collected. the prosobranch mollusc, Epitonium ulu, associated with New Guinea. Three specimens in the DuShane Col- a solitary coral. Pacific Science 16:508. in m, lengths lection (ex Withrow Collection), taken 30 Bell, J. L. 1985. Larval growth and metamorphosis of a proso- 11, 11.5, 13 mm, with 8, 9, 9 whorls. branch gastropod associated with a solitar\ coral. Pro- ceedings of the Fifth International Choral Reef Congress,

Tahiti 5:159-164, 6 figs. Epitonium sp. Bosch, H. F. 1965. .\ gastropod parasite of solitary corals in

Hawaii. Pacific Science 19:267-268, 1 fig. An unidentified species of £pi7o;!nu>i was reported taken Boury, E. de. 1912. Description de Scalidae nouveaux ou peu from under Ftingia paunwtensis (Stutchbur\-, 1833) off connus. Journal de Conch\ liologie 60t2):87-108, 1 pi. Saudi Arabia in the Red Sea b\ Sabelli and Taviani Exploring Ex- Dana, J. D. 1846. Zoophytes. United States (1984). Three specimens in varying growth stages were pedition during the vears 1838-1842 under the command

collected with an egg mass at a depth of 2 m. The authors of Charles Wilkes, L .S.N., Vol. 7, p. 1-741 Philadelphia. with a very thin fossils. United States Ex- described their specimens as umbilicate Dana, J. D. 1849. Descriptions of shell, 22 obsolete costae and interspaces crossed by fine ploring Expedition, 1838-1842, under the command of Dana. \'ol. spiral grooves. This is similar to the description of Epi- Charles Wilkes, U.S.N. Geology. By James D. X, .Appendix, 681-729, pars. Atlas, xxi, pis. pars. Phila- tonium ulu, but comparison with E. ulu is needed. p. delphia, 1849. The extensive paper by Jousseaume (1911) treating 58 Jousseaume, F. P. 1911[1912]. Faune malacologique de la species of epitoniids from the Red Sea is of little help in Mer Rouge. Memoires de la Societe Zoologique de France identifying species from that area. Unfortunately, his 24:180-246, pis. .5-7. figures are too small to be of value, and some of his Kay, E. A. 1979. Hawaiian marine shells Reef and shore descriptions are vague. fauna of Hawaii. Section 4: Mollusca. Bernice P. Bishop Museum Special Publication 64(4). Bishop Museum Press, CONCLUSIONS Honolulu, Hawaii, 653 p., 2 figs. Kersiake, 1958. A new Epitonium from eastern .Australia. With an upper surface exposed to the light and a pro- J. Proceedings of the Royal Zoological Society of New South tected concave under surface, species of the genus Fun- Wales for 1956-57, p.' 157-158, 1 fig. as host for several species of epitoniids with gia serve Kiener, L. 1838-39. Species general et iconographie des co- very delicate shells, thin, fragile walls, and numerous quilles vivantes; Genre Scalaire. Rousseau, Paris 7:1-22,

costae that are easiK shattered. Whether the wentletraps pis. 1-7 (text: 1839; plates: 1838). feed on the mucus from the host polyps or the polyps Kilburn, R. N. 1985. The family Epitoniidae (Mollusca: Gas-

themselves is unknown. Taylor (1975) concluded that tropoda) in southern Africa and Mozambique, Annals of Epitonium ulu can, under laboratory conditions, feed the Natal Museum 27(l):239-337, 171 figs. Lamarck, de M. de. 1816. Histoire naturelle des animaux sans on the sea anemone Aiptasia. All other evidence indicates vertebres, Paris 2:1-568. that E. ulu deposits eggs only under Fungia. The coral Pilsbry, H. A. 1921. Marine mollusks of Hawaii. Proceedings is used as the spawning site, the eggs being laid in ge- of the Academy of Natural Sciences of Philadelphia 72: latinous masses, each capsule connected to another by 360-382. ability of epitoniids to locate and two threads. The these Scientific results of the vo\age of Chal- Quelch, J. J. 1886, utilize the underside of F'ungiidae requires further study. lenger. Report on the reef corals collected b\ H.M.S. Chal- An increasingly high degree of specificity is being found lenger during the years 1873-76. London. Zoology 16:1- w ith certain epitoniids seemingK having permanent as- 203. sociations with Fungia corals. Robertson, R. 1963. Wentletraps (Epitoniidae) feeding on sea anemones and corals. Proceedings of the Malacological ACKNOWLEDGEMENTS SocietN of London 35(2&3):51-63, pis. 5-7. Robertson, R 1965 Coelenterate-associated prosobranch gas- My thanks to Robert Robertson for encouragement, and tropods .•Viinual Reports of the .American Malacological for reading the manuscript and making useful sugges- Union for 1965, 32:6-8. H. DuShane, 1988 Page 35

Robertson, R. 1970. Review of the predators and parasites of Taylor, J B 1975 Planktonic prosobranch veligers of Ka- stony corals, with special reference to symbiotic proso- neolie Bay Pli D. dissertation, I'niversity of Hawaii, Ho-

branch gastropods. Pacific Science 24(l):43-54. nolulu, 599 p. rapa in the Snlu Sea. Taylor, R. 1977. rate in juvenile carnivorous proso- Root, J. 1958. Rapa Hawaiian Shell J. Growth News 7:7-8, branchs (Mollusca: Gastropoda) of Kaneohe Bay, Oahu, Sabelli, B. and Marco Taviani. 1984 Red Sea record of a Hawaii. Proceedings Third International Coral Reef Sym-

Fungia-associated epitonid sic]. Bollettino Malacologico posium Rosenstiel School ut Marine and Atmosplieric Sci- 20(l-4):91-94, ence, University of Miami, Miami, Florida 3(l):253-259,

Sowerby, G. B., II. 1844 (July) Descriptions of new species 8 figs.

of Scalaria, collected by Mr. H. Cuming, to be figured in Wells, J. W. 1956. Treatise on invertebrate paleontology. Part the fourth part of Thesaurus Conchyliorum. Proceedings F. Coelenterata. Geological Society of America and Uni-

of the Zoological Societ) of London, pt 12:1-38. versity of Kansas Press, xvii -I- 498 p., 2,700 figs. Stuchbury, T. 1833. .\\\ account of the mode of growth of young corals of the genus Fungia. Transactions of the Linnean Society of London 16:493-498, pi. 32, figs. 6a, b. THE NAL TILLS 102(1 ):36-.39, 1988 Page 36

A New Species of Vasum (Gastropoda: Turbinellidae) from off Somalia

William K. Emerson Waller E. Sage, III Department of ln\ertebrates American Museum of Natural History New York, NY 10024-5192, USA

ABSTRACT zanii (Bianconi, 1851), Bufonaria fernandesi Beu, 1977, Vasum crosseanum (Souserbie, 1875), Tudicula zartzi- Vasum stephanti new species is described from moderately barica Abbott, 1958, Metula bosu:ellae Kilburn, 1975, deep water off Cape Gardafui (Ras Asir), Somalia and com- CucuUaca labiata (Lightfoot, 1786), and Chlamys tonn- pared with congeners. sendi (Sowerb>, 1895).

SYSTEMATICS INTRODUCTION Family Turbinellidae Swainson, 1840 In the past few years, deep-sea commercial fisheries op- Subfamily Vasinae H. & .\. .\dams, 1853 erations off the northeastern coast of SomaUa have resulted in the discovery of several new or otherwise in- Genus Vasum Roding, 1798 teresting species of moilusks. Lorenz (1987:11) described Remarks: See Abbott (1959) and \okes (1966) for re- P.seudosiiunia wieseonim new species from the region views of this subfamily. off (^ape Ras Hafun in about 300 m, and recorded the presence oi Festilyria jestiva (Lamarck, 1811), S(romi>us oldi Emerson, 1965, and Cijpraea broderipii Sowerby, Vasum stephanti new species 1832, from the trawl samples. Waller (1986:39-46) de- (figures 1-6) scribed Somalipecten crannwrorurn new genus, new Diagnosis: Similar to Vasum tubiferum (.-^nton, 1839) species, from "off Somalia, depth 150-300 m ", obtained in general appearance, but differs in having a more tri- from Taiwanese fishermen, and also provided a list of angular outline, three equalK well-de\ eloped columellar associated species. Another species recently described plaits (in place of three major, plus one or two minor from off Somalia is Vulutocorbis rosavittoriae Rehder, plaits), a nearly uniformly milk-w hite shell w ith a white, 1981. glazed aperture and parietal wall (compared to an or- Through the good offices ot John Bernard, our atten- ange-brown to yellowish shell with the parietal wall a tion was called to an umiamed species of Vasum trawled light tan w ith \er\ large splotches of chestnut to purple- in Somalian waters. We take pleasure in describing this brown), and in the presence of two or three rows of spines new species in honor of Adolphe Stephant, w ho obtained at base of shell (instead of one row). the specimens from Danish shrimpers and generously provided Mr. Bernard with the data and specimens. Oth- Description: Shell moderately large for genus, attaining er species of moilusks reported by Mr. Stephant (in Hit., 108-f- mm in length. Solid, heavy, turbinate, and strongly August 19, 1987) to have been taken during these trawl- spined. Spire ele\ated with a short, smooth, bulbous nu- ing operations include h'cstihjria jcstiva, titwrnbits oldi, cleus of 1'2 whorls (figures 5, 6). Postnuclear whorls 7 S. piicatus (Roding, 1798), Phalium microstoma (von (adult specimens lack complete spires), the body whorl Martens, 1901), P. bituberadosnm (von Martens, 1903), with 7 to 8 well-developed, curved to strongly recurved, Ficus investigatoris (E. A. Smith, 1894), Cytnatium ran- flaringly grooved and terminally open, subsutural spines.

Figures 1-6. V'a.sinn stephanti new species. 1, 2. Paratype, AMNll 225988. 3, 4. Holot\pe, AMNH 225987. 5, 6. Paratype, x AMNH 225989 (details of spire). All from type locality: off Cape Gardafui, Somalia, see text; figures 1-4 approximately %. figures 5, 6, x 2. W. K. Emerson and W. E. Sage, III, 1988 Page 37 Page 38 THE NAUTILUS, Vol. 102, No. 1

1. stephanti new species. Shell measurements A row of similar but much shorter spines below the first Table Vasum in and number of whorls; width measured including spines, row, followed by 5 coarse spiral cords and intervening mm n = 12. Spires incomplete e.xcept for 5, 10 raised lines. Base of shell with 2 to 3 spiral rows of moderateK developed, groove spines and lower surface l.rliUtl] Wiiltli = \\1 with weakK developed spiral lines. Parietal wall thickened, slightK raised, glazed. (Columella with 3 plicae, Paratype 1 107 68 posterior 2 better developed; first posterior plica semi- Parat) pe 2 Holotype bifid in 1 specimen. Outer lip moderateK thin, slightK' Paratype 3 reflected, crenulated. Umbilicus funnel-shaped and in Parat) pe 4 most specimens widely open. Base color of shell milky Paratype 5 white, spire stained buff. CJolumella glazed. Aperture Paratype 6 white, with a slight bluish tinge. Periostracum moder- Paratype 7 ateK thick, tannish brown, and somewhat foliaceous. Soft Paratype 8 parts not seen. Operculum brown, corneous, unguiculate, Paratype 9 apicalK nucleate, filling most of the aperture with foot Referred fulK u ithdrawn. Inner surface marginally thickened on Parat\|)c II) basal and abcolumellar sides, central area depressed and with irregular concentric rings; outer surface scabrous.

Type locality: 13-16 km east, 80-96 km south of Cape Gardafui (Ras Asir), Somalia, trawled by shrimp fisher- man in 183 to 220 m, December, 1986.

Range: Known only from the type localit\ and in the Gulf of .'Kden off the Bari coast of Somalia.

Material examined: Holotype, AMNH 225987, 102.36 mm, ex E. Schelling Collection (figures 3, 4); paratype

2, AMNH 225988 (figures 1, 2), paratype 5, AMNH

225989 (figures 5, 6), paratype 8, AMNH 225990, ex J.

Bernard Collection; paratypes 1, 3, 4, 7, 9 J. Bernard Collection; paratypes 6, 10 A. Stephant Collection, all from the t\pe locality; referred specimen, H. Lee Col- lection, Alula, Bari coast, Somalia (.see Table 1).

Remarks: As noted above, Vasum stephanti new species most closely resembles in shell morphology the endemic Philippine (Cuyo-Palawan group) species V. tubiferum (Anton, 1839:70; Kobelt, 1876:155, pi. 9, fig. 3; Abbott,

1959:20, pi. 4, fig. 1; Spring.steen & Leobrera, 1986:105,

pi. 28, fig. 6). Anton's taxon and the closeK related V.

turbinellus (Linne, 1758:750; Abbott, 1959:17, pi. 1, figs.

2, 3), which ranges from East Africa to the western Pa-

cific, are inhabitants of shallow water, as are the other four Indo-Pacific species assigned to Vasum {sensu stric- to) by .Abbott (1959). One of these, V. rhinoceros (Gme-

lin, 1791 ), from Kenya and Tanzania, is somewhat similar but has a lower-spired, heavier shell with massive nodules, thickened and reflected outer lip, and a brown- blotched to light yellow parietal wall (Abbott, 1959:21,

pi. 4, figs. 3, 4). Strongly spino.se specimens with immature outer lips of the Brasilian V. cassiforme (Kiener, 1840), cited from low tide to 60 m (Rios, 1985:115), are superficially similar to the new species. (See Abbott & Dance, 1982:209, 210 for polychrome illustrations of these taxa.) The Australian Altivasurn Hedle\, 1914, and several species of Indo-Pacific Tudicula H. & A. Adams, 1864 are known to occur in moderate depths (to 220 m). Some of the spinose species originalK assigned to Tu- dicula (e.g., T. zanziharica Abbott, 1958, Iron) the western Indian Ocean, and T. rasilistorna Abbott, 1959), from W. K. Emerson and W. E. Sage, III, 1988 Page 39

kiener, L. C;. 1840[-41]. Species general et icoiiographie des Rios, E. C. 1985. Seashells of Brazil. Fundagao Sociedade do

coquilles vivantes. . . . Paris, Genre Turbinelle 6(59-71): Rio Grande, Funda^ao Universidade do Rio Grande, Mu- 1-50, 1841; pis. 1-21, 1840 (Vasum cassijormc (Kiener) seo Oceanografico. 329 p., 102 pis. dates from the citation on plate 9 of "Turhinclla ca.ssi- Souverbie, S. M. 1875. Description d une espece nouvelle formis \'alene." [= Valenciennes]; the text was pnhlislied appartenant au genre Titrhinella. Jonrnal de (>'onchylio- in 1841). logie 23(4):297-298. F. Leobrera. 1986. Shells of the Kobelt, H. C 1876. Systematisches Conchylien-Cabinet von Springsteen, F. J. and M.

Martini und Chemnitz. ... Niirnberg. Purpur.schnecken, Philippines. Carfel Seashell Museum, Manila, 377 p. Band 3, Abt. 3a, pt. 251:121-164. Vokes, E. H. 1966. The genus Va,si(ni (Mollusca: Gastropoda) Linne. C, von. 1758. Systema naturae per regna tria naturae, in the New World Tulane Studies in Geology 5(l):l-36.

10th ed. Stockholm, \ol. 1, Regnum animale, 284 p. Waller, T. R. 1986. A new genus and species of scallop (Bi- Description of new Ovulidae species valvia: Pectinidae) from off Somalia and the definition of Lorenz, F , Jr. 1987 a from Somalia (Gastropoda: Ovulidae). La Conchiglia, Rome a new tribe Decatopectinini. The Nautilus 100(2):39-46. 19(214-215):11-12. THE NAUTILUS 102(1 ):40-45, 1988 Page 40

Density, Spatial Distribution, Activity Patterns, and Biomass of the Land Snail, Geophorus bothropoma Moellendorff (Prosobranchia: Helicinidae)

Kurt Auffenberg Troy Auffenberg Malacology Division Florida State Museum University of Florida Gainesville, FL 32611

ABSTRACT birds, Odum & Kuenzler, 1955; frogs, Turner. 1960; insects, Alexander, 1961, Iwao, 1970; mammals, Stumpf & The results of preliminary work done in the Republic of the Mohr, 1962). While the database with which we worked Philippines during July, 1981 on the density, spatial distribu- was rather small, the ease ot anaKses and the application, activity patterns, and biomass of the Philippine Island bility of the results suggest that such to sim- land snail Geophorus bothropoma (Prosobranchia, Helicinidae) approaches are presented. ilar questions in terrestrial snail ecolog) would be highly Density was estimated at 0.55/m^ utilizing the quadrat meth- beneficial. od. Using the parameters of mean crowding and patchiness, the species was shown to be uniformly distributed spatially, possibly due to mutual repulsion. Individual activity distance STUDY AREA per 24 hour period varied greatly (0.0-275.0 cm) and was The study area is located in the Republic of the Phil- possibly correlated with rainfall Utilizing the convex polygon and ellipsoid methods, minimum individual activity ranges for ippines, Luzon Island, Camarines Sur Province. Cara- the duration of the study were estimated to be 2.4-4.2 m^. moan Municipality, 1.0 km south of Barrio Ilawod. The Biomass (live weight) of adults was estimated at 0.13 g/m'^. work was conducted 10 m off the trail between Ilawod The species is diurnal and does not home. and Gota Beach on a large (8 x 9 x 3 m high) limestone boulder with almost vertical sides. Key words: Gastropoda; Prosobranchia; Helicinidae; Geopho- rus; density; spatial distribution; activity; biomass; Philippine Much of the Caramoan Peninsula consists of well- Islands. developed limestone karst of Miocene reef origin. Be- cause the area has been relativeK recently uplifted, the mountains are rather steep. Hills and ridges often consist INTRODUCTION of masses of large limestone boulders with chasms tens of meters deep. Runoff is rapid and though rainfall may

There are relatively few published studies dealing with be high seasonally, the surface environment is often rel- the density, spatial distribution, activity patterns, or bio- atively dry. Surface water collects in solution pits on mass of land snails (see Discussion section for citations). limestone and in hollows of trees. Of those that do exist, almost all have dealt with pul- Due to the rugged topography, the area remains blan- monates of temperate regions, while tropical terrestrial keted in a multi-canopied, tropical evergreen forest. prosobranchs remain virtually unstudied. The present Vegetational communities are extremely diverse. The paper reports on the results of a preliminary study con- forest is dominated by tall dipterocarp trees in both upper ducted July 7-17, 1981 on Geophorus bothropoma Moel- (30-40 m) and lower (5-20 m) canopies; a dense her- lendorff, 1895 (Quadras & Moellendorff, 1895:148; Wag- baceous layer of mainK ferns exists near the ground ner, 1908:152, pi. 29, figs. 20-23), a large (13.0-16.0 mm surface. in diameter), limestone rock-dwelling helicinid endemic The Caramoan Peninsula has no definite dry season. to the Caramoan Peninsula, Luzon, This represents one While it tends to rain throughout the \ear (ca. 2,900 of the very few times quantitative ecological methods mm), most rainfall occurs from September through Jan- have been utilized for the study of terrestrial mollusks uary, with a minor peak in July associated with the onset in all of Southeast Asia. In addition, the present paper of the typhoon season. introduces to ecological malacology several analyses used As in other tropical rainforests, the Caramoan Pen- by zoologists working in groups other than mollusks (i.e., insula shows little seasonal temperature variation. Mean 1 ,

K. Auffeiiberg and T. Auffenberg, 1988 Page 41

iluil\ temperature fluctuations are greater tPian mean Anal) sis of the activity of Geophorus buthropoma was seasonal variation. However, there is a marked vertical made by subdi\idiiig the I m- quadrats, plotting the tliermocline within the forest, in which temperatures exact location of individuals and measuring straight line from the surface to 1 m are most stable; daily variation distances (nearest cm) from the last observation with a increases at greater heights above the ground. Due to flexible tape. This provided a crude estimate of the min- the dense vegetation at the study site, the effects of solar imum distance tra\eled since the last ob.servation. Cal- railiation and wind are minimal. Therefore, Geophorus culations are based only on data provided by two or more butliropoma lives in the most stable part of the local consecutive daily observations (N = 27). warm, moist tropical forest environment. The above eco- Estimations of activity ranges (N = 5) are based on logical data is from Auffenberg (in press). the convex polygon method and the ellipsoid method,

which is based on a covariance matrix of capture loci METHODS (Jeunrich & Turner, 1969). The convex polygon method involves plotting all the recapture points, drawing the An 8 m- (2 m tall, 4 m wide) vertical grid of 1 m- quadrats smallest convex polygon containing these points and then was constructed against the east side of the boulder to determining the area. This method, defined by the equa- measure density and activity. All adult Geophorus both- tion: ropoma within this grid (N = 14) were numbered with small dots of fingernail polish on the dorsal surface of x.y^+i - Xi+iYi A = (2) the shell. This increased shell visibilit\ , but is not believed to have increased predation due to the nocturnal habits is relatively simple but is biased b\ sample size. The of most potential predators. No additional individuals values tend to increase as the number of capture points were located during subsequent visits. The site was vis- increases. ited nine times during the next 11 days at 10 a.m. for The ellipsoid method is not biased by sample size and appro.ximateK 1 hour observations. In addition, the site that individuals are active outside the observed was visited sporadically throughout the daylight hours assumes range of activity (see Jennrich & Turner, 1969, for dis- and seven times at night (8-10 p.m.). This provided a set of successive censuses, not a random quadrat sampling. cussion). This method is defined by; Densit\ estimates are based on the ciuadrat sampling 67r|S| (3) method. This method simply involves counting individuals occurring within a quadrat of fixed size. where |S| is the determinant of the capture point co- Analyses of spatial distribution are based on m* (mean variance matrix; crowding) and m* m (patchiness) of Lloyd (1967). Mean

is given the formula: crowding by S S q S = (4) s s

- (1) defined by the equations;

1 x;^ where x, = the number of individuals in the jth quadrat = the total number of quadrats. and q S„ - o 2 (y. y)^ Mean crowding is defined as the mean number of other n - 2 individuals per quadrat per individual. It expresses the degree of spatial crowding experienced by an individual = = —!— - S. S,. — "2 (X, x)(y, because of others of the same species (or different species; n 2 see Lloyd, 1967). A particularly important feature of this - - statistic is that it is relatively independent of spatial dis- X 7v X, tribution t\ pe, number of samples, and size of the means. The most complete discussion concerning the utility of = - y. (5) this statistic is by Iwao (1977). It should be stressed that y 2j this statistic does not measure crowding in an ecological sense, as man\' variables (i.e., food availabilit\', territo- Live adult snails (N = 90) were collected near the rialit) ) must be examined before this can be determined study site and their weights (in groups of 10 individuals) (Lloyd, 1967). were taken to the nearest milligram on a single beam,

Patchiness (m* /m) is the ratio of mean crowding to double pan scale. Fifty specimens were then macerated; mean densit\ (Iwao, 1968). This index provides a relative the shells and opercula were rinsed, dried for several measure of aggregation. It equals unity (1) in random days and reweighed. distribution, is greater than unity in contagious distri- Voucher specimens of Geophorous bothropoma are butions and is less than unity in regular (uniform) dis- housed in the mollusk collection of the Florida State tributions (Iwao, 1977). Museum (UF 56667). Page 42 THE NAUTILUS, Vol. 102, No. 1

Table 1. Densit\ and spatial distribution parameters based on quadrat method, n = individuals recaptured per visit, m/m- (lciisil\, m* = mean crowdiiij;, m* m = patchiness. Visit 6 is (irnitted from densit\ and spatial distribution analyses Page 43 K. Auffenberg and T. Auffenberg, 1988

Estimates of individual activity ranges (N = 5) (figure Barnes & Weil, 1944, 1945; Wells, 1944; Dainton, 1954a,b; 1963; Cameron, 1970; Crawford- 2) are quite consistent. The convex polygon method (cor- Blinn, 1963; Henne, rected for sample size bias; Jennricli & Turner, 1969) Sidebotham, 1972; Baker, 1973; Bailey, 1975; Shachak yielded activity ranges of 2.3-3.8 m- (x = 3.1 ± 0.3, et al., 1975; Heatwole & Heatwole, 1978; Deisler, 1987). SD = 0.5 m-). the ellipsoid method yielded slightly (but It is apparent from these studies that the initiation and insignificantly), larger area estimates of 2.7-4.2 m- (x = continuation of activity is a variable response to complex, perhaps 3.3 ± 0.3, SD = 0.6 m-). simultaneous climatic changes (some minute, immeasurable) in the microenvironment. More field and Biomass: Mean total live weight per adult individual laboratory studies must be undertaken before this will the study area was 0.59 (N = 90). Mean shell weight in g be understood. In previous studies snail movements were (including operculum) per individual was 0.35 (N = g most numerous shortly before or after nightfall and short- 52), yielding a mean live biomass per individual of 0.24 ly before dawn. However, Geophorus bothropoma is mean density estimated by the quadrat method g. The diurnal rather than crepuscular, and the congener G. of this species in this area is 0.55 individuals/ (table 1) trochiformis (Sowerby, 1842) was also observed active m-. Thus the estimated biomass is 0.13 g/m-. throughout the day. We believe this behavioral pattern

is not a general trend in tropical terrestrial prosobranchs. Moellendorff, 1895 and Japonia DISCUSSION Cyclophorus ceratodes ciliata (Sowerby, 1843) and J. stephaiiophora (Moellen- Density and spatial distribution: Previous studies on dorff, 1895) living in the same study area are decidedly species comparable in size to Geophorus bothropoma crepuscular in their activity patterns (personal observa- have been few (i.e.. Mason, 1970; Richardson, 1975; tion). Cameron, 1982), and none concerns terrestrial proso- Few data are available on activity and home site ranges branchs. The only prosobranchs with available data are of terrestrial mollusks. Heatwole and Heatwole (1978) the small hydrocenid Georissa monterosatiana (Godwin- found individual home site ranges of the camaenid Car- .•\usten & Nevill, 1879) and the diplommatinid Opis- acolus caracoUus (Linne, 1758) to vary greatly (0.08- thostoma retrovertens Tomlin, 1938 of Malaysia. Berry 59.0 m-). Since Geophorus bothropoma does not home (1966) found these species occurring in higher densities we prefer to call the area enclosed by the recapture sites on moss covered rocks than on moss-free rocks, but did (figure 2) the activity range. The term homesite range not address spatial distribution. In the Caramoan area implies that the individual possibly returns to a certain we found Geori.'isa rufescens (Moellendorff, 1887) in ex- area after foraging outside that area. We assume that tremely localized and highly aggregated populations on the activity range of Caracolus caracoUus is much larger vertical rock faces. This is very different from the large than the enclosed polygon determined by recapture sites.

Geophorus bothropoma, which is found on virtually every Long-term study is needed to provide a meaningful vertical rock face and is uniformly distributed spatially. estimate of the activity range of Geophorus bothropoma. In this stud\-, intersample variation in estimated mean The data presented here only provide an estimate of the densities is probabK largely due to the limited size of minimum size of the activity range expected during a the study area. The choice of study area size and number wet part of the year. Activity ranges are probably sig- of samples must have some biological basis to estimate nificantly smaller during the dry season. population density and spatial distribution adequately. Homing is apparently based on chemoreception (see

This is usually done by preliminary sampling or guessing Cook, 1979, for review). The advantages and/or disad- (Ivvao, 1977). However, we do feel confident that the vantages of homing have not been properly addressed. proper quadrat size (1 m^) was chosen because the ob- A permanent resting site should provide relative safety served dail\ activity was approximately 1 m (x = 93.7 ± from predators and fulfill the physiological requirements 13.31 cm). Density estimates were possibly also biased of the individual at rest. The physiological and/or be- by the rugged microhabitat. The limestone karst provides havioral basis for Geophorus bothropoma being unable man> cracks, crevices, and solution pits in which a snail to (or not having to) home is not known. Although the could remain unseen, despite our thorough searches. operculum provides protection from small predators and Utilizing Iwao's (1968) interpretation of m*/m regres- desiccation, this does not explain this species' lack of sions, we determined that individuals of Geophorus both- homing behavior. Geophorus bothropoma most com- ropoma were uniformK distributed and may have ex- monly rests on vertical rock faces. Even when inactive, hibited "mutual repulsion. Infra-specific competition it does not retract fully into the shell, leaving the oper- has never been properly substantiated in studies on ter- culum non-functional. Because Cyclophorus ceratodes

restrial mollusks and we cannot substantiate it here. Moellendorff, 1895 and Japonia ciliata (Sowerby, 1843) were found Nevertheless, density was relatively low, despite the and J. stephanophoru (Moellendorf. 1895) seemingly adequate food and resting site resources that to return to the same solution pits after foraging (personal could have been expected to sustain substantialK' higher observation), non-homing is not a general trend in trop- snail densities. ical terrestrial prosobranchs.

.Activity: Previous studies have shown terrestrial mol- Biomass: Few data on the biomass of terrestrial mollusks to be very cyclic in their activity patterns (i.e.. lusks are available (Strandine, 1941; Jennings & Bark- Page 44 THE NAUTILUS, Vol. 102, No. 1

ham, 1975; Richardson, 1975; Cameron, 1982) and none numbers, activities, and distribution. Part 2. Journal of concern prosobranchs, nor do they pro\ ide eas\ or direct .\nimal Ecology 1471-105 Berry, .A 1966 Population structure Quctuations comparisons. Man\ more species must be studied before J and in the snail fauna of a Malayan limestone hill. general trends can be demonstrated. Journal of Zoology, London 150:1 1-27. Fragments of one marked specimen were found in a Blinn, W. C. 1963. Ecolog\ of the land snails Mesodon thy- solution pit, possibly preyed on by the nocturnal shrew, roidus and Allogona profunda. Ecology 44(3)498-505. Suncus marinus (Linne, 1766), an introduced species. Cameron, R. A. D. 1970. The effect of temperature on the This species was seen on two occasions at the study site. activity of three species of helicid snail (Mollusca: Gastrop- shells of Geophorus bothropoma gnawed open at Many oda). Journal of Zoolog\ . London 162:303-315. the periphery were found in solution pits nearbs . The Cameron, R A D 1982 Life histories, densit\ and biomass diurnal, snail-eating monitor lizard Varanti.s grayi Bou- in a woodland snail communit\ . Journal of Molluscan Stud- lenger, 1885 rarely preys on this species (Auffenberg, in ies 48:159-166 Cook, A. 1979. in 18:315- press). Homing gastropods. Malacologia 318. Although short-term data such as those presented here Crawford-Sidebotham, T. 1972. The influence of weather add much to our knowledge of this species, onl\' with J. upon the activity of slugs, Oecologia 9:141-154. long-term life-histor\ studies will ([uestions be answered Dainton, B H. 1954a. The activit\ of slugs. I The induction with confidence. with morphological infor- Combined ot activity b) changing temperatures. Journal of Experi- mation, sound behavioral and ecological data uill un- mental Biology 31:165-187. doubtedly contribute greatK to a better understanding Dainton. B H. 1954b The activity of slugs. II, The effect of ot the systematics and e\olution of Philippine helicinids. light and air currents. Journal of Experimental Biology 31: 187-197.

Deisler, J. E. 1987. The ecology of the Stock Island Tree Snail. ACKNOWLEDGEMENTS Orthalicus reses reses (Say). Bulletin of the Florida State Museum, Biological Sciences 31(3):107-145. We would like to thank the U.S. Fish and Wildlife Service Heatwole, H. and A. Heatwole. 1978. Ecology of the Puerto and the New York Zoological Society for herpetological Rican camaenid tree-snails Malacologia 17(21:241-315. grants to Walter Auffenberg that allowed us the oppor- Henne, F. C, 1963. The effect of light and temperature on tunit) for the present stud>. The Philippine Parks and the locomotorv activit\' of Polygyra albolahris (Sa\ ), Bios 34:129-133, Wildlife office was very helpful in issuing appropriate Iwao, S. 1968, A new regression method for the aggregation permits. The illustrations were drawn by Armando Gar- pattern of animal populations. Research in Population cia and Marcos Guinaldo. We appreciate the patience, Ecology 10:1-20. time, and insight provided bv Irvy Quitmyer, Mark Iwao, S. 1970. Analysis of contagiousness in the action of Michael W. Binford of the Florida State Brenner, and mortality factors on the Western Tent Caterpillar popu- Museum during the computer assisted regression anal- lation b\ using the m*-m relationship. Research in Pop- ysis. Glenn A. Goodfriend provided suggestions during ulation Ecology 12:100-110, early data analysis. Robert .\. (>ameron provided many Iwao, S, 1977, The m*-m statistics as a comprehensive method constructive suggestions for analyses and format. We also for analyzing spatial patterns of biological populations and its application to sampling problems. In: Morisita. M (ed). thank Arthur J. Cain, Robert Hershler, and Fred G. Studies on methods of estimating population density, bio- Thompson for reviewing the manuscript. Special thanks mass and productivity in terrestrial animals. JIBP Synthe- are extended to our father, Walter Auffenberg, for his sis, University of Tokyo Press 17:21-46. tireless assistance in the field and for his time and guid- Jennings. T. J and J P Barkham. 1975, Slug populations in ance during the writing of the manuscript. mixed deciduous woodland. Oecologia 10:279-286.

Jeiuirich, R. I, and F B. Turner. 1969 Measurement of non- circular home range. Journal of Theoretical Biology 22: LITERATURE CITED 227-237, Lloyd, M, 1967, 'Mean Crowding Journal of ,\niinal Ecolog) Alexander, R D. 1961, .Aggressiveness, territoriality and sex- '36(l):l-30, ual behaviour in field crickets (Orthoptera: (iryllidae). Be- Mason, F. 1970. Snail populations, beech litter production, haviour 17:130-223. C. and the role of snails in litter decomposition. Oecologia 5: Auffenberg, W. In press. The behavioral ecology of Gray's 215-239. Monitor Lizard. University Presses of Florida E, P, and E. Kuenzler, 1955, Measurement of ter- Bailey, S. R. 1975. The seasonal and daiU patterns of loco- Odum, J 72:128-137, motor activity in the snail Helix asperse Muller, and their ritory and home range size in birds. .\uk (Quadras, F, 1895, relation to environmental variables. Proceedings of the J, and O, F, von Moellendorff Diagnoses Malacological Society of London 41:415-428. specierum novarum ex insults Philippinis auctoribus, Baker, A. N. 1973. Factors contributing towards the initiation Nachrichtsblatt der deutschen Malakozoologischen Ge- of slug activity in the field Proceedings of the Malaco- sellschaft 1895(9-10):137-149. logical Society of London 40:329-333. Richardson, A. M. M. 1975, Energy Dux in a natural population of the land snail, Cepaea nemoralis Linne. Oeco- Barnes, H. F. and J. W. Weil 1944 Slugs in gardens: their

numbers, activities, and distribution Part 1 Journal of logia 19(2):141-164. Animal Ecology 13:140-175 Shachak, M., Y. Orr. and Y. Steinberger. 1975, Field obser-

Barnes, H. F. and j VV Weil 1945. Slugs in gardens: their vations on the natural history of Sphincterochda (S.) zona- K. Auffenberg and T. Auffenberg, 1988 Page 45

ta (Bourguignat, 1853) (=S. boissieri Charpentier, 1847). in Yellowstone Park, Wyoming. Ecological Monographs Argamon, Israel Journal of Malacology 5(l-4):20-46. 30:251-278.

Strandine, E J. 1941. Quantitative study of a snail population. Wagner, A. 1908, Die Familie der Jlelicinidae. In: Syste- Ecology 22:86-91. malisches Conchylien-Cabinet von Martini und Chemnitz Stumpf, W. A. and C. O. Mofir. 1962. Linearit\ of home ia8):113-184.

ranges of California mice and other animals Journal of Wells, G. P. 1944. The water relations of snails and slugs. III. Wildlife Management 26:149-154. Factors determining activity in Helix pomatia L. Journal Turner, F. B. 1960. Population structure and dynamics of a of Experimental Biology 20:79-87. Western Spotted Frog, Rana p. pretiosa Baird and Girard, ^^ THE NAUTILUS 102(1 ):46, 1988 ^^S^

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Volume 102, Number 2 April 29, 1988 ISSN 0028-1344

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Volume 102, Number 2 April 29, 1988 ISSN 0028-1344 CONTENTS

Joseph R. Pawlik The egg capsules, embryos, and larvae of Cancellaria John B. O'Sullivaii cooperi (Gastropoda: Cancellariidae) M. G. Harasewych

Martin Avery Snyder Latirus nuirtini (Gastropoda: Fasciolariidae), a new species from Honduras 54

Robert Hershler Shell variation of springsnail populations in the Cuatro Lee-Ann C. Havek Cienegas Basin, Mexico: Preliminary analysis of Limnocrene Fauna 56

Eva Pip Niche congruency of freshwater gastropods in Central North America with respect to six water chemistry parameters 65

B. W. Kilgour

THE NAUTILUS 102(2):47-53, 1988 Page 47

The Egg Capsules, Embryos, and Larvae of Cancellaria cooperi (Gastropoda: Cancellariidae)

Joseph R. Pawlik John B. O'Sullivan M. G. Harasewych' Scripps Institution of Oceanography Monterey Bay Aquarium Department of Invertebrate Zoology UniversitN of California, San Diego 886 Cannery Row National Museum of Natural History La Jolla, CA 92093, USA Monterey, CA 93940 Smitfisonian Institution Washington, DC 20560, USA

ABSTRACT tions of its egg capsules and embryonic and larval development, and reviews the available data on the The egg capsules, and the embr\onic and larval development reproductive biology and larval development of the Can- of the cancellariid gastropod Cancellaria cooperi Gabb are cellariidae. described. Egg capsules are spatulate in form, having long, narrow stalks that support the eggs above the surrounding sand. Egg capsules contain 4,000-5,000 eggs (165 ^ni in diameter), MATERIALS AND METHODS which undergo typical prosobranch development to hatch as planktotrophic veligers after 27 da> s at 15 °C Larvae in culture Twenty-three specimens of Cancellaria cooperi were col- grew from 305 ^m to 890 ^m in shell length over 30 days, but lected on the artificial reef "Torrev Pines #1", off San died before metamorphosis. Limited comparative data suggest Diego, CA (32°53'12"N, 117°50'50"'w) at depths of 20- that long stalked egg capsules are known only in members of 22 m using SCUBA. The animals were maintained in an the Cancellariinae, that opercula, absent in all adult Cancel- aquarium containing sufficient sand for complete snail lariidae, are present or prominent in the late larval stages of burial (4-8 cm depth) and supplied with a continuous at least some species, and that developmental type cannot be flow of seawater (12-16 °C). Snails were allowed to feed inferred from protoconch morphology using the criteria of Shu- electric rays basis, to (1974) in a majority of cancellariid species. on on a bimonthly and had been maintained in this manner for at least 6 months prior to Key words: Reproduction; development; larvae; eggs; egg cap- the onset of oviposition. sules; Cancellariidae; Cancellaria. Individual egg capsules were freed from the aquarium bottom and maintained in beakers containing continu-

ously aerated, 1 ixm filtered seawater at 15 °C. Developing INTRODUCTION embryos were removed with a glass pipette through an incision cut along the narrow edge of the egg capsule.

Little is known about the reproductive biology and early Hatched larvae of Cancellaria cooperi were cultured development of most of the approximately 200 species following procedures described by Paige (1986), except that constitute the neogastropod family Cancellariidae. that 1 /um filtered natural seawater containing 40 mg/ The few published reports (Morch, 1869; Thorson, 1935, liter each of the antibiotics streptomycin sulfate and so- 1944; Knudsen, 1950; MacGinitie, 1955; Kilburn & Rip- dium penicillin G was used instead of artificial seawater. pey, 1982; Bouchet & Waren, 1985) are limited to de- Larvae were fed a mixture of the green flagellates Iso- scriptions of egg capsules and, in some cases, ova or larval chrysis galhana (Park, 1949) and Pavlova lutheri (Droop) shells attributed to cancellariids, usually on the basis of Green, 1975 at 10^ cells/ml. Prior to photography, larvae the proximit\' of living snails. were narcotized in a 1:3 mixture of a saturated aqueous During studies of the diet and feeding behavior of solution of chlorobutanol in seawater.

Cancellaria cooperi Gabb, 1865, a species that is attract- Eggs and larvae for SEM examination were fixed in ed to and speciBcall) parasitizes the Pacific electric ray 2% formalin in seawater, and stored in 70% ethanol. Torpedo califarnica .\>res, 1855 (O'Sullivan e/ ai. 1987), Specimens were critical-point dried and photographed a number of these snails were observed producing egg using a Hitachi S-570 Scanning Electron Microscope. capsules. The present study supplements our know ledge of the natural history of this cancellariid with descrip- RESULTS

Beginning April 21 and continuing through Ma\ 19, 1986, a total of 15 egg capsules were laid by at least three snails (mean shell length = 62.3 mm), with nine Author for correspondence. capsules being produced sequentially b>- a single female. Page 48 THE NAUTILUS, Vol. 102, No. 2

Figure 1. Cancellaria cooperi Gabb. Apertural and right side view of female specimen collected by SCUBA in 20-22 m at artificial reef "Torrev Pines #1", off San Diego, CA {32°53'12"N, !17°50'50"W), USNM 846054. 1.0 x.

Oviposition generally occurred at night or in the morn- Figure 2. Egg capsule of Cancellaria cooperi Gabb. Frontal ing. Snails emerged partialK from the sand, with their and side views 1.0 x. detail of hatching aperture 6.0 x. mp, raised foot spread anteriorK and enfolded posteriorly, mucous plug; li, membranous sack. the emerging stalk protruding from the folded portion of the foot. Over the course of several hours, the egg case which was enveloped in a membranous, transparent sack emerged from the ventral pedal gland and was released, (figure 2, li). The lenticular walls of the egg case were whereupon the stalk straightened and the egg case was slightly concave, creating a constriction along the mid- supported well above the surface of the sand. line of the case that displaced the eggs to either side.

The spatulate capsules (figure 2; table 1) consisted of Spiral cleavage commenced within 12 hours of ovi- lenticular egg cases supported on long, narrow (250 fim position, and the 8 cell stage (figure 5) was reached by diameter) stalks. Egg capsules were roughly rectangular the second day. Thereafter, the embryos became in- in transverse section, with narrow keels running along creasingly irregular in form (figures 6-8). All the embryos the outer edge of each corner. A pre-formed hatching within an egg case underwent de\elopment; there were aperture, containing a membranous plug (figure 2, mp), no nurse eggs or unfertilized eggs in the cases examined. was situated at the uppermost end of each egg case. By 10 da\s after deposition, the stomodeal in\agination Capsules were affixed to the bottom ot the arjuarium by was e\ident (figure 9). .\fter 12 da\s, the protoconch and holdfasts that spread from the base of the stalk. operculum, both of conchiolin, were clearly discernable Each egg case contained an estimated 4,000-5,000 (figures 10, 11), and torsion was complete after 14 days spherical eggs (figures 3, 4; diameter = 164.5 ± 6.7 ixm, (figures 12-14). The velar lobes were formed and in- N = 10), suspended in a clear, gelatinous matrix, all of creasing in size by the 16th day (figures 15, 16). On the

Figures 3-14. Embryonic and larval development of Cancellaria cooperi Gabb at 15 °C. 3. Living etnbr\os, 1 day old 4. One-

(l.i\-(]ld embryo, critical-point dried. Scale bar = 25 ^m. ,5. lJ\iiig embrxos, 2 days old. 6. Living embr\o, 4 days old. 7. Four- da\-old embryo, critical-point dried Scale bar = .50 fim. 8. Living embr\os, 8 days old. 9. Ten-dav-old embr\o. critical-point dried. Scale bar = 50 /nm 10. Li\ing embryos, 12 days old 1 L Hight side and ventral views of same 12-da\-old embr\o, critical-

point dried. Scale bar = 50 ^m. 12. Living embryos, 14 days old \'.i. 14. X'cntral (13) and dorsal (14) views of 14-da\-old embrsos, critical-point dried. Scale bar = 50 ^m asr, apical sensory region; dml, dorsal maiille lip; Ik, larval kidney; o, operculum; pb, polar body; pc, protoconch; pt, prototroch; s, stomodeum. J. R. Pawlik et ai, 1988 Page 49 Page 50 THE NAUTILUS, Vol. 102, No. 2

Figures l,'j-22. Embryonic and lar\al development of Cancellaria cooperi Gabb at 15 °C. 15. Living embryo, 16 days old 16. Sixteen-da\-old t'nibr\o, critical-point dried. Scale bar = .50 ;um. 17. Li\ing. newK -hatched larva with carmine particles in the gut. 18. Shell ultrastrticture of newK-hatched larva, plane of fracture parallel to growing edge. Scale bar = 2 nm 19. .^pertural, dorsal and apical views of shells of newly-hatched larvae. Scale bar = 100 ^in. 20. Living larva, 30 days after hatching 21. Shell and operculum of larva, 30 days after hatching. Scale bar = 250 fim. 22. .-Vpical portion of adult shell (LACM 40-95.1), arrow- indicating transition from protoconch to teleoconch, scale bar = 500 fim.

mo, month; o, ()j)crculuni; po, posterior ciliary band; pr. preoral ciliar\ band; sto, storiiach; vl, velar lobe.

20tli day after capsule depobitiou, eiiibr\ os began to move da\s at 13 °(.' the nicnibrancjus plug occluding the hatch- more freely within the egg case. The gelatinous matrix ing aperture dissolved, and swimming larvae escaped. in which the developing embrvds had been suspended Shells ol the new K hatched veligers (figures 18, 19) had became less viscous, and the swimming enil)ryos aggre- a mean diameter of 304,6 /um (N = 10, SD = 3.5), and gated randondy w illiin the case. ,\ftor an average of 27 were 3.9 ^m thick. The veligers possessed an operculum a/., Page 51 J. R. Pawlik et 1988

Table 1 . Measurements of egg capsules of Cancellaria cooperi Gabb. All measurements in mm (N = 10)

Character Mean Range SU

Total length Page 52 THE NAUTILUS, Vol. 102, No. 2

Table 3. Protoconch measurements for cancellariid species for which mode of development is not known Data are presented in the format of mean/standard deviation. D = diameter in mm; \'(il = number of w horls or volutions, measured according to Jablonski and Lutz (1980:332).

Species 1) V„l D \ ol

Cancellaria reticulata: L'SNM 619108 (N = 10) 0.99/0.11 C-ancellaria spengleriana. USNM 664965; USNM .344417 (N = 6) Trigonnstoma scalare: USNM 846304 (N = 1) Olssonella smithiU USNM 806986; USNM 450577; USNM 667720 (N = 5) Sarona mitraeformis: Petit collection (N = 3) Scalptia ohiiquala: L'SNM 629063 (N = 4) Cancellaria similis. USNM 664967 (N = 5) et Page 53 J. R. Pawlik al, 1988

In: Rhoads, D. C. and R. A. Lutz (eds.). Skeletal growth Paige, J. A. 1986. The laboratory culture of two aplysiids, of aquatic organisms: biological records of environmental Aplysia brasiliana Rang, 1828, and Bursatella leachii plei change. Plenum Press, New York, p 323-377 (Rang, 1828) (Gastropoda: Opisthobranchia) in artificial Jabloiiski, D. and R. A, Lutz. 1983. Larval ecolog\ ot marine sea water. The Veliger 29(l):64-69.

benthic invertebrates: paleobiological implications. Bio- Radwin, G E. and J. L. Chamberlin. 1973. Patterns of larval logical Reviews 58:21-89. development in stenoglossan gastropods. Transactions of Kiiburn, R and E Rippey. 1982. Sea shells of southern Africa the San Diego Society of Natural History 17(9):107-117. McMillan South Africa, Ltd., Johannesburg. 249 p Shuto, T. 1974. Larval ecology of prosobranch gastropods and

Knudsen, J. 1950. Egg capsules and development of some its bearing on biogeography and paleontology. Lethaia 7: marine prosobranchs from tropical west Africa. Atlantide 239-256. Report 1:85-130. Thorson, G. 1935. Studies on the egg-capsules and develop- MacGinitie, G. E. 1955. Distribution and ecology of the ma- ment of Arctic marine prosobranchs. Meddelelser om rine invertebrates of Point Barrow. Alaska. Smithsonian Gronland 100(5):1-71, Miscellaneous Collections 128(9):1-201. Thorson, G, 1944. The zoology of east Greenland: marine Morch. O. \ L. 1869. Catalogue des Mollusques du Spitz- gastropoda prosobranchiata. Meddelelser om Gronland bergen recuillis par H. Kroyer. Memoire de la Societe 121(13):1-181.

malacologique de la Belgique, Tome 4, Bru.xelles. Thorson, G. 1950. Reproductive and larval ecology of marine

O'Sullivan, J. B., R. R. McConnaughey. and M. E. Huber. 1987. bottom invertebrates. Biological Reviews 25:1-45. A blood-sucking snail: the Cooper's nutmeg, Cancellaria cooperi Gabb, parasitizes the California electric ray. Tor- pedo californica .\yres. Biological Bulletin 172(3):362-366. THE NAUTILUS 102(2):54-55, 1988 Page 54

Latirus martini (Gastropoda: Fasciolariidae), a New Species from Honduras

Martin Avery Snyder

7-4.5 Newtown Road \'illanova, PA 19085

ABSTRACT squarish, greater in length than siphonal canal; anal canal weakly developed; axial sculpture of 13-16 prominent Latirus martini new species is described from shallow water ribs, crossed by numerous, pronounced spiral cords with off KoalaM Island. approximateK 40 miles north ot the Hon- grooves between; cords alternateK larger and smaller, duras mainland. This ne« ta.xoii differs from its nearest Carib- first t\\ o on body whorl about ecjual in thickness, forming bean relative, Latirus angulatus (Roding, 1798), by its sculpture, coloration, shape, and size. The 13-16 axial ribs on the ridge and deep suture; approximateK 45 cortls on body body whorl constitute roughly twice the number found in other whorl running onto anterior end of siphonal canal, 25 Caribbean members of this genus cords on the penultimate whorl, with every fifth cord somew hat stronger than others; spiral cords darkish orange-brow n turning to dark brow n between axial ribs; INTRODUCTION grooves between cords light caramel brown w ith yellow cast; shell appears dark orange-brown, with lighter axial

In the summer of 1985 10 specimens of a new Latirus ribs; columella with 3-4 distinct plicae (figure 1); teeth were taken under coral rubble b\ divers off the north translucent, shiny, white; outer lip crenulated, with in- coast of Roatan Island, Honduras. These shells were ob- dentations corresponding to spiral sculpture; smooth por- tained by Mr. Thomas Honker of Florida, who kindly tion of inner lip extending approximately to opposing passed them on to the author. The specimens were all side of the first axial rib, followed anteriorly by 14-16 collected alive, the soft parts discarded, and the opercula irregular w hite lirae on inside of bod\ w horl; operculum glued back in place on cotton. For this reason no soft \ellow -brow II in color, chitinous, nearK filling aperture. parts were examined, although the opercidum is de- Type locality: 15-20 feet, under coral rubble, north scribed. Since that time, some additional specimens have coast, Roatan Island, Honduras, summer, 1985. been taken from the same locality.

Specimens of the new species are deposited in the Type material: Holotv pe, ANSP 361064 (27.5 mm); collections of the Academx of Natural Sciences of Phila- paratype 1, DMNH 169442 (23.8 mm); paratype2, USNM delphia (ANSP), the Delaware Museum of Natural His- 859070 (26.9 mm); paratypes 3-5 in the author's collec- the National of Natural tory (DMNH), and Museum tion (34.0 mm. 24.8 mm, 23.6 mm). .\11 paratv pes from Institution .additional History, Smithsonian (USNM). the t\ pe locality. specimens are retained in the collection of the author. Etymology: The species is named for the author's eldest son. Family Fasciolariidae Gray, 1853 Subfamily Perislerniinae Trvon, 1880 Discussion: This species appears to be most closeK related to the highly variable Latirus angulatus (Roding, Genus Latirus Montfort. 1810 1798). Several forms of the latter were illustrated by Type species: Latirus auraiitiacus Montfort, 1810, In Bullock (19(58, 1974). Specimens of /.. angulatus from Islands, 120 miles north ot Hon- nionut) p> [= L. gibbulus (Gmelin, 1791 j]. the Swan approximateK duras (Bullock, 1968: fig. 5, plate 2), and a specimen from the northern coast of South .\merica (Bullock, 1974: Latirus martini new species fig. 21) are superficialK similar to L. martini. The col- (figures 1, 2) oration is similar but the shape and sculpture are com- Description: Shell heavy, small to medium in size (,21.7 pletel) typical of L. angulatus and thus readiK distin- mm to 38.4 mm in length), somewhat squat, with spire guishable from L. martini. Latirus angulatus has 7-9

' iiearK s length of shell; profile inflated, w ith 8-9 whorls; axial ribs, about half the iiund)er of /.. martini. The spiral protoconch (figure 2) of 1 '/2 whorl, smooth, bulbous, cords on L. angulatus luc weaker than those of L. mar- translucent pale orange-brown in color; aperture oval to tini. The tvpical coloration is also quite different. In a M. A. Snyder, 1988 Page 55

Generally, L. angulatus is a more elongated shell with the spire usualK- more than half the length of the shell. Possilile confusion could also occur with Leucozonia

nas.sa (Gnieliii, 1791 ) which is somewhat similar in pro-

tile and coloration. This shell lacks the strong a.xial sculpture of L. martini, and has a thick, black-brown oper-

culum. A characteristic narrow \\ hite band at the base, commonly terminating in a small spine on the outer lip, serves to distinguish this species from L. martini. Finally, confusion might arise with the recently-described species Latirus vermeiji (Petuch, 1986). Latirus

vermeiji has less pronounced surface sculpture, is dis- tinctlv orangish in color, and has a caramel-orange col-

ored inner lip, whereas the inner lip in L. martini is yellowish brown. The roughly even spiral cords in L. vermeiji are more pronounced with every other cord being white in color.

LITERATURE CITED

Bullock, R. C. 1968. The fasciolariid genera Latirus, Doli- cholatirus. and Teralatinis (Mollusca: Gastropoda) in the

Western .Atlantic. Master s thesis, Universit\ of Maine,

Orono, 109 p., 8 pis. Bullock, R. C. 1974. A contribution of the systematics of some West Indian Latirus (Gastropoda, Fasciolariidae). The

Nautilus 88(3):69-79, 26 figs. Gmelin,J. 1791. Systema naturae, 13th ed. Part 6:3021-4120. Montfort, P. D. de. 1810, Conchyliologie systematique et

Figures 1. 2. Lalinis martini new species. 1. Holotype, ANSP classification methodique des Coquilles ... 2. Paris, 676 p. 361064. 2. EarK « horls of parat\ pe, DMNH 169442, both from Petuch, Edward J. 1986. New South American gastropods in north coast of Roatan Island, Honduras, under coral rubble in the genera Conus (Conidae) and Latirus (Fasciolariidae).

5-7 m. Proceedings of the Biological Society of Washington 99( 1 ): 8-14. Roding, P. 1798, Museum Boltenianum, Hamburg, 199 p. dark brown specimen of L. angulatits, the raised cords Tryon, G. W. 1881. Manual of conchology 3. Tryon, Phila- and that portion of the whorl near the suture are dark delphia, 310 p., 87 pis. brown, in areas between cords where there is fine or- namentation, the shell is light cocoa-tan to orange-brown. THE NAUTILUS 102(2):56-64, 1988 Page 56

Shell Variation of Springsnail Populations in the Cuatro Cienegas Basin, Mexico: Preliminary Analysis of Limnocrene Fauna

Koberl Hershler Lee-Ann C. Ilayek

Department of Invertebrate Zoology Mathematics and Statistics National Museum of Natural History National Museum of Natural History Smithsonian Institution Smithsonian Institution Washiniitiin, DC 20560, USA Washington, DC 20560, USA

ABSTRACT ulations, the analysis of w hich is considered crucial to understanding the speciation process (Gould & Johnston, Geographic variation in shell morphometry is analyzed for 1972; Endler, 1977). To date, researcii on this snail fauna three locally endemic springsnail (Gastropoda: Hydrobiidae) has largeK been taxonomic (Ta\ lor, 1966; Hershler, 1985), species occurring sympatrically in nine limnocrenesof the Cua- although geographic variation of one endemic species tro Cienegas Basin, Coahuila, Mexico. Despite some correlation was partly analyzed (Hershler, 1985; Hershler & Minck- of size-related variables across species, groupings of populations ley, 1986). Taylor (1966) and Taylor and Minckley (1966) based on multivariate analyses were not very similar among noted apparent diversity in extent of differentiation species, nor were they strongly concordant with current drainage configurations in the basin. Groups of populations of Mexi- among local species: Mexipyrgus Taylor, an endemic pyrgu.s churinceanus Ta\ lor having different patterns of shell restricted to large (> 25 m-) springpools (limnocrenes) sculpture and color banding (and once considered separate and stream outflows, is variable enough to have been species) were not separated similarK on basis of shell size and origiiialK considered as six nominal species (Ta\lor, 1966; shape. Inter-population differentiation of these snails was ap- synonymized to monotypy by Hershler, 1985), whereas equivalent to that of Symphuphilus minckleyi proximately other snails appear morphologically uniform, at least in Taylor, whereas Mexithaunia quadripaludium Taylor was less the portion oi the basin that has been well studied (i.e.. variable all but the southeastern lobe; Hershler, 1985). It was suggested that heightened differentiation of Mexipyrgus

resulted from marked discontinuity of its habitat: INTROULCniON

. . it seems that habitat of this genus is more likely to One of the more remarkable aciuatic faunas of the New be discontinuous than that of other snails in the area. World occurs in the small (1,200 km-), intermontane Mexipyrgus lives in soft flocculent ooze or mud in the valley of Cuatro Cienegas, Coahuila, Mexico, which har- lagunas, thus not in the shallows where wave action re- moves the fine particles Extensive marsh\ areas with small bors at least 26 locall) endemic forms (Cole, 1984; McCoy, streams connecting larger water bodies provide no suitable 1984; Minckley, 1984; Hershler, 1985). Local aquatic widespread habitat. (Taylor, 1966:188) taxa show a great diversity in extent of differentiation relative to adjacent biota, ranging from slightK differ- Variation within Mexipyrgus has largeK- been dis- entiated pojMilations to highly divergent genera, sug- cussed in terms of shell sculpture and color banding gesting both long-term persistence of aquatic habitat and (Taylor, 1966), characters absent from or poorl) devel- multiple invasions of the valley over a broad time scale oped in other local snails, and there has been no attempt (Minckley, 1969). .\quatic organisms are deployed among to contrast intraspecific variation among members of the diverse, spring-fed acjuatic habitats that comprise h\'e to snail launa using a set of common characters, such as

seven local drainage systems (Minckle\ , 1969; LaBounty, shell morphometric variables. In this paper we provide 1974), providing what has been termed a "matchless such a comparison between Mexipyrgus churinceanus natural laborator> " (Taylor & Minckley 1966:22) for eco- Taylor and the two other species (both local endemics) logical and cvolutionars- study. common in basin limnocrenes. Mexithaunia quadripa- Spriiigsnails (C»astropoda: Hydrobiidae) of the basin ludium Ta\ lor and S'yniphophilus ininckU'yi Ta\ lor (see are diverse [nine genera (five endemic), 13 species (nine figure 2). Specifically, we seek to answer the following endemic); Hershler, 1985], and occur abundantly in a questions: 1) Do these distantly related snails (Hershler, large number of easiK' accessible sites, providing an ex- 1985) show commonalitN of pattern of geographic vari- cellent (>pportunit\ to study geographic variation of pop- ation? 2) Does shell morphometric variation among pop- R. Hershler and L.-A. C. Havek, 1988 Page 57

Figure 2. Scanning electron micrographs (printed at same enlargement) of cleaned shells ol springsnails from Laguna Tio

Candido, Cuatro Cienegas, Mexico: a, Mexipyrgus churinceanus (shell height, 6.34 mm); b, Nymphophilus minckleyi (6.45 mm); c, Mexithauma quadripaludium (6.22 mm).

For each species, about 15 live-collected, fully mature adults, recognizable by their complete and thickened inner shell lips, were randomly selected from collections from each site and dried for morphometric analysis. Af- ter whorls were counted (WH), shells were imbedded in clay in standard apertural aspect (figure 2) and shell outlines were draw n using a camera lucida mounted on

a WILD M-5 dissecting microscope (12 x or 25 x ). Points on these drawings were digitized and values for the fol-

Figure 1. Map of the central portion of the Cuatro Cienegas lowing "standard" shell parameters (1-4) and shell shape Basin, show ing major drainage features and collecting localities descriptors (5-8) (from Raup, 1966, and elsewhere) were

(numbered as in table 1 ), The inset (lower right) shows locations generated: of the state of Coahuila (dashed line) and Cuatro Cienegas (dot) within Mexico. 1) Shell height (SH) 2) Shell width (S\V) 3) Length of bod\ whorl (LEW) 4) Width of body whorl (WBW) ulations of Mcxipyrgus churinceanus exceed that of the 5) Translation rate (T) tv\o other limnocrene species (above) that occur s\ m- 6) Whorl expansion rate (W) patrically with this species? 3) Are Mexipyrgus popu- 7) Distance of generating curve from coiling axis (D) lations that are well differentiated in terms of shell sculp- 8) Aperture shape (SA) ture and color banding patterns similarl\ separated by shell size and shape variation? A calculated variable (S), consisting of the addition of SH and SW, was generated to serve as a more realistic MATERIALS AND METHODS measure of size than either shell length or width. Cal- culation of shape parameters largely followed methods

Nine limnocrenes were sampled (shown in figure 1 and of Kohn and Riggs (1975), with the exception being W, described in the Appendix), representing the majority of which was measured as the mean of a series of squared springpools in the study area (all of basin excluding south- ratios of perpendicular distances from coiling axis to eastern lobe) where all three forms are common. Only sutures (shell in apertural and not apical aspect) at half isolated sources (Sites 1-4, 9) or, in cases where springs whorl intervals. The apertural suture was not used, due were connected by stream, upflow pools (Sites 5-8) were to frequent loosening of coiling during last half whorl of considered, in order to reduce possible effects of gene growth, nor were sutures used from eroded adapical sec- flow from contiguous populations (Hershler & Minckley, tions of the spire. Digitizing was done using the CONCH 1986). These springs belong to four separate drainage software program (Chapman et ai, 1988; methodology systems of the basin and harbor forms of Mexipyrgus fulK described therein) and a GTCO Micro-Digi Pad referable to four nominal species (see belowj. 12x12 linked to a KAYPRO 2000 microcomputer. Mexipyrgus churinceanus was collected by sieving soft Descriptive statistics for all morphological variables substrates, while the other two species, associated with were obtained for each species and locality. The hy- hard substrates, were gathered by w ashing travertine and potheses of homogeneity of mean differences and vari- macrophytes in a bucket. Material was fixed in dilute ances across localities were tested for each species. An formalin, and preserved in 70!^ ethanol. ANO\"A model was selected for each variable of each Page 58 THE NAUTILUS, Vol. 102, No. 2

Table I. Descriptive statistics for each species at each locahty. Data given are mean, standard deviation, and sample size (in parentheses). L = locality; variable abbreviations are civen in te\t

\ ariable R. Hershler and L.-A. C. Havek, 1988 Page 59

Tabic I. Oititiimcd

Variable Page 60 THE NAUTILUS, Vol. 102, No. 2

Mexipyrgus c h u rincean us 12 -

10 E E

6 -

_i 1 I ; L. R. Hershler and L.-A. C. Havek, 1988 Page 61

Table 3. Results of Principal Compoiit'iits .\iial\scs on cacli species. Only factor coefficients having weights > 0,25 are listed. I = Mexipyrgus churinceanus, 11 = Mcxilhauina quadripalndium. Ill = Nijmphophihts inincklcyi, Kig. = eigenvalue; % V. = % of variance explained. Page 62 THE NAUTILUS, Vol. 102, No. 2

Despite these correlations, concordance of pattern

among plots of PC scores for the three species is not

impressive, although there is similarit) in order along size-related PCI (figure 4). Note that the tightest clustered centroids for Mexipyrgus churinceanus (Localities

1, 3, 4) are vvideK separated for the other two species.

SiniilarK . di\ t-rgeiit "outK iiig centroids lor gi\en sf>ecies

{Mcxithaiinia quaclripaludium, I, S. minckleyi, 3) are not so differentiated in the other species. As mentioned abo\e, the populations of Mexipyrgus considered in this stud\ are referable to four nominal species (fide Ta\ lor. 196(-)) on basis of shell sculpture and

color banding; Mexipyrgus churinceanus (Localities 1,

2, 4); Mexipyrgus rnojarralis (5); Mexipyrgus lugoi (6- S); and Mexipyrgus carranzae (9) (the form present at

Localit) 3 is distiiicti\e and not clearK referable to any nominal species). These nominal species are poorly seg- 1 J regated on the PC axes: extent of separation of rnojarralis Mexipyrgus churiiiceanus (5) and carranzae (9) from other centroids, for instance,

is exceeded by that seen among three populations referable to churinceanus in Los Remojos spring complex (6-

8).

DISCUSSION

For all three species, most inter-population variation in- volved shell size and size-correlated variables. Common- ality of geographic variation patterns was indicated by significant correlation of population means tor some of these variables across two of three species pairs. The extent of this commonality was not, however, impressive when groupings of populations based on multivariate anal) ses were examined. A strong correlation between geographic variation patterns and current drainage configurations was not apparent for any of the species, suggesting that in this example development of intraspecific di\ersit\ of shell inorphometrv ma\ be related to ecological as well as Mexithauma quadripaludium historical factors (see Chernoff, 19S2, for general discussion of this subject), although we acknowledge the possibilitv that the poor correlation w ith current drainage configuration ma\ be obscured b\ historicalK complex basin hydrograpln (Miiickle\, 19(i9; Hershler \ Minck- ley, 1985). Springpools concentrated around the northern (5)@ tip of Sierra de San Marcos (where the study area is located) are higliK unilorm in water quality (Miiickley

in Cole, 1968): single measurements taken b\ us during the study indicated, for instance, that temperature and conductivity ranged among the nine springpools from 3 25.5-34.5 °C (seven localities diftering b\- < 4 °C) and ® 1,825-3,500 micromhos cm (seven localities differing by < 430 micromhos, cm), respectively. Pools do dilter con-

Figure 4. 'rliree-dimeii.sioiial plots ot PC cenlroids for the

three species t\ axis. PCI; Y, PC2; Z. PC:.'3). standardized and Nymphophilus minckleyi viewed in perspective. Sizes of balls indicating centroids are scaled to heighten perspective. R. Hershler and L.-A. C. Hayek, 1988 Page 63

Table 5. Results of four-group Discriminant Analyses on drainage system for each species. Standardized function coefficients (>

0.25) and pooled within group correlation coefficients ( > 0.25) of each function with each original variable are listed. I = Mexipyrgus churinceanus: II = Mexithauma quadripaludiutn; III = \'ymphophilus minckleiji; Eig. = eigenvalue; % V. = % of variance explained; ('.C. = canonical correlation Page 64 THE NAUTILUS, Vol. 102, No. 2

Hershler, R. 1984. The h\(lrobiid snails (Gastropoda: Risso- drainage of basin now feeding canals. USNM 857909, acea) of the (^uatro C^ienegas Hasin: systematic relation- 857918, 857927. ships and ecology of a unique fauna. Journal of the .Ari- Sites 2-4 represent three springs in the area known as zona-Nevada .-Keademy of Science 19:61-76 El Garabatal, located north of Poso de la Becerra and Hershler, R. 1985. Systematic revision of the Hydrobiidae

east of Becerra s outflow . El Garabatal drains to the north (Gastropoda: Rissoacea) of the Cuatro Cienegas Basin. west, Coahuila, Mexico. Malacologia 26:31-123. and and may be considered either a separate, small Hershler, R. and VV. L. Minckley. 1986. Microgeographic drainage or a subset of the Rio Mesquites system, the variation in the banded spring snail (Hydrobiidae: Mexi- largest drainage in the basin. pyrgits) from the (;uatro Caenegas Basin. (;()ahuila. Mex- 2. Laguiias de Juan .Santos (5-IX-86), 12.8 km W-SW ico. Malacologia 27:357-374. ot Ciuatro Cienegas. Largest of El Garabatal springs con- Hershler, R and U VV. Sada. 1987. Springsnails (.Gastropoda: sisting of series of relatively shallow (depths to 2.0 m), Hydrobiidae) of Ash Meadows, .•\margosa Basin, Califor- inter-connected lagunas fed by springs emerging along nia-Nevada. Proceedings of the Biological Society of pool margins. Pool area somewhat larger than that of Washington 100:776-843. Poso de la Becerra. Extensive marshes bordering near- Kohn. A. J. and A. C. Riggs. 1975. Morphometry of the Conus entirety of spring's perimeter. Water lily stands exten- shell. Systematic Zoology 24:346-359 sive. Stream outflow extending several hundred meters LaBounty. J. F. 1974. Materials for the revision of cichlids from northern Mexico and southern Texas. LS.\ (Pcrci- before terminating in shallow marshv area. USNM formes: (^ichlidae). Ph.D. thesis (unpublished), Arizona 857912, 857921, 857930.

State University. Tempe. 120 p. 3. Unnamed small spring, know n to biologists working

McCoy, C. J. 1984. Ecological and zoogeographical relation- in area as North Spring (3-IX-86), 12.0 km W-SW of ships of amphibians and reptiles of the Cuatro Cienegas Cuatro Cienegas. Small springpool ca. 12 x 18 m. about Basin. Journal of the .'Vrizona-Nevada Academy of Science a meter deep, w ith ca. 20^ coverage b\ water lily . Out- 19:49-59 flow entering second pool (not a spring); di.scharge from Minckley, V\' L. 1969. Environments of the bolson of Cuatro latter e.vtending 50 m before disappearing into hole. Cienega.s. Coahuila, Mexico L'niversity of Texas at El Paso. USNM 857910, 857919, 857928. Science Series Number 2, p. 1-65. 4. Unnamed large spring (.5-IX-86), 11.8 W-SW Minckley, W. L. 1984. Cuatro Cienegas fishes: research re- km ot Cuatro Cienegas. Roughly circular springpool, ca. \ iew and a local test of diversity versus habitat size. Journal 70 of the Arizona-Nevada Academy of Science 19:1.3-21. m across; depths not exceeding 3.5 m. Water lily dense Minckley, VV. L. and G. A. Cole. 1968. Preliminary limnologic in center of pool. Outflow feeding small marsh. USNM information on waters of the Cuatro Cienegas Basin, Coa- 857916, 857925, 857934. huila. Mexico. Southwestern Naturalist 13:421-431 5. "West" Laguna in El Mojarral (4-I\-86). 9.0 km Haup. D. M 1966. Geometric analx sis ot shell coiling. Journal SW of Cuatro Cienegas. Moderate-sized spring (26 x 59 of Paleontology 40:1178-1190.' ni; Arnold, 1972:12), with depths to 4.5 m. Orifices few Taylor. D. VV. 1966. A remarkable snail fauna Irom Coahuila, in number, cavernous. Water lily stands few, relatively Mexico. Veliger 9:152-228. thin. Water exiting spring \ia both a shallow, surficial Taylor, D. VV. and W. L. Minckley. 1966. Neu world for stream and single, tubular, subsurface \ent. Stream out- biologists. Pacific Discovery 19:18-22. flow entering Wilkinson, L. 1986. SYSTAT:' the system for statistics. SYS- "East Laguna, which in turn drains into T.-VT, Inc., Evanston, IL. Rio Mesquites. USNM 857908, 857917, 857926. Sites 6-8 are inter-coiuiected springpools known locally as Los Remojos. 9.0 km S-SW ot Cuatro Cienegas. APPENDIX Site 6 drains into a large pool recei\ ing discharge from a second pool fed by outflows from Sites 7 and 8. System Springpools sampled are numbered as in figure 1. Lo- draining into Rio Mesquites. cality names are either those of Minckley (1969) or reflect 6. Northernmost of Los Remojos springs (5-IX-86). local usage. Locality data represent air distances troni Pool ca. IS X 32 m. Depths generalK > 1.5 m; water Cuatro Cienegas. Dates of collection are given in paren- lily common. USNM 857913, 857922,' 857931. theses. Catalog numbers (USNM) for voucher material 7. Intermediate Los Remojos spring (5-IX-86). Pool (dry shell plus alcohol specimens) from each locality are ca. 28 X 47 m. shallow (< 1.0 m). Water lib uncommon. given in follow ing order: Symphophihis ininrklciji Tay- USNM S57914. 857923. 857932, lor, Mcxipyrgiis cliiirinrcaiuis Ta\ lor, ami Mcxithauiua 8. Southeriunost of Los Remojos springs (5-IX-86). Pool quadripaluilium Taylor. moderately large (22 x 42 m), with depths increasing

1. Poso de la Becerra [(south pool) 3-1X-86], 13.7 km in southern end of pool to 4.0 m. Water lib common. W-SW of C^uatro C^ienegas. A once enormous spring area USNM 857915, 857924, 857933. (over a kilometer long) signilicantK reduced in size hy 9. Laguna Tio (-andido (5-IX-86), 12.5 km S-SW of canal development in early 19605 (see Taylor, 1966:162; C.uatro Cienegas. Large spring (ca. 45 x 100 m) exten- Minckley 1969: figs. 15, 16). Currently consisting of two sively vegetated by water lily and other macrophytes. large springpools (each ca. 50 x 125 m-) connected by Depths generally from 2.0-4.0 m. Outflow extending short section ot stream. Spring orifices few and large, eastward, comprising part of a major system positioned occurring in tleepest (to 7 ni) portion of pools. Water liK south of Rio Mesquites drainage. USNM 85791 1, 857920, stands extcnsi\c Spring and outflow i-onstituting a major 857929. THE NAUTILUS 102(2):65-72, 1988 Page 65

Niche Congruency of Freshwater Gastropods in Central North America with Respect to Six Water Chemistry Parameters

Eva Pip Department of Biology University of Winnipeg Winnipeg, Manitoba R.3B 2E9 Canada

ABSTRACT The objective of the present study was to examine niche relationships for the freshwater gastropods in cen- Occurrences of freshwater gastropods were studied at 430 sites tral North America in six water chemistry dimensions: in central North America with respect to total alkahnity, total total alkalinity, total dissolved solids, chloride, sulphate, dissolved solids, chloride, sulphate, phosphorus, and dissolved reactive phosphorus, dissolved organic matter. Mean niche positions for the gastropods were moly bdenum and organic calculated for the six combined water chemistr\ parameters matter (DOM). These factors were chosen because a pre- by comparing mean values for each species using agglomerative vious study (Pip, 1987) showed that these variables con- hierarchical cluster analysis. Total niche relations were exam- tribute towards variation in species richness of freshwater ined by calculatmg and summing the amount of overlap in the gastropod communities within the study area. In the observed ecological tolerance ranges for the six parameters for present study a niche was defined as the range between each species pair, and appKing cluster and principal compo- the minimum and maximum values observed for each nent anaKses. The results showed that the species occupied a species in the study area, and was thus analogous to the broad spectrum of niches, with progressively increasing eco- concept of the "realized niche" discussed bv Hutchinson logical range. Gastropods with low mean inorganic values and (1957). narrow niche widths were interpreted as specialists for waters with low concentrations of dissolved inorganic materials. Species which clustered together in terms of central tendencies of occurrence often differed from each other in terms of overall MATERIALS AND METHODS niche similarity, and vice versa. Competition may have been more important in waters with low inorganic concentrations. A total of 430 sites was examined in central North Amer- ica (47°-54°N and 94°-106°W) during the May-Septem- ber, 1972-85 seasons. All sites contained water year-round. INTRODUCTION Ponds (< 10 ha) comprised 41.0% of the sites sampled, lakes (> 10 ha) 41.5%, rivers (> 2 m deep) 10.3%, and A number of workers have demonstrated the importance creeks (< 2 m deep) 7.2%. Each site was examined for of water chemistry in the distribution of freshwater gas- snails by wading or canoeing; snails on macrophytes were tropods {e.g.. Boycott, 1936; Hubendick, 1947; Aho, 1966; obtained by dredging with a rake or by using SCUBA.

Harman & Berg, 1971; Dussart, 1976; Pip, 1978, 1985, Search time at each site was limited to 1 hr. 1986; Okland, 1979; Okland & Okland, 1980). However, Surface water samples were collected at most sites, but aside from the pioneering study of Finnish lakes by Aho at depths of >3 m where macrophytes did not reach the et al. (1981). relatively little is known regarding niche surface a van Dorn sainpler was used. The samples were relationships between freshwater gastropod species that placed on ice in darkness and frozen within a maximum coexist within a given geographical area. of 48 hr. Samples were analyzed using methods rec- The ecological niche of a species can be described as ommended b\ the American Public Health Association an infinite h) perspace representing ail physical and biot- (1971). ic aspects of the environment in which the species occurs. While most of the sites were sampled only once, ap- Nevertheless, practical considerations restrict most niche proximately 50 of the locations were resampled at dif- studies to a few selected parameters that are thought to ferent times of the growing season and in different \ ears. be important for the species (Levins, 1968 in Aho et al., For such sites, extreme low and high water chemistry 1981). .According to the latter v\orker, the niche of a values were used for statistical anaKses. Because of en- species with respect to a given parameter can be ap- vironmental heterogeneity , large lakes were sampled at proximated by observing the species distribution over a different locations which were treated as separate sites. range of values of the parameter in a number of envi- Mean and niche comparisons were made using ag- ronments. glomerative hierarchical cluster analysis (Sneath & Sokal, Page 66 THE NAUTILUS, Vol. 102, No. 2

1973). Ward's method, a minimum variance technique and principal component analysis were obtained from (Wishart, 1969) was used for clustering. Cluster fusion SPSS. Inc.. (Chicago. Illinois. distances were calculated by means oi the squared Eu- clidean distance measure: RESULTS

Distance,, = S, (Xj — y,)- The ranges of concentrations of the inorganic chemical

Because Ward's method is an intense!} clustering pro- parameters encompassed at the stud\ sites have been cedure, ranking of the species in the dendrograms was presented elsewhere (Pip, 1986). The inorganic param- checked for misclassifications by comparing with the eters showed a number of significant (P < 0.05) positive ranking obtained using the unweighted pair-group meth- intercorrelations at the stud\ sites (table 1) and could od (Cliltortl k Stephenson, 1975), a weakly clustering therefore be regarded as a block with main similar trends. strateg) (Sneath & Sokal, 1973). The two methods showed However DOVl was significantK correlated onK with similar relative ranking; thus mi.sclassifications were not moK bdenum reactive phosphorus and total dissoK ed sol- apparent. Since the relative importance of the six pa- ids. rameters differed with species and situation (Pip, 1987), Five or more sets of w ater chemistr\ data were avail- ail parameters were weighted etiualK in the cluster anal- able for 36 species in the present stud\ (table 2); only yses. these species were included in cluster and principal com- In construction of the dendrogram for mean niche ponent analyses. The bulk of the mean, maximum, and position (tigure 1), mean values for each of the six pa- minimum values on w hicli anahses were based are gi\en rameters were used to construct a data matrix for the by Pip (1986) for all parameters except DOM, which is comparison of taxa with respect to the mean positions of given in table 2. their niches. Because of differences in units and statistical The dendrogram for mean niche positions (figure 1) distributions of the six chemical parameters, values were with respect to the six combined parameters reflected converteil to standardized Z scores, each parameter w ith the lf)cations in the ecological ranges w here each species a mean of and a standard deviation of 1. Ward s method w as most often found. C^lnster 1 consisted of species w ith was applied to the Z scores. the lowest mean values lor total dissolved solids and low

The first step in niche comparison was the construction values for all other inorganic parameters. On the other ot a combined niche congruenc\ matrix for the six chem- hand, Armiger crista, Physa jcnncssi. and Fossaria mo- ical parameters. For each parameter, the amount of o\er- dicella differed the most frcmi all other species; these lap in the observed ecological ranges of each species pair gastropods showed the highest mean values for total dis- was di\ided by the combined range of the two species solved solids, as well as high \alues for all other param- (range between lowest minimum and highest maximum eters (Pip, 1986), including DOM. Other species in the values). Parameter \alues were not standardized. The six spectrum could show low or high \alues of DOM which overlap proportions were then summed for each species were largely unrelated to the concentrations ot inorganic pair (maximum possible value 6.0). The combined coef- parameters; for example Valvata tricarinata, Fossaria ficients formed a niche congruency matrix that was ana- decampi. and Stagnicola catascopium show ed the lowest

i\ zed using cluster anaKsis as above to gi\e the dendro- mean values for DOM ol all species except Campcloma gram in figure 2. decisum (table 2). The niche congruency matrix was also examined using The relations between species in terms ot niche con- principal component analysis (Tatsuoka, 1971), based on gruency for the six parameters are represented by the the error correlation matrix. Components were f)rthog- dendrogram in figure 2. The groups that emerged formed onalK rotated so that the lirsl principal component lay a spectrum ol the ranges of chemical environments in along the direction of greatest variance. which each species was observed. Clusters 1 and 2 con- Statistical programs used for dendrogram construction sisted of species that showed narrow tolerance ranges for

Table 1. Correlations between water chemistrv parameters at the stud\ sites Upper diaRonal = r. lower diaijonal = p. \ = 446- 468. —

E. Pip, 1988 Page 67

E<»

o2o - — C Si?3 o «> 2 S5 - — Q. » e = o = ° ; S ? E o °- ' — JO O o c E c a. o° > -o S •o ° o K !: ^ 2 o Z <« — o O O" C^ O- 3 o£ «)^- w — O U J E o O K 3 c >. ~^ o O o o = ! w CO *rt >»•»—«»-•» © O e c ; « O 0>« c =^ E E « .. jr o 3 o "^ < CL U- < ll- O < I Q. Q. > I I I I I I I I——————r-~T— Page 68 THE NAUTILUS, Vol. 102, No. 2

— •» O fl) > 5 ? * > R — •- il o c o > m f o _1 O X E. Pip, 1988 Page 69

Table 2. Mean, maximum, and minimum values of dissolved the inorganic parameters at low corieeMitratioiis (Pip, organic matter (absorbance of acidified water at 27.5 nm) ob- 1986), The narrow tolerance ranges ot these species con- served for the gastropods in the study area. tributed towards larger interspecific differences, as seen from the larger cluster fusion coefficients; niche overlap with species that had broad ranges was small, while the probability that significant proportions of the niches of any two species w ith restricted ranges would not coincide was increased b) the narrowness of the respective ranges. As a result, relative niche differentiation was better defined in waters with low inorganic concentrations. Species witii broader ecological tolerance ranges, on the other iiand, could occur in waters with high inorganic concentrations, although in most cases they could also tolerate comparatively low values. These species formed the clusters on the upper part of the dendrogram in figure

2, and were more numerous than species which were restricted to a narrow range of environments. The greater similarity between members of each of these clusters derived from the larger degree of overlap that necessarily arose as a result of the broad ranges. Cluster 6, composed of Stagnicola paliistris, Fossaria modicella, and Physa jennessi, represents species that occurred in the broadest range of chemical environments. Niche relations were also examined using principal component anal\ sis applied to the niche congruency error matrix, which extracted two major components that accounted for 55.4% and 22.7% of the variance, respectively. The remaining components each accounted for 5% or less of the remaining variance. The niche relations for individual species are plotted with respect to the first two principal components in figure 3. The distributions that emerged reflected the groupings of figure 2, although clusters 1 and 2 from figure 2 showed close affinities (groups A and B) in figure 3. Species in groups A and B showed narrow niche widths and low mean parameter values. Species in groups E and particularly F showed wide ecological ranges; these groups also included the most common species in the stud\ area (Pip, 1978). The first principal component of the niche congruence- error matrix was significantly correlated w ith mean values for the respective species of all six parameters (total alkalinity r = 0.49, P = 0.001; total dissolved solids, phosphorus, and chloride, all r = 0.47, P = 0.002; sulphate r = 0.36, P = 0.016; DOM r = 0.33, P = 0.023; all N = 36). This large number of correlations derived from intercorrelations of these parameters at the sites examined (table 1). The strong correlations with the first component are interesting, in that the values in the original congruency matrix were not themselves correlated with any of the chemical parameters. The second principal component showed no significant correlations with the mean values of the parameters examined and was apparentl) related to an unmonitored factor.

DISCUSSION

Cluster analysis showed that the gastropods within the study area occupied a broad spectrum ot chemical niches. Page 70 THE NAUTILUS, Vol. 102, No. 2

PC I '55.4%

3 ,i.«''9 33 7 2l\3S\ '24/24" Z. •-.<^ 'if

PC 2- 22.7%

. .- -1,0.——.— E. Pip, 1988 Page 71

separate macrophyte species within the same water bod\ 3 also reflected the geographical distributions and types (Pip & Stewart, 1976). Interspecific differences in diet of habitats frequented by the respective species. Species have also been reported by Reavell (1980). Other types in groups A and B were present largely in Precambrian of habitat partitioning may occur in certain situations, Shield w aters characterized by low total dissolved solids, for example with respect to depth (Lacoursiere ct al., and were absent or rare west of the Shield boundary. 1975), temperature (Boag, 1981), and turbulence (Calow, Species in group C frequented ponds and small lakes 1973). (Pip, 1986) with intermediate water chemistry values. The proportionally greater niche differences between Group D occurred frequently in lakes. Group E species species found at low inorganic concentrations were re- occurred in a very w ide variety of water chemistry types, flected b\ the lack of significant association between the and differed w ith respect to preferences for water body species in clusters 1 and 2 in figure 2 at the same sites type, in some cases lacking discernable preferences en- within the study area [e.xcept for M. decepta and H. tirely. Species in group F occurred in the most extreme campanulatum (Pip, 1978)]. Greater niche differences ranges of water chemistry, but were found most fre- in unproducti\e environments ma\' have been associated quently in ponds (Pip, 1986). with more intense competition (Emlen, 1973) than in While the niche means and widths utilized here are nutrient-rich situations, where niches among many species those observed for the study area as a whole, ecological were similar, and a number of interspecific associations ranges may vary for a given species in different geo- were observed. Eutrophic waters show greater produc- graphical regions (Pip, 198.5), and are likeh different for tivity of algae and macroph\ tes, w hich constitute much individual populations. The importance of particular pa- of the food of freshwater gastropods (Reavell, 1980). rameters ma\ also vary among habitats with different Food has been proposed as an important factor regulating chemical and physical characteristics (Pip, 1987). Thus gastropod abundance and distribution [e.g., Dillon & the exact position of the niche may be a plastic attribute Benfield, 1982). Species richness of gastropod commu- that can vary to some extent with situation as well as nities in the study area is positiveK correlated with tro- time. phic state (Pip, 1987). This correlation derives from the tendencies of many species to occur more often at higher LITERATURE CITED values of phosphorus (Pip, 1978, 1986), despite the observation that of the species can also tolerate low- of the littoral most Aho, J. 1966. Ecological basis of the distribution values of this factor. freshwater molluscs in the vicinity of Tampere, South Fin- Competition has been put forward as an important land. Annales Zoological Fennici 3:287-322. Ranta, Vuorinen. 1981. Species composition factor governing gastropod distribution (e.g., Lassen, Aho, J, E. and J. of freshwater snail communities in lakes of southern and 197.5). Species which are taxonomically and morpholog- western Finland. Annales Zoological Fennici 18:2.33-241. ically similar {i.e., "sibling species") (Aho et al., 1981) American Public Health Association. 1971. Standard methods often were not grouped within the same clusters with for the examination of water and wastewater. American respect to niche similarity (e.g., the species of each Stag- Public Health .Association, New York, 874 p. nicola, Fossaria, Helisorna) (figures 2 and 3), indicating Boag, D. .\. 1981. Differential depth distribution among intrageneric ecological differentiation, w hile other con- freshwater pulmonale snails subjected to cold tempera- geners (Gyraulus circumstriatus and G. deflectus) were tures. Canadian Journal of Zoology 59:733-737. grouped together. Aho et al. (1981) found a similar lack Boycott, C. 1936. The habitats of the freshwater molfuscs in of consistency in the distributions of gastropods in Fin- Britain. Journal of Animal Ecology 5:118-186. P. 1973. Gastropod associations w ithin Malham Tarn, nish lakes and suggested that competition is only one of Calow, Yorkshire. Freshwater Biology 3:521-534. a number of factors that contribute towards distribution. Clifford, H. T. and VV. Stephenson. 1975. An introduction to Conversely, most of the groupings w hich emerged in numerical classification. Academic Press, New York, 229 p. figures 2 and 3 contained a wide representation of dif- Dillon, R. T., Jr. and E. F. Benfield. 1982. Distributions of ferent taxonomic and morphological entities. Accord- pulmonate snails in the New River of Virginia and North similar ingly, while the niches within each group were Carolina, U.S.A.: interaction between alkalinity and stream with respect to the parameters examined, species behav- drainage area. Freshwater Biology 12:179-186. ior ecological requirements were likeK to be differ- of freshwater molluscs in and Dussart, G. B. J. 1976. The ecology ent, decreasing competition. Furthermore, differences north west England in relation to water themistry. Journal with respect to other parameters ma\ also have been of Molluscan Studies 42:181-198. 1973. Ecology: an evolutionarx approach. .\d- important, for example t\ pe of water bod\ and bottom Emlen, J. M. dison- Wesley Co., Reading, \i.\, 493 p. substrate (Pip, 1986), t\pe of vegetation (Pip, 1978), or Harman, W. N. and C. O. 15erg. 1971. The freshwater snails other, unmonitored, chemical and physical variables. of central New York with illustrated ke\s to the genera Principal component analysis yielded a representation and species. Search: Cornell L niversit\ Agricultural Ex- of the species in a hyperspace, of w hich the first two perimental Station 1:1-68. for three-quarters of dimensions accounted more than Hubendick, B. 1947. Die Verbreitungsverhaltnisse der lim- the variance. While the first dimension was clearly re- nischen Gastropoden in Sudschweden. Zoologische Bid- lated to the types of chemical environments in which rucke Uppsaliense 24:419-.559. the species most often occurred, the groupings in figure Hughes, R. N. 1980. Strategies for survival of aquatic organ- Page 72 THE NAUTILUS, Vol. 102, No. 2

isms. In: Barnes, R. S. K. and K. H. Mann (eds.). Funda- southwestern edge of the Precambrian Shield. Canadian mentals of aquatic ecosystems. Blackwell Scientific Pub- Field-Naturalist 99:76-85.

lications, Oxford, p. 162-184. Pip. E. 1986. The ecology of freshwater gastropods in the Hutchinson, G. E. 19.57. Concluding remarks. C^old Spring central Canadian region. Nautilus 100:56-66. Harbor Sym[X)sium. Quantitative Biology 22-41.5-427 Pip. E. 1987. Species richness of freshwater gastropod com- Lacoursiere, E., G N'aiiiancourt, and R. Goiilure 1975. Re- munities in central North America Journal of MoUuscan lation entre les plantes aquatiques et les gasteropodes (Mol- Studies 53:16.3-170.

lusca. Gastropoda) dans la region de la centrale nucleaire Pip, E. and J M Stewart. 1976 The dynamics of two aquatic

Gentilly 1 (Quebec). Canadian Journal of Zoology 53:1868- plant-snail associations. Canadian Journal of Zoolog\ 54: 1874. 1192-1205. Lassen, H. H. 1975. The diversity of freshwater snails in view Reavell, P. E. 1980. A study of the diets of some British of the equilibrium theory of island biogeography. Oeco- freshwater gastropods. Journal of Conchology 30:253-271. logia 19:1-8. Sneath, P. H. A. and R. R. Sokal. 1973. Numerical taxonomy. \V. H. Freeman and Co., San Francisco, Okland, J. 1979. Distribution of en\ ironmental factors and 573 p. fresh-water snails (Gastropoda) in .Norway: use of Euro- Stanley. S. M. 1979. Macroevolution, pattern and process. VV.

pean invertebrate surve\ principles Malacologia 18:211- H. Freeman and Co., San Francisco, 332 p. 222. Tatsuoka. M. M. 1971. Multivariate analysis: techniques for educational and psychological research. \\ ile\ Okland, J. and k. A. Okland. 1980. Acidification threatens John & Sons, p.' trout diet. Research in Norway 1980, p. 21-27. New York, 310 Pip, E. 1978. A survey of the ecolog\ and composition of Wishart. D. 1969. Mode analysis: a generalization of nearest

submerged aquatic snail-plant communities, Canadian neighbour which reduces chaining effects. In: Cole, .\. J. Journal of Zoology 56:2263-2279. (ed). Numerical taxonomy. Academic Press, London, p. Pip, E. 1985. The ecolog\ of freshwater gastropods on the 282-308. THE NAUTILUS 102(2):73-77, 1988 Page 73

Factors Affecting the Distribution of Sphaeriid Bivalves in Britannia Bay of the Ottawa River

B. \^ . Kilgour G. L. Mackie

Departmt'iit of Zoology Universit) of Guelph Guelpli, Ontario NIG 2W1 Canada

ABSTRACT mm mesh screen on top) samples were taken, except at 0.36 m where four samples were taken. A sediment sam- Distribution of Sphaeriidae in Britannia Bay of the Ottawa ple was taken at each depth from which three subsamples River was not homogeneous and was primarily affected by (approximately 100 ml) were taken. for determi- sediment particle size and depth. Total sphaeriid diversity is Two maximal at particle sizes near 0.18 mm. Total sphaeriid density nation of percent organic matter content and one for and abundance of the genus Pisidiurn increase with decreasing geometric mean particle size. water depth. Distribution of species within the genera Pisidiurn From each benthic sample, all sphaeriids and algae and Musculium is variable. These data indicate that compar- (entangled masses of Lyngbya sp. and Vaucheria sp.) isons of pisidiid distribution should consider at least depth, and were hand picked. Sphaeriids were counted and iden- particle size in the sampling design. tified to species. Algae was dried at 70 °C and weighed to the nearest 1.0 mg. Total sphaeriid diversity (Shannon & Weaver, 1949) and density (N/m-), and individual INTRODUCTION species abundances were calculated for each benthic sample. The importance of sphaeriids as part of the aquatic com- The organic content of the sediments was estimated munity has lead to a number of studies dealing with the by ashing two of the sediment samples from each depth distributional patterns of this group (Healey , 1978; Mack- and expressing the result as percent loss on ignition. Geo- ie et ai. 1980). Most studies, liowever, ha\e dealt with metric mean particle size was determined with the third distribution patterns among a group of lakes, rivers, or sediment sample from each depth using the methods sections of a river. Usually it is assumed that variation outlined by Lotspeitch and Everest (1981). among sites is greater than the variation within sites. Stepwise multiple regressions (Ostle & Mansing, 1975) However, data in Avolizi (1976) and Holopainen (1979) were performed to determine which environmental fac- suggest that the abundance of some species can be vari- tors significantly affected total sphaeriid diversity and able within a single site, but little work has been done density, and the abundances of individual genera and to determine the extent of this variation. species. Only those variables found to be independent

Britannia Bay , a section of the Ottawa River just above by simple correlations (table 1), were used coincidentally Ottawa and Hull, Canada, is relatively unpolluted, and in the regression models. is known to contain a diverse sphaeriid fauna ( 15 species), Box plots (McNeil, 1977) determined that all data, including members of Musculium. Pisidiurn, and except diversity values, were most normally distributed Sphaerium (Mackie, 1971). The purpose of this study when the depth means were transformed by logjQ. All was to determine if the depth distribution of sphaeriids simple correlations and stepwise multiple regressions were in Britannia Bay is homogeneous, and if distribution is calculated using the normalized data. a function of the sediment particle size, organic matter content, algal biomass, and depth. RESULTS

MATERIALS AND METHODS Depth means of algal biomass, and ?? organic matter content are presented in figures 2 and 3. Both parameters During mid-.\ugust. 1985. benthos and sediment samples increased with increasing water depth. Trends in sedi- were taken at eight depths (0.25. 0.35, 0.50, 1.0, 2.0, 3.0. ment particle size are given in figure 4. Compared to 4.0, and 6.0 m) along a transect in Britannia Bay of the other depths, sediments were coarser at the 1.0, 2.0. and Ottawa Ri\er (figure 1). 6.0 m depths. At each depth, five Ekman grab (15 x 15 cm, 0.32 Twelve sphaeriid species were found in Britannia Bay Page 74 THE NAUTILUS, Vol. 102, No. 2

laoa-

80D-

<00-

o.so 1.0 2.a «.0 6.0

Depth (in)

Figure 2. Mean algal biomass at each depth along the transect with 95^< confidence limits indicated b\ vertical lines.

(table 3). Density at 0.25 m was less than that at 0.36 and 0.5 m (figure 7). Abundance of the genus Fisidium significantK increased w ith decreasing water depth (table 3). A similar relationship was found for P. variabile. Abundance of P. casertanum significantK increased with finer sediments and lower organic matter content. None of the other seven Pisidium species were correlated with the en\i- ronmental variables. Abundance of the genus Musculiniu did imt correlate with any of the variables tested, .\bundance ot A/, trans- Figure 1. Map of the Ottawa River, and an enlarged map of versum increased significantK with decreasing water Britannia Bay above Ottawa indicating location of tfie transect. Depth contours on enlarged map are in metres. depth (table 3). Abundance of A/, secitris correlated pos- itively with algal biomass. No relationship was found tor the genus Sphaerium with P. casertanum the most abundant species. Mean which was represented b\ onK one species, S. striati- abundances at each depth for each species are presented nuin. in table 2. Diversit) values were highest at the shallow depths (0.2.5-0.50 in) and at .'3.0 and 4.0 m (figure 5). Stepwise multiple regression indicated that diversity increased with decreasing particle size (table 3). Figure 6 indicates a unimodal distribution oi diversity in relation to geometric mean particle size. Maximum diversilN occurred near a particle size of 0.18 mm. Regression analysis indicated that total sphaeriid density significantly increased w ith decreasing water depth at 2-

Table I. C^nrrelatinns between some environmental variables.

Sigiiilitant rorrclaliniis [P < O.O.'j) are indicated 1)\ asterisks (•)

.Mgal Organic 0.25 0.36 0.50 1.0 2.0 3.0 t. 6.0 biomass matter Depth Depth (m) Organic matter 0.787* Figure 3. Mean organic matter content of the sediments at Depth 0.839* 0.863* each depth along the transect with 95^t confidence limits in- Particle size f) Vyl -0,043 322 dicated l>\ \frtital lines. B. W. Kilgour and G. L. Mackie, 1988 Page 75

2.5-

1.5-

0.50 l.D 6.0

Depth (m)

Figure 4. Geometric mean particle size at each depth along Figure 5. Mean diversity values at each depth along the tran- the transect. sect with 95^0 confidence limits indicated bv vertical lines.

DISCUSSION (Rogers, 1976), production (Hamill, 1975), growth and reproduction (Mackie & Qadri, 1978), and may improve In Britannia Bay, total sphaeriid diversity significantK- in- survival in young clams (Gale, 1976). Fine substrates, creases with finer sediments. The finer sediments at 3.0 however, may reduce oxygen a\ailability (Meier-Brook, and 4.0 m, which correspond with high diversity values, 1969; Hartnoll, 1983). The particle size at which diversity contradict the positive correlation which exists between is maximized may represent the optimal grain size for particle size and depth in rivers (Hamill, 1975). The sand efficient burrowing and /or oxygen/ nutrient availability'. bar between the 3.0 and 4.0 m sites (figure 1) may have Total sphaeriid density increases in shallow water in affected the distribution of sediments in the same manner Britannia Bay. From the literature, it appears that re- that P'olke and Ward (1957) describe for a sand bar in source availability and utilization may be affected by the Brazos River. depth. Bacteria and phytoplankton (especially diatoms), The unimodal distribution of diversity in relation to which are food resources for sphaeriids (Holopainen, particle size (figure 6) indicates that diversity is maximal 1985), are generally more abundant in shallow water (0- at a particle size near 0.18 mm, and decreases with finer 2.0 m) than in deep water (> 5.0 m) (Hargrave, 1970). or coarser material. Finer substrates enhance burrowing The drop in total sphaeriid density at 0.25 m which

Table 2. Mean abundance (No. /m-) of individual species at each depth along the transect. Page 76 THE NAUTILUS, Vol. 102, No. 2

2.5- B. W. Kilgour and G. L. Mackie, 1988 Page 77

haviour of the fingernail clam, Sphaeriuni transvcrsum. Mackie, G. L. 1971. Some aspects of the distributional ecology Malacologia 9:121-125. of macrobenthos in an industrialized portion of the Ottawa

Green, R. H. 1971. .\ multivariate statistical approach to the River near Ottawa and Hull, Canada M.Sc. thesis, Uni- Hutchinsonian niche: bivalve molluscs of central Canada. versity of Ottawa, 161 p Ecologv 52:543-556. Mackie, G. L. and S. U. Qadri. 1978. Effects of substratum

Hamili. S. E. 1975. Production of sphaeriid clams and am- on growth and reproduction of Musculium securis (Bi- phipod crustaceans in the Ottawa River near Ottawa-Hull, valvia: Sphaeriidae). The Nautilus 92:135-144.

Canada. M.Sc. thesis. University of Ottawa, 98 p. Mackie, G.L.S. U.Qadri, and R. M. Reed. 1978. Significance Hargrave, B. T. 1970. Distribution, growth, and seasonal of litter size in Musculiuryi securis (Bivalvia: Sphaeriidae). abundance of Hyallela azteca (Amphipoda) in relation to Ecology 59:1069-1074.

sediment microflora. Journal of the Fisheries Research Mackie, G. L., D. S. White, T. W. Zdeba, and N. A. Thomas Board of Canada 27:685-699. 1980. A guide to freshwater mollusks of the Laurentian Hartnoll, R. G. 1983. Substratum. In: Earll, R. and D. G. Great Lakes with special emphasis on the genus Pisidium.

Erwin (eds. ). Sublittoral ecology: the ecologv of the sub- U.S. Environmental Protection Agency, Duluth, MN, 144 p.

littoral benthos. Clarendon Press, 0.\ford, p. 97-124. McNeil, D. R. 1977. Interactive data analysis: a practical

Healey, M. C. 1978. Sphaeriid mollusc populations of eight primer. John Wiley and Sons, Toronto, 186 p. lakes near Yellowknife, Northwest Territories. Canadian Meier-Brook, C. 1969. Substrate relations in some Pisidium Naturalist 92:242-251. species (Eulamellibranchiata: Sphaeriidae). Malacologia 9:

Holopainen, I. J. 1979. Population dynamics and production 121-125. ol Pisiditim species (Bivalvia, Sphaeriidae) in the oligo- Ostle, B. and R. W. Mansing. 1975. Statistics in research: basic trophic and mesohumic lake Paajarvi, Southern Finland. concepts and techniques for research workers, 3rd ed. Iowa

Archiv fiir Hvdrobiologie 54iSuppl ):466-508. State University Press. Ames, 596 p.

Holopainen, I. 1985 Feeding biology of Pisidiidae (Bivalvia) Rogers, G. E. 1976. Vertical burrowing and survival of sphae- with special emphasis on functional morphologv of the riid clams under added substrates in Pool 19, Mississippi digestive tract Lammi Notes 12:5-9. River. Iowa State Journal of Research 51:1-12. Lotspeitch, R. B and F. H Everest. 1981 A new method for Shannon, C. E. and W. Weaver. 1949, The mathematical reporting and interpreting te.xtural composition of spawn- theory of communication. University of Illinois Press, Ur-

ing gravel United States Forest Service Research Note bana, 125 p. PN\\-369. THE NAUTILUS 102(2):78-81, 1988 Page 78

Hijpselostonia holimanae New Species, a Pupillid Land Snail from Thailand

Fred G. Thompson Harry G. Lee Florida State Museum 709 Lomax Street University of Florida Jacksonville, FL 32204, USA Gainesvilie, FL .32611, USA

ABSTRACT angle. Last whorl flat below peripheral angle, and strong-

Hypselostorna holimanae new species (Gastropoda, Pulmonata, 1\ shouldered above. Occasional specimens may be weak-

Pupillacea, X'ertiginidae) is described from a limestone range 1\ furrowed below periphery. Last whorl ascending at Kancliaiiaburi. near Thailand Its morphological characteristics about 10° to longitudinal axis of spire (figures 2, 4). Neck are distinct to the extent that no close relationship between this of last w horl becoming narrowed behind aperture, and form and other nicmbers of the genus is apparent. Hypselos- extending forward for about ',5 of minor diameter of last torna tubifcrum (Benson, 1856) from Burma is it closest phy- whorl (figure 3); indented externally over junction of logenctic and geographic congener. The two species are alike angulo-parietal lamella and slightK so over colinnellar in details ol sculpture and umbilical width, but differ widely lamella. Base of shell broadK umbilicate due to lateral in shell shape and aperture barriers. Similarities of reduced apertural dentition among other species of Hypselostorna are expansion of last whorl. Umbilicus about 0.40-0.53 times cnnsidered to be due to convergence. minor diameter of last w horl as measured across the basal angle. Whorls 4.6-4.9. Protoconch consisting of about 1.5 whorls that appear smooth under light microscopy; at INTRODUCTION higher magnifications the whorls are sculptured with a dense mesh of fine reticulating threads that have an

The land snail fauna of Thailand is \ery poorly known. underlying spiral arrangement (figure 5). Whorls of te-

The genus Hypselostorna is wideK distributed from Bur- leoconch sculptured with raised spiral threads that are ma through CJamhodia, Vietnam, Mala\a, the Loo C^hoo nearK uniformly distributed o\er the surface of the shell Islands and the Philippine Archipelago. Yet not a single (figure 4). Threads weak liut distinct on spire; most con- species has hitherto been recorded from Thailand. The spicuously developed on last whorl (figure 6). Spiral species de.scribed in this paper was collected by Stephen threads interrupted at irregular intervals b\ incremental C. Holimaii while serving as a Peace Corps volunteer in growth striations, which in some specimens ma\' cause Thailand. During his few opportunities of leisure, Mr. the spiral sculpture to appear cancellate or beaded on Holiman collected mollusk specimens for his mother, the spire. Color dark brow n with a light brow n aperture Mrs. Stanley (Bonnie) Holiman of Jacksonville, Florida, and white lamellae within the aperture. Face of aperture who has a.ssemhled a private collection of fair importance translucent w ith fine radiating lirow n lines due to raised becau.se of the data that accompanies the specimens. The spiral threads on opposite surface Peristome broadly ex- mollusks from Thailand were submitted to us by Mrs. panded and nearK uniformly wide around aperture. Ap- Holiman for identification. The new species of Hypse- erture barrier with four teeth located on inner rim of lostorna described in this paper shows strong similarity the aperture just behind expanding peristome and ar- to the generic t\ pc species from Burma, and differs con- ranged o|5posite each other in a cross-configuration. .An- spicuously troi7i other known species. We are honored gulo-parietal lamella rectilinear, torming a single short to name this snail after Mrs. Holiman in recognition of undulating blade; bifid as is typical for genus, with an- her bringing this species to our attention. gular segment smaller and separated from parietal portion by a weak notch. Palatal and basal plicae short and lateralK llattcned; coiiliiied to inner ritu of peristome. Hypselostorna holimanae new species Columellar lamella tubercular and located on a slight (figures 1-6) callus. Description: Shell small, about 2.6-2.9 mm wide and Measurements in mm for the holot> pe and three para-

2.3-2.6 mm high; about 0.8.5-0.9.5 times as high as w ide. t\ pes (I'F 1 13428) selected to show \ariation follow , The

Shell turlMii-shaped with a moderateK long conical spire minor diameter is the transverse w idth of the bod\ w horl

(figures l-.'3, holotype). Last whorl conspicuously en- posterior to the neck of the aperture. Other measure- larged, and with a distinct peripheral angle and a basal ments were made of standard parameters. F. G. Thompson and H. G. Lee, 1988 Page 79

Figures 1-6. Hypselostoma holimanae new species. 1-3. Holotvpe (UF 113427). x 26. 4-6. Parat\pe (UF 113483). 4. x 40. 5. X 120, 6. X 160. Page 80 THE NAITILUS, Vol. 102, No. 2

Specimen Height Maj. w. Min \v .\per w. L'mbil Whorls Holotype F. G. Thompson and H. G. Lee, 1988 Page 81

chela (Gastropoda. Pulnioiiata, Nertiginidae) with a cat- Solem, A. 1981. Small land snails from Northern Australia, alogue of all the species hitherto described Bulletin of the 1: species of Gtjiiotraclicia Tonilin, 1930 (Mollusca, Pul- Raffles Museum 21 5-47 monata, Nertiginidae). Journal ot the Malacological So-

Jutting, W S. S. van Benthem 1961. Additional new species ciety of Australia 5:87-100 and new localities of the family X'ertiginidae and the gen- Thompson, F. G. and K. Auffenberg 1984. Hypselostoma era Oophana and Opisthostoma from Mala>a. Bulletin of latispira, a new pupillid land snail Irom the Philippine the Raffles Museum 26:34-48, pis. 8-14. Islands. Proceedings, Biological Societv of Washington 97:

Jutting, W. S. S, van Benthem. 1962. Coquilles terrestres 86-89. nouvelles de quelques collines calcaires du Cambodje et Thompson, Fred G. and S. Peter Dance 1983. Non-marine du Sud X'ietnani. Journal de Conchyliologie 102:3-15. mollusks of Borneo, II. Pulmonata: Pupillidae, Clausili-

Pilsbrv, Henr\ .\. 1916-18. Manual of conchology, Ser. II, idae. III. Prosobranchia: Hydrocenidae, Helicinidae Bul-

24. Pupillidae, Gastrocoptinae. Philadelphia, p. i-xii, 1- letin Florida State Museum 29:101-152. 380, pis. 1-49. THE NALTILL'S 102(2):82-87, 1988 Page 82

Eledone gaucha, a New Species of Eledonid Octopod (Cephalopoda: Octopodidae) from Southern Brazil

Manuel Haimovici Dt-partamento de Oceaiiografia Fuiida^ao Universidadf de Rio Grande Cx. Postal 47-1, Rio Crande RS 96.200 Brazil

ABSTRACT This work has been partially supported by a grant of the Conselho Nacional de Pesquisas Cientificas e Tec- A lieu species of Eledone is described from tlie southwestern nologicas (CNPq), Proc. 403293-83 and the surve\ fi- Atiaiitif at depths of 60 to 160 m, off Rio (Jrande do Sul, Brazil. nanced bv the Comissao Interministerial de Recursos do The characters that distinguish this species from the other species of the genus are presented, as well as a morphometric com- Mar (CIRMj. parison with the sympatric Eledone massyae Voss, 1964.

Eledone gaucha new species (figures 2-14, table 1) INTRODUCTION Material examined: Holot\ pe; 6 32.5 mm ML, K \' Several cephalopods were collected during a survey of Atlantico Sul, cruise 13/83, S'ta. 37, 32°58'S, 5ri9'W, 56 the demersal resources of the inner shelf of Rio Grande m, trawl, 17 Nov. 1983, MORG 23544. Parat\pes: 7 6 do Sul between Solidao (30°40'S) and Chui (34°20'S) 27-41 mm ML and 5 9 32-34 mm ML, R \' Atlantico at depths to 100 m (figure 1) by the R/V "Atlantico Sul" Sul, cruise 13/83, Sta. 37, 32°58'S, 51°19'W, 56 m, trawl, of Funda^ao Universidade de Rio Grande (FURG). Hai- MORG 23544; 2 <5 32-34 mm ML, R/\' Atlantico Sul, movici and Andriguetto (1986) stated that two species cruise 13/83, Sta. 39, 32°50'S, 50°45'W, 87 m, trawl, 18 \" of the octopod genus Eledone were found. One of them, Nov. 1983, MORG 23545: 1 S 21 mm ML. R Atlantico E. rnassijae, was described by \ oss (1964) and the second Sul, cruise 13/83, Sta. 2, 31°46'W, 100 m, 9 Nov. 1983, was a new species. Both sympatric species possess the MORG 23846; 1 9 33 nun ML, R/\' Atlantico Sul, cruise generic character of papillae at the tips of the non- 10/83, Sta. 51, 33°43'S, 52°13'W, 60 m, trawl. 29 Aug. hectocot) iized arms of the males. Morphological analysis 1983, MORG 23547; 4 9 28-33 mm ML, R/\' Atlantico presented here as well as ijiochemicai dilferences fouml Sul, cruise 10/83, Sta 57, 33°13'S, 51°25'\\'. trawl, 60 m, by Levi et al. (1985) separate these two similar species. 30 Aug. 1983, MORG 23548; 1

Zoologia da L ni\ersidade de Sao Paulo (MZUSP), Museo Aug. 1983. MC:NLP 4682; 1

University of Miami Mollusks Laboratory (UMML) and MNHN 14763; 1 S 35 mm ML and 1 9 42 inm ML, R/V National Museum ot Natural History, Smithsonian In- Atlantico Sul, cruise 3/83, Sta. 42, 33°02'S, 51°30' \V, 130 stitution (USNM). m, trawl, 17 June 1980, MNRJ 5626, MNRJ 5627. M. Haimovici, 1988 Page 83

DESCRIPTION 29° Animal small, maximum observed mantle length 65 mm (figures 2, 3). Mantle firm, not very thick, ovoid and

elongated (MVVl 3: 61.0; 2: 59.2) separated from head by small constriction. Body surface smooth with some papillae on the dorsal mantle and head. Head narrow er

than mantle (HWl S: 36.2; 9: 37.3); eyes slightly protu-

berant. One supraocular cirrus. Funnel long (FLI S: 44.6;

9: 45.4) with anterior half free (FFuLI 6: 21.6; 5: 21.1); funnel organ (figure 4) W shaped. Arms long and rather slim. Arms length order

1 >2>3>4 in most specimens \\ ith dorsal arms markedly longer than others. All arms longer in males (except hec-

tocotylized arm) than in females (ALl I to IV S: 273.2-

239.8-134.9-208.1; 9: 250.0-213.8-200.6-191.0). Suckers small, uniserial, well separated and deeply set into arms. Suckers somewhat crowded near tips of the arms on females. Two rows of minute fleshy papillae on all non-hectocotylized arms of males (figure 10). Number of suckers on basal half of the first right arms varies from 17 to 23 (ASC 6: 20.1; 9: 19.4), suckers slightly larger on all arms of males (lASI S: 7.7; 9; 6.5). Web extends over half the length of arms and decreases from dorsal to ventral surface; web formula most trequentlv A:B:C:D:E. Web indices similar in both sexes; 24.4-24.1-21. 6-18.6-15.6 for males and 24.4-23.9-21.1- 18.3-14.9 for females. Third right arm in males hectocotylized (figure 9) (HcAI: 58.9). Ligula small (LLI: 8.8), without differentiated calimus (figure 9); spermatophore grove deep. Gill count in external hemibranch from 7 to 10, most

trequentlv 8 in males and 9 in females (Gilc S: 8.2; 9: 8.9). Males reproductive system with no special figures (figure 11). Penis long and tubular (PLI: 23.4) with a rather short diverticulum (PdLI: 7.8). Spermatophores undifferentiated (figure 13) from 12 to 20 mm (SpLI: 45; SpLWT: 1.56). Number of spermatophores in a sample of 40 mature males from 7 to 92 (mean 32.1). The proximal o\ iduct long, the oviductal glands small, the distal oviducts shorter and somewhat stouter (figure 12). Intraovaric eggs oval (figure 14); maximum lengths of apparently mature eggs approximately 8 mm. Num- ber of developing eggs in a sample of 42 maturing females ranged 10-55 (mean 30.2). Buccal mass well developed, with small anterior salivary glands and larger posterior salivary glands. Esoph- agus connects to developed crop leading to muscular stomach and smaller spiral caecum united by two ducts to the large digestixe gland. Intestine thin and leads to the anus adjacent to ink sac opening. Ink sac superficialK embedded in the digestive gland (figure 7). Radula with a tricuspid rachidean tooth, three lateral teeth and a

marginal plate (figure 8). Color of living animals changed from brow n to almost white dorsally always remaining clear ventrally. Color of specimens preserved in alcohol purplish gray dorsally and pale \ellow ventralK'. Inner surface of the arms, mouth, and ventral mantle with few chromatophores. NAUTILUS, Vol. 102, No. 2 Page 84 THE

;-. : :1'. . V" r;" ' Vj

-'T^

r \ mm ....

I,,

10 M. Haimovici, 1988 Page 85

Table 1. Ranges and means of measurements and indites of 10 males and 10 females each of Eledone massyae V'oss, 1964 and Eledune gaticlui new species, from southern Brazil Page 86 THE NAUTILUS, \ol. 102, No. 2

shorter and much thinner in £. gaucha. ExternalK, the

best distinctive character is the arm length pattern. Eledunc caparti was described b\ .Adam (1950) based on five specimens, two males and three females collected in the equatorial west .Africa at depth ranging from 60 to 170 m. No figures or tables were included in the original description. The decreasing arm length and web depth of £. caparti are similar to those of the new species. However, £. caparti does not have a supraocular cirrus, has enlarged suckers at the base of the lateral arms of the males, and the number ot suckers on the dorsal arms

is almost double that in E. gaucha. The radula of £.

caparti has an A2 seriation and the spermatophore is insufficiently described for comparisons. Eledone thysanophora was described by \'oss (1962) based on a single male specimen collected in a tide pool in western South Africa. The morphometric description

is short, but the number of papillae on the tips of the non-hectocotylized arms and the structure of the spermatophore, w ith the inner w all ot the horn portion lined with teeth, differentiate £. thysanophora irom the new species. Summary descriptions of £. cirrhosa (Lamarck, 1798) and £. moschata (Lamarck, 1798) are presented in Roper et al. (1984), and the species are compared b\ Rees (,1956). Both species can be distinguished from £. gaucha by several characters. Eledone cirrhosa has moderately short arms, a ridge along the mantle, non-hectocot\lized arms of males w ith a single row of compressed sucker- Figures 13, 14. Rpprocliictive products of Eledone gaucha like cirri, and spermatophores with spines. Eledone mos- new species. 13. Sperinatophore. 14. Egg. chata has subequal arms, 11 to 12 filaments on the outer hemibranch of the gills, big, sausage-shaped eggs 15 mm long, and a characteristic musk odor. onl\ one eledoiiid. E. inaasyae. A survey of the MORG collection by the author showed one specimen of E. gau- ACKNOWLEDGEMENTS cha (MORG 15'341) lornierK classified as E. massijae. It is expected that reviews in other collections w ill expand J. M. Andriguetto Filho and J. A. Alvarez Perez helped the range of £. gaucha. with collection of the specimens, G. L. Voss and an anon- The sympatric species E. nias.syae and E. gaucha ini- ymous reviewer improved dramaticalK the final version, tially look similar but many differences may be seen in and E. G. Rios stimulated m\ interest in mollusks. To all a more detailed stud\'. In ortler to compare both species of them m\' sincere gratituile. morphologicalK, the same indices were calculated for 10 males and 10 females of E. massyae {collected in the same survey) which were fixed, preserved, and measured LITERATI RE CITED in the same way as the new species (table 1). Eledone .Adam, W. 1950. Notes sur les cephalopodes .\.\11. Deux nou- gaucha is smaller antl has a narrower mantle and head. velles especes de la cote africaine occidentale. Bulletin arms are thinner, longer, The and decrease in size while Institut Ro\ al des Sciences Naturelles de Beigique 2(i(45): in £. massyae all arms are approximately the same length. 1-9.

The hectocotvlized is shorter the arm and web depth Haimovici, M. and J. M .Andriguetto Fo. 1986. Celalopodes decreases troni the dorsal to the ventral surface in E. costeiros capturados na pesca de arrasto do iitoral sul do gaucha. while in E. massyae the web is shorter only Brasil .Archives de Biologia e Tecnologia. Parana 29(3): 473-495. between the ventral arms. The funnel organ is W shaped Levi, A., M. Haimovici. and M. B. Concei^ao. 1985. Car- in E. gaucha, w shaped in £. massyae. The number of J. acteriza9ao eletofortica de dois morfotipos do genero Ele- inner and outer gill lamellae is one unit lower and the done (Cephalopoda: Octopodidae). Resumes do XII Con- arms sucker count two units higher in E. gaucha. .Arm gresso Brasileiro de Zooiogia. (lampinas. 27 Jan to 1 Feb. sucker indices are similar in of species, males both but 1985:39. in females they are smaller in E. gaucha. Perhaps the Palacio, J. F. 1977, .A stud\ of the coastal ceplialopods from best single diagnostic character to distinguish mature Brazil with reference to Brazilian zoogeography. Ph.D. males of both species is the spermatophore, which is thesis, University of Miami, 311 p. M. Haimovici, 1988 Page 87

F. E. L. Voss. 1983. for Palacio, J. F. 1978. Vosseledorie charrua. a new Patagonian Roper, C. and G. GuideHnes taxonomic ceplialopod (Octopodidae) with notes on related genera, descriptions of cephalopod species. Memoirs of the Na- bulletin of Marine Science 28(2):282-296. tional Museum of Victoria 4449-63. species ol F.lcdone Voss, L. 1962. Soutli African cephalopods. Transactions of Rees, W J. 1956. Notes on the European G. with special reference to eggs and larvae. Bulletin British the Royal Societ)' of South Africa 36(Part 4):245-272. Museum of Natural Histor\ .3(6):283-292, pis. 9-10, Voss, G. L. 1964. A note on .some cephalopods from Brazil

Roper, C. F. E., M. J. Sweene>, and C. E. Nauen. 1984. FAO with a description of a new species of octopod: Eledone species catalogue, Vol. 3. Cephalopods of the world. FAO massyae. Bulletin of Marine Science of the Gulf and Ca- Fisheries Synopsis No. 125, 3:227. ribbean 14:511-516. THE NAUTILUS 102(2);88, 1988 Page 88

News and Notices

.54TH ANNUAL MEETING OF THE AMERICAN 16TH CONCHOLOGISTS OF AMERICA MALACOLOGICAL UNION CONVENTION CHARLESTON. SOUTH CAROLINA FORT MYERS, FLORIDA RADISSON FRANCIS MARION HOTEL SHERATON HARBOR HOUSE HOTEL June 19-24, 1988 July 11-15, 1988

The 16th convention of the Conchologists of ,\merica is The 54th annual meeting of the American Malacological scheduled for Jul\ 11-15, 1988. This con\ention will be Liiioii \\ ill be held June 19-24, 19SS in (Charleston, South headquartered in the Sheraton Harbor House Hotel in Carolina. Charleston is a historical cit\, many parts of Fort M\ers, Florida, and hosted b\ the Southwest Florida which have been beautifully restored, as has the Radisson Concliologist Societ\ . Con\ention acti\ ities are centered Francis Marion Hotel, which is located downtown, with- around informative, shell-related programs, fascinating in walking distance of many restaurants, shops and other field trips and interesting tours. .^ fund-raising auction, attractions. Charleston is easily accessible both by air and dealers bourse, and banquet are all planned. A field trip b\ interstate highway. to the Sarasota Fossil Pits is scheduled for Saturda\ , Jul\- Three symposia are planned: Applications of Nucleic 16, 1988. Acid Techniques to the Study of Molluscan Evolution, Pre-registration forms and packets for the Convention, convened b> Dr. M.G. Harasewych, Department of In- Hotel and Field Trip reservations are available from: vertebrate Zoology, National Museum of Natural His- tory, Smithsonian Institution; S\ stematics and Evolution Gene Herbert of Non-marine Mollusks, convened by Dr. Robert 19168 Meadow Brook Court N.W. Hershler, Department of Invertebrate Zoology, National North Fort Myers, FL 33903 USA Museum of Natural History, Smithsonian Institution; and Telephone (813) 731-2405 History of Malacolog\, convened by Dr. W. Backhuys, or

E.J. Brill, Inc Leyden, The Netherlands. Al Bridello 2265 West Gulf Drive # 240E In addition to these ssmposia, contributed papers and Sanibel, FL 33957 poster presentation, scheduled events w ill include a tour Telephone (813) 472-1637 of historic Charleston, guided field trips to terrestrial and marine molluscan communities, an auction to benefit the

s\ inposium fund, and a bantiuet.

For further information, please contact:

Richard E. Petit President, AMU P.O. Box 30 North Mvrtle Beach, SC 29582 USA Telephone (803) 249-1454 INSTRUCTIONS TO AUTHORS

THE NAUTILUS publishes papers on all aspects of the All line drawings must be in black, high quality ink, biology and s\stematics of mollusks. Manuscripts de- clearly detailed and completely labeled. Photographs scribing original, unpublished research as well as review must be on glossy, high contrast paper. All figures are articles will be considered. Brief articles, not exceeding to be consecutively numbered (figs. 1, 2, 3, ... , NOT

1000 words, will be published as notes and do not re- figs. la. lb, Ic, . . . NOR plate 1, fig. 1 . . .). Illustrations quire an abstract. Notices of meetings and other items must be arranged in proportions that will conform with of interest to malacologists will appear in a news and the width of a page (6% inches or 171 mm) or a column notices section. (SVi inches or 82 mm). The maximum size of a printed figure is 6% by 9 inches or 171 by 228 mm. All illus- Manuscripts: Each original manuscript and accompa- trations must be fully cropped, mounted on a firm, white nying illustrations should be submitted in triplicate. Text backing, numbered, labeled and camera ready. The au- must be typed on one side of 8'/2 x 11 inch white paper, thor's name, paper title and figure number(s) should ap- double spaced throughout (including literature cited, ta- pear on the back. Original illustrations must be between bles and figure captions), with at least 1 inch of margin one and two times the desired final size. It is the author's on all sides. All pages must be numbered consecutively. responsibility that the line weight and lettering are ap- If printed on a word processor, the right margin should propriate for the desired reduction. Original illustrations be ragged rather than justified. Authors should follow will be returned to the author if requested. Color illus- the recommendations of the Council of Biology Editors trations can be included at extra cost to the author. Style Manual, which is available from the Council of Biology Editors, Inc., Rockville Pike, Bethesda, 9650 MD Voucher Material: Deposition of type material in a 20814, U.S.A. The first mention of a scientific name in recognized public museum is a requirement for publi- the text should be accompanied the taxonomic au- by cation of papers in which new species are described. thority, including year. Latin names and words to be Deposition of representative voucher specimens in such printed in italics must be underlined; leave other indi- institutions is strongly encouraged for all other types of cations to the editor. Metric and Celsius units are to be research papers. used. The sequence of sections should be: title page, abstract Processing of Manuscripts: Upon receipt, every manu- page, introduction, materials and methods, results, dis- script is acknowledged and sent for critical review by at cussion, acknowledgements, literature cited, tables, fig- least two referees. These reviews serve as the basis for ure captions, figures. The title page should include the acceptance or rejection. Accepted manuscripts are re- title, author's name{s) and address(es). The abstract page turned to the author for consideration of the reviewers' should contain the title and abstract, which should sum- comments. A finalized version of the manuscript is re- marize in 250 words or less the scope, main results and turned to the editor and sent to press. Two sets of proofs conclusions of the paper. The abstract may be followed are sent to the author for correction. Changes other than by a maximum of 8 key words. All references cited in typesetting errors will be charged to the author at cost. the text must appear in the literature cited section and One set of corrected proofs should be sent to the editor vice versa. In the literature cited section, all authors as soon as possible. Authors with institutional, grant or must be fully identified and listed alphabetically. Follow other research support will be billed for page charges at a recent issue of THE NAUTILUS for bibliographic style, the rate of $55.00 per printed page. noting that journal titles must be unabbreviated. Infor- An order form for reprints will accompany the proofs. mation on plates and figures should be cited only if not Reprints may be ordered directly from the printer. included in the pagination. Tables must be numbered Manuscripts, corrected proofs and correspondence re- and each placed on a separate sheet. A brief legend must garding editorial matters should be sent to: Dr. M.G. accompany each table. Captions for each group of illus- Harasewych, Editor, Division of Mollusks, NHB stop 118, trations should be typed on a separate sheet and include National Museum of Natural History, Smithsonian In- a key to all lettered labeling appearing in that group of stitution, Washington, DC 20560, USA. illustrations. i .

: 4 ' •:. I / • ; ' 1 f >.!.;;;.-^:(:-.;. THE NAUTILUS

Volume 102, Number 3 August 29, 1988 ISSN 0028-1344

A quarterly devoted to malacology.

Marine Biological Laboratory . LIBRARY

SEP 6 1988 i

Woods Hole, Mass. EDITOR-IN-CHIEF Dr. Aurele La Rocque Dr. Geeral J. Vermeij Dr. M. G. Harasawych Department of Geology Department of Biology Division of Mollusks The Ohio State University UniversitN of Maryland National Museum of Columbus, OH 43210 College Park, M 20740 Natural Histor\ Voss Smithsonian Institution Dr. Gilbert L. Dr. James H. McLean Washington, DC 20560 Division of Biology and Living Department of Malacology Resources Los Angeles County Museum of Rosenstiel School of Marine and Natural History .Vtmospheric Science ASSOCIATE EDITOR 900 Exposition Boulevard L'niversit\ of Miami Dr. R Tucker A!)bott Los Angeles, CA 90007 4600 Rickenbacker Causeway .\merican Malacologists, Inc. Miami, FL 33149 P O Box 2255 Dr. Arthur S. Merrill Melbourne, FL 32902 SL BSCRIPTION INFORMATION % Department of Mollusks Museum of Comparative Zoologx The subscription rate per volume Harvard University is US $20.00 for individuals and CONSULTING EDITORS C;ambridge, MA 02138 US $30.00 for institutions. Postage Dr. RiJdiger Bieler outside the L iiited States is an Department of Mollusks additional US $2.00 for surface Dr Donald R. Moore Delaware Museum of and US $1000 for air mail. All Division of Marine Geology Natural Histor\ orders should be accompanied by and Geophysics P.O Box 39.37' payment and sent to: THE Rosenstiel School of Marine and Wilmington, DE 19807 NAUTILUS. P.O. Box 3430, Silver .'\tmospheric Science Spring, MD 20901. University of Miami Dr. William K. Emerson 4600 Rickenbacker Causeway Department of Change of address: Please inform Living Invertebrates Miami, FL 33149 the publisher of \our new address The .\merican Museum of at least 6 weeks in advance. All Natural History Mr. Richard E. Petit communications should include New York, NY 10024 P.O. Box 30 both old and new addresses (with North Myrtle Beach, SC 29582 zip codes) and state the effective Mr. Samuel L. H. Fuller date. 1053 Mapleton Avenue Dr. Edward J. Petuch Suffield. CT 06078 THE NAUTILUS (ISSN 0028- Department of Geology 1344) is published quarterly by Florida Atlantic University Dr. Robert Hershler Trophon Corporation, 363 Boca Raton, FL 33431 Division of Mollusks Crescendo Way, Silver Spring, MD National Museum of 20901. Natural History Dr G. Alan Solem Smithsonian Institution Department of Invertebrates Second Class postage paid at Silver Washington, DC; 20560 Field Museum of Natural History Spring, MD and additional mailing Chicago, IL 60605 offices. Dr. Richard S Houbrick Division of Mollusks POSTMASTER: Send address Dr. David H. Stansbery National Museum of changes to: Museum of Zoology Natural History THE NAUTILUS The Ohio State I'niversity Smithsonian Institution P.O. Box 3430 Coliimlnis, OH 43210 Washington. DC 20560 Silver Spring, MD 20901

Mr. Richard I. Johnson Dr Ruth D. Turner Department of Mollusks Department of Mollusks Museum of Comparative Zoology Museum of (Comparative Zoology Harvard I'niversity Har\ard l'niversil\ C:ambridge, MA 02138 Cambridge, MA 02138 .

€9 N An^ LJicaiXio/ L U S THE i j LIBRARY Volume 102, Number 3 August 29, 1988 SEP 6 1988 ISSN 0028-1344 CONTENTS Woods Hole, Mass.

Timothy M. Askew A new species of pleurotomariid gastropod from the western Atlantic 89

IM. G. Harasewych Spongivory in pleurotomariid gastropods 92 Shirley A. Pomponi Timothy M. Askew

James H. McLean A new species of Macrarene (Turbinidae: Liotiinae) from Ricardo Silva Absalao Brazil 99 Renato Luiz dos Santos Cruz

Geerat J. Vermeij Nerita fortidentata, a new gastropod from the Neogene of Timothy M. Collins Panama, with comments on the fossil record of Nerita in tropical America 102

Anthony D^Attilio A new species of Favartia from the eastern Pacific Barbara W. Myers (Gastropoda: Muricidae) 106

William K. Emerson Conus baccatus G. B. Sowerby III, 1877: A Panamic Salter E. Sage III faunal constituent 110

Kenneth J. Boss References to Molluscan Taxa Introduced by Linnaeus in the Systema Naturae (1758, 1767) 115

Mark E. Gordon Frederick Benjamin Isely: Biographical Sketch and malacological contributions 123

William F. Adams Rediscovery of Planorbella magnifica (Pilsbry) in Andrew G. Gerberich southeastern North Carolina 125

Donald M. McKinstry Bite by Octopus joubini: a case report 127

Oscar J. Polaco Occurrence of mites in Mexican land snails 129 Wolfgang Mendl

Richard E. Petit Axelella, new name for Olssonella Petit, 1970, a preoccupied taxon (Mollusca: Cancellariacea) 130

THE NAUTILUS 102(3):89-91, 1988 Page 89

A New Species of Pleurotomariid Gastropod from the Western Atlantic

Timothy M. Askew Harbor Branch Oceanographic Institution, Inc. 5600 Old Dixie Highway Fort Pierce, FL 34946, USA

ABSTRACT Perotrochus charlestonensis new species

(figure 1) Perotrochus charlestonensis, a new species of pleurotomariid,

is described from off the coast of South Carolina. This is the Description: Shell (figure 1) moderately large (maxi- 12th Recent pleurotomariid taxon to be described from the mum diameter 87.4 mm, minimum diameter 80.1 mm, western Atlantic. Its habitat is described, environmental data height 73.0 mm), broadly turbiniform, very thin, fragile; are provided, and comparisons made with closely related con- spire angle 89°, spire slightly convex in profile; proto- geners. conch of 1.0 whorls, translucent, glassy; transition to te-

Key words: Gastropoda; pleurotomariid; Perotrochus; slit shells; leoconch marked by axial costae, with selenizone ap- western Atlantic; JOHNSON-SE,\-LINK parent by second postnuclear whorl; teleoconch of 8'/3 whorls; early whorls nearly flat-sided, becoming progressively more inflated; selenizone near suture in early INTRODUCTION whorls, shifting to slightly below mid-whorl by fifth post-

nuclear whorl; anal slit depth at upper margin 89°, at Since the discovery of the first living species of the pre- lower margin 57°; anal slit width 4 mm; suture adpressed; dominant!) Mesozoic and Paleozoic family Pieuroto- periphery rounded; base inflated, convex, non-umbili- mariidae in the western Atlantic over a century ago cate; nacreous umbilical callus extending '/6 the distance (Fischer & Bernard!, 1856), 24 Recent species and sub- from axis to periphery; spiral sculpture of 21 uniformly species have been described, usually on the basis of one sized spiral cords between suture and anal slit, 20 cords or a very few specimens. The habitat of these animals, of variable thickness between anal slit and periphery, 40 generally steep-wailed, hard substrates at depths in e.xcess cords along base; selenizone with 0-5 broad cords, num- of 100 meters, accounts for their infrequent collection ber increasing with shell size; axial sculpture of weak by such methods as trawling, dredging, and grab sam- nodes on early whorls (88 on fourth postnuclear whorl), pling, and, therefore, for the paucity of data on the forming cancellate sculpture; axial sculpture decreasing, biology and distribution of most species. Since the pub- sculpture limited to spiral cords by sixth postnuclear whorl lication of a review of the Recent pleurotomariids that above selenizone, and seventh postnuclear whorl below included six species from the West Indies (Bayer, 1966), selenizone; aperture broadly ovate; columellar lip slightly three species (Bayer, 1967; Rios & Mathews, 1968; Leme thickened, weakly recurved; color creamy white with & Penna, 1969) and two subspecies (Okutani & Goto, diffuse brownish orange axial streaks and blotches; na- 1983, 1985) have been described from the western At- creous layer visible through porcellaneous layer, creating lantic. iridescent hue; color lighter on base than on dorsal sur- Another new species of pleurotomariid, described face; selenizone margins with cream colored lines most herein, was collected while conducting fish population evident on penultimate and body whorls; aperture na- studies approximately 90 nautical miles east of Charles- creous, iridescent; operculum multispiral (7 whorls), ton, South Carolina, utilizing the submersible JOHN- horny, brownish-yellow, translucent; soft parts unknown. SON-SEA-LINK I (Harbor Branch Oceanographic In- stitution, Inc., Fort Pierce, Florida). Bottom topography Type localitv: 90 nautical miles east of Charleston, South the area at stud\ was extremely rugged, making sampling Carolina (32°43'80"N, 78°05'60"W), in 213 m, R/S by any other means difficult. JOHNSON-SEA-LINK I, dive 1250, August 6, 1982.

SYSTEMATICS Holotype: USNM 859961, maximum diameter 87.4 mm.

Family Pleurotomariidae Swainson, 1840 Etymology: Named after the t\pe localit\ , commonly Genus Perotrochus P. Fischer, 1885 referred to as the Charleston Lumps. THE NAUTILUS, Vol. 102, No. 3

Figure 1. Perotrochus charlestunensis new species. Apertural, right lateral, apical, and basal views of the holotype (USNM 859961 ), off Charleston, South Carolina (32°43'80"N, 78°05'60"W), in 213 m, maximum shell diameter 87,4 mm.

Ecology: This species is known only from the type lo- to black phosphorite. This area was formed during the cality, an area of extremely rugged terrain where the middle Tertiary and has remained stable since the Mio- bottom topograph}' consists of steep, large hills and val- cene (Baturin, 1982). leys. Topographical features (figure 2) consist of a pave- Marine life in the vicinity indicates an area of high ment of relithified phosphorite and fibrous concretionary productivity resulting from warmer Gulf Stream waters. apatite composed of calcium phosphate and other min- Large snowy grouper [Epinephehts niveatus (Valen- erals (Manheim et ai, 1980). This pavement, which ranges ciennes, 1828)] and blue-lined tile fish [Caulolatilus mi- in thickness from 10 cm to almost a meter, has been crops (Goode & Bean, 1878)] are abundant in the area undermined in some areas, causing large pieces to break together with many species of small, deep-reef fish, which off and fall down-slope forming rubble and boulder zones. generally inhabit rocky terrain. Common invertebrates Hills range in height from several meters to about 30 include basket and brittle stars, sea urchins, solitary and meters. Valleys contain sand composed primarily of brown colonial anemones, solitar\ corals, arrow, spider, and gal- T. M. Askew, 1988 Page 91

mer Gutherz, National Marine Fisheries Service, Pas- cagoula, Mississippi; and Tom Smoyer, Harbor Branch Oceanographic Institution photographer, for his excel- RCXKSOULOER LECX3E lent photographs. This is Harbor Branch Oceanographic Institution Contribution No. 642.

HOCK RUBBLE LITERATURE CITED

Baturin, G. N. 1982. Phosphorites on the sea floor —origin, composition and distribution. Developments in Sedimen-

'-a.-.. tology 33:111-112. i^^, SAND PEBBLE Bayer, F. M. 1963. A new pleurotomariid gastropod trawled BARREN SAND in the Straits of Florida by R/V Gerda. Bulletin of Marine Science of the Gulf and Caribbean 13(3):488-492. Figure 2. Cross-section of bottom topography at collection Bayer, F. M. 1966. New pleurotomariid gastropods from the site. Hills range in height from 3 to 30 m. western Atlantic, with a summary of the Recent species. Bulletin of Marine Science 15(4):737-796. Bayer, F. M. 1967. Another new western Adantic pleuroto- atheid crabs, barrel and encrusting sponges, and hy- marian gastropod. Bulletin of Marine Science 17(2):389- droids. A pink featherlike hydroid covers many of the 397. broken boulders along the ridge tops. Prevalent gastro- Bouchet, P. and B. Metivier. 1982. Living Pleurotomariidae pods include Perotrochus amabilis (Baser, 1963), Cal- (Mollusca: Gastropoda) from the South Pacific. New Zea- liostoma sayana (Dall, 1889), Stenorhytis pernobilis land Journal of Zoology 9:309-318. Dall, VV. H. 1887. [A letter containing notes on Antillean (Fischer & Bernardi, 1857), Aurinia gouldiana (Dall, mollusks.] Conchologist's Exchange 2(1):9-10. 1887), and Pterynotus phaneus (Dall, 1889). Dall, W. H. 1889. Reports on the results of dredging ... in

Remarks: Perotrochus charlestonensis is a member of the Gulf of Mexico (1877-1878) and in the Caribbean Sea the species complex consisting of P. midas Bayer, 1966, (1879-1880), by the U.S. Coast Survey steamer "Blake"

. . . Report on the Mollusca. Part II. Gastropoda P. pyramiis Bayer, 1967, P. africanus (Tomlin, 1948), XXIX. — and Scaphopoda. Bulletin of the Museum of Comparative P. teremachii (Kuroda, 1955), P. tangaroana Bouchet & Zoology, Harvard 18:1-492, pis. 10-40. Metivier, 1982, and an undescribed species from off Fischer, P. 1885. Manuel de conchyliologie et de paleonto- northwestern Australia (Group B, Bayer, 1966:745). All logie conchyliologique. Histoire naturelle des mollusques are characterized by having large, thin shells with in- vivants et fossiles. Fascicule 9:785-896. Libraire F. Savy, flated whorls and proportionally large, broadly ovate ap- Paris, xxiv -I- 1369 p., 23 pis. (1887). ertures. This new species most closely resembles P. af- Fischer, P. and A. C. Bernardi. 1856. Description dun pleu- ricana and Perotrochus sp. (Bayer, 1966; fig. 29) from rotomaire vivant. Journal de Conchvliologie 5:160-166, Japan, but differs from these taxa in having a thinner pi. 5. shell w ith more inflated whorls, and a more convex profile Fischer, P. and A. C. Bernardi. 1857. Descriptions d'especes 5:292-300, pis, 8, 9. of the spire. Perotrochus africanus has a more stepped nouvelles. Journal de Conchyliologie Kuroda, T. 1955. A new Pleurotomaria from Japan with a spire, a more strongly recurved and thicker columella, note on a specimen of P. rumphii Schepman collected and a broader umbilical callus (Vi distance from axis to from Taiwan. Venus 18(4):21 1-221, pis. 8, 9, periphery). Of the western Atlantic species, P. charles- Leme, J. P. L. and L. Penna. 1969. Ocorrencia de Mikado- tonensis is most similar to P. pyramus. but is more than trochus no Brasil com descri^ao de uma nova especie. twice the size, and is much higher-spired. Perotrochus Papeis Avulsos de Zoologia (Sao Paulo) 22(21 ):225-230. charlestonensis also somewhat resembles Perotrochus Manheim, F. T., R. M. Pratt, and P. F. McFarlin. 1980. Com- midas, but lacks its characteristic flat, blunt spire and position and origin of phosphorite deposits of the Blake angular periphery. Perotrochus charlestonensis occurs Plateau. The Society of Economic Paleontologists and Min- in shallower water (213 m) than either P. pyramus (420- eralogists, Special Publication No. 29:117-137. T. Y. 1983. new subspecies of Adanson's 648 m) or P. midas (600-770 m). Okutani, and Goto. A slit shell from Bermuda. Venus 42(4):305-311. Okutani, T. and Y. Goto. 1985. A new subspecies of Pero- ACKNOWLEDGEMENTS trochus quoyanus from Bermuda. Venus 44(1):27-31. Rios, E. C. and H. R. Mathews. 1968. Nova especie de Pleu- I thank the following persons, without whose help and rotomariidae do Brasil (Mollusca: Gastropoda). Arquivos guidance this paper would not be possible: Dr. Richard Estaceo de Biologia Marinha da Universidade Federal do Cooper, University of Connecticut; Dr. Roger Theroux Ceara 8(l):65-68. Fisheries Services, and Joseph L zmann. National Marine Tomlin, J. R. le B. 1948. A new species of Pleurotomaria. Woods Hole, Massachusetts; Dr. Walter Nelson and El- Journal of Conchology 23:2, pi. 1. THE NAUTILUS 102(3):92-9S, 1988 Page 92

Spongivory in Pleurotomariid Gastropods

M. G. Harasewych Shirley A. Pomponi Department of Invertebrate Zoology Timothy M. Askew National Museum of Natural History- Harbor Branch Oceanographic Smithsonian Institution Institution, Inc. Washington, DC 20560, USA 5600 Old Dixie Highway Ft. Pierce, FL 34946, USA

ABSTRACT 1935:1129; Hyman, 1967:.360; Yonge k Thompson, 1976: 52); a diet of encrusting invertebrates, predominantly Direct in situ observations of feeding together with analyses sponges (Hickman, 1984a:29); or detrital feeding and of the gut contents of Perutruchus ruidas and P. amabilis in- vegetarian diet (Fretter & Graham, 1976:1). The pres- dicate that these species feed predominantK and selectively on ence of sponge spicules in the gut contents, however, is sponge tissue. Foraminiferal and diatom tests previously reported in the gut contents of pleurotomariids are derived from not necessarily evidence of spongivory. Although mem- planktonic sediment coating the surfaces of sponges and do not bers of the pleurotomariacean family Haliotidae are significantK contribute to the nutrition of these gastropods. The known herbivores (Leighton, 1961; Leighton & Boo- family Pleurotomariidae represents an adapti\e radiation to lootian, 1963; Shepherd. 1973; Fretter & Graham, 1976: likely occurred after the divergence of spongivory that most 6), Leighton and Boolootian (1963:229) reported sponge the herbivorous Scissurellidae and Haliotidae from the pleu- spicules, comprising 1-2% of the gut content \olume, in rotomariid progenitor in the late Paleozoic, but prior to the 25-50/c of the specimens of Haliotis cracherodii they appearance of umbilicate pleurotomariids with deep anal ca- examined. Similarly, Herbert (1987:289) identified sponge nals in the Upper Jurassic. spicules, foraminiferans, annelid setae, and crustacean Key words: Pleurotomariidae; diet; sponges; spongivory; Pero- remains in the alimentary systems of members of the trochus; Mikadotrochus, Entemnutrochus. Solariellinae (Trochidae), a group adapted to feeding on superficial and interstitial detritus. Three species of pleu- INTRODUCTION rotomariids maintained in aquaria have been reported to feed on a wide variety of foods, including starfish,

Since the discovery of the first living species of the family bivalve meat (Arakawa et al., 1978) and sliced, raw fish Pleurotomariidae in the mid-nineteenth century, nu- (Matsumoto et al, 1972; Sekido et al, 1976). merous papers have been published on various aspects The increasing use of research submersibles in deep of the biology and anatomy of these, the most primitive sea investigations has made possible in situ observations living gastropods (e.g., Bouvier & Fischer, 1899, 1902; on the feeding of two species of pleurotomariids. These Woodward, 1901; Fretter, 1964, 1966; Yonge, 1973; observations, supported by gut content analyses of col- Hickman, 1984a, b). As pleurotomariids are predomi- lected voucher material, form the basis of this report. nantly restricted to hard substrates and bathyal depths in the Recent fauna, sampling has pro\ed difficult and MATERIALS AND METHODS most published observations are based on limited material, often poorly preserved. During the course of numerous di\es aboard the research The common occurrence of sponge spicules, forami- submersible JOHNSON-SEA-LINK-II throughout the

niferal tests, and diatoms in the alimentary systems and northern and central Bahamas, li\ ing Perotrochus midas fecal pellets of pleurotomariids has been variously in- Bayer, 1966, were observed on 16 occasions at depths terpreted as being indicative of a spongivorous diet con- ranging 670 to 853 meters. Observations were noted and sisting of a single species of sponge (Woodward, 1901: specimens photographed when practical, using a BEN- 252); a diet consisting principally of sponges (Thiele, THOS 35 mm camera, with 85 mm lens. Two of the

Figures 1, 2. Perotrochus midas Bayer, 1966, and Strongylophora hartmani \'an Soest, 1981, in situ. 1. Perotrochus midas feeding on Strongylophora hartmani. JSL-II dive 1501, #2, west end of Island at Ciouldings Cay, New Providence Island, Bahamas, 25°00'00"N, 77°34'06"\V. in 766 m. October 20, 1987. Specimen not collected Note area of sponge consumed by snail. 2. Perotrochus midas and Strongylophora hartmani along steep wall, JSL-II dive 1505, Chub Cay, Berry Islands, Bahamas, 25°22'22"N, 77°50'25"W, in 777 m. October 23, 1987. Specimen not collected. Note layer of planktonic sediment. Page 93 M. G. Harasewvch et al, 1988 Page 94 THE NAUTILUS, Vol. 102, No. 3

specimens were collected, one together with its prey, upon, the sponge Strongylophora hartmani Van Soest, using the hydraulic arm and clamshell scoop. These spec- 1980 (class Demospongiae, order Haplosclerida) on six imens were fixed in formalin and preserved in 70% eth- occasions (figures 1, 2). Two specimens of P. midas were anol. collected, one (specimen 1) together with the sponge on

Fifty-six specimens of Perotrochus arnabilis (Bayer, which it was feeding. The esophagus of both snails was 1963) were observed and collected during four dives distended and full of sponge tissue. Comparisons of spic- aboard the research submersible Nekton Delta on the ules taken from the esophagus of both specimens of Per-

"Charleston Lumps", an area of rough bottom topog- otrochus midas (specimen 1, figure 5) with those of Stron- raphy approximately 90 miles east of Charleston, South gylophora hartmani (figures 3, 4) confirmed the identity

Carolina (32°43'78"-32°44'90"N, 78°05'68"-78°06'00"\V) of the prey species. The rectum of specimen 1 contained (Askew, 1988:91), at depths ranging from 200 to 230 m. spicules of S. hartmani (figures 6, 7) as well as spicules Samples were collected by suction through a 2 inch (51 tentativeK attributed to a species of PachastreUa (figure mm) diameter hose. Specimens were frozen on dry ice 7). The rectal contents of specimen 2 included spicules and maintained at — 80°C. of S. hartmani, but spicules of an unidentified sponge After identification of the intact sponge on which a belonging to the order Hadromerida (figure 8) comprised specimen of P. midas was feeding, a spicule sample for an estimated 80% of the \olume. The contents of the SEM examination was prepared by digesting a portion esophagus and rectum from both specimens of P. midas of the sponge in 70% nitric acid (HNO5) until only sili- consisted almost entirely (estimated >95% by volume) ceous material remained. Two specimens of P. midas of sponge spicules, with foraminiferal and ostracode tests and three specimens of P. amalnlis were dissected, and accounting for most of the other identifiable remains. sections of the esophagus between the esophageal valve, Of the 53 specimens of Perotrochus amabilis observed situated behind the buccal mass (Fretter, 1964:181, fig. and collected, the majority were on phosphorite blocks 5, ov; 1966:609, fig. 2, v), and the long sphincter at the ranging in size from several centimeters to over a meter opening of the stomach (Fretter, 1966:609) were excised. in length. Only three specimens were on or near a sponge. Also removed were portions of the rectum (Fretter, 1966: In all three instances, the sponge, which was small and fig. 1, r; 1964: fig. 2, r). These sections were teased apart roughly spherical (<6 cm diameter), could not be col- and examined under a dissecting microscope, then treat- lected. Examination of the contents of the esophagus and ed with warm diluted bleach (1-2% sodium hypochlorite, rectum of these three specimens revealed a more het- NaOCl) to dissolve organic material while leaving the erogeneous assemblage of sponge spicules (about 80%), siliceous and calcified remains. The preparations were diatoms, and foraminiferal tests (about 20%) (figures 9, rinsed in distilled water, filtered through 0.4 ^m Nucle- 10). In no instance could more than an estimated 50% pore membrane filters, and the filters mounted directly of the contents of either the esophagus or the rectum be onto SEM stubs. Samples were coated with carbon and attributed to a single species of sponge. gold, and photographed using a Hitachi S-570 scanning electron microscope. A transverse section of the intact DISCUSSION sponge was critical point dried prior to SEM examination.

The following voucher material is deposited at the Direct feeding observations as well as gut content anal- National Museum of Natural History: yses indicate that Perotrochus midas feeds on the sponge Strongylophora hartmani b> rasping large, deep depres- Perotrochus midas: Specimen 2. USNM 857097, JSL-II sions (>2 cm diameter, >1 cm depth) in its surface dive 1501, #2, West end of Island at Gouldings Cay, (figure 1). In each of the two specimens dissected, the New Providence Island, Bahamas, 25°00'00"N, voluminous esophagus was distended, and contained a 77°34'06"W, in 766 m. October 20, 1987. corresponding volume (2-3 cm') of sponge tissue. Rectal Perotrochus amabilis: Specimens 1, 2, and 3. USNM contents also consisted almost exclusively of sponge spic- 846900, DELTA Dive 560, 129 km due east of Charles- ules, though from species belonging to the orders Cho- ton, South Carolina, USA, 32°43'95"N, 78°05'72"W, in ristida and Hadromerida. The most abundant groups of 198-210 m. May 3, 1987. deep water sponges (in terms of biomass) in the tropical western Atlantic and Caribbean are the orders Haplo- RESULTS sclerida and Choristida, with the most common species often being highly silicified (Pomponi, unpublished ob- Of the 16 sightings of Perotrochus midas. this species servations). Although six of the 16 sightings of P. midas was observed in close proximity to, or actively feeding were on or near a haplosclerid sponge, it is unclear

Figures 3, 4. Strongylophora fiartmani Van Soest, 1981. 3. Transverse section, outer surface at top (SEM), scale bar = 300 ^lm. 4. Nitric acid spicule preparation (SEM), scale bar = 200 urn. Figure 5. Perotrochus midas, contents of mid-esophagus of spetimen

1, showing high concentration of Strongylophora hartmani spicules (SEM), scale bar = 200 ^m. Figures 6, 7. Perotrochus midas, rectal contents of specimen 1. 6. Strongylophora hartmani spicules (SEM), scale bar = 200 ^m. 7. Spicules of S. hartmani as well as of ?Pachastrella sp. (arrows) (SEM), scale bar = 200 ^m. Figure 8. Perotrochus midas, rectal contents of specimen 2, with spicules of unidentified sponge, order Hadromerida (arrows) (SEM), scale bar = 200 ^m. M. G. Harasewych et al, 1988 Page 95 Page 96 THE NAUTILUS, Vol. 102, No. 3

Figures 9, 10. Perotrochus amabilis (Bayer, 1963), contents of mid-esophagus of specimen collected 90 miles due east of Charleston, SC, in 210 m. 9. Spicule of an unidentified species, order Poecilosclerida (SEM), scale bar = 20 ^m. 10. Spicules of Strongylophora sp. (small arrow) and an unidentified species, order Choristida or Spirophorida (large arrows) (SEM), scale bar = 100 ^m.

whether prey species are selected, or mereK reflect the pieces of the sponge , and the brush teeth used to "rasp relative abundance of sponges in a nutrient-poor envi- away some of the flesh from the spicules". The occur- ronment. In either case, available evidence indicates that rence of brush or filament-tipped teeth, long considered sponges comprise the major component of the diet of P. unique to Pleurotomariidae, in unrelated, sponge-feed- midas, and that the small amounts (<5%) of foraminif- ing mesogastropods of the genus Seila. has led Hickman eral and ostracode tests as well as other planktonic sed- (1984a:35) to conclude that this tooth morpholog\ is a iment found in the esophagus and rectum of this species functional adaptation to sponge predation and not a phy- were present on the surface of sponges (figures 1, 2), and logenetically constrained, conservative, morphological probably do not contribute significantly to its nutrition. feature. Although far more specimens of Perotrochus amabilis Although exceptions have been documented (e.g., Gra- were observed and collected, fewer were in the proximity ham, 1939; Perron, 1975), archeogastropods are gener- of a sponge. Due to the smaller size of this species and all\ considered to be herbivores (Yonge & Thompson. the inability to collect the sponges, no direct observations 1976). This has led several authors (Yonge, 1973; Hick- of feeding can be documented. E.xamination of the con- man, 1984a) to imply that carnivory was not the original tents of the esophagus and rectum of three specimens mode of feeding of pleurotomariids. Yonge (1973) fur- indicate that sponge spicules comprise approximately ther suggested that the change to a carnivorous diet may 80% of the bleach insoluble mass, the remaining fraction have been associated with the ecological shift of this consisting mostly of foraminiferal tests and diatom frus- family from shallow water reefs, which it inhabited dur- tules. Fretter (1964:182) reported similar contents in the ing the Paleozoic and Mesozoic, to deeper water (>200 stomach of this species and concluded that the animal m), hard substrates by the end of the Eocene. was a microphagous scavenger. We suggest that P. ama- As members of all three Recent pleurotomariid genera bilis, like P. midas, feed exclusively on sponges, but, due have radulae with filament-tipped teeth, it would appear to their smaller size and correspondingly shorter snout, that spongivory in Pleurotomariidae predates the diver- these animals are not able to penetrate as deeply into the gence of the genus Entemnotrochus Fischer, 1885, char- tissues of the sponge, and feed on surface tissues covered acterized by a deep, broad umbilicus and an anal slit with deposits of planktonic sediment, thus accounting extending nearK 180° back from the aperture, from the for the higher proportion of diatoms and foraminifera genera Perotrochus Fischer, 1885, and Mikadotrochus in the gut. Lindholm, 1927, both with shallow (<90°) anal slits and With a single exception (Barnard, 1963), all published without umbilici. Inclusion of the genus Conotomaria reports on the gut contents of pleurotomariids (table 1) Cox, 1959, which is also characterized by having shells list sponge spicules as a major component. Woodward w ith deep anal slits and umbilici, in the same clade as

(1901:252) was the first to speculate that the distinctive Entemnotrochus would date this divergence, and there- radula, shared by all pleurotomariids, was adapted for fore the adaptation to spongivory, prior to the Late Ju- spongivory, with the hooked teeth "tearing away great rassic (Knight et al., 1960). The restriction of the family M. G. Harasewych et ai, 1988 Page 98 THE NAUTILUS, Vol. 102, No. 3

mentary canal of the style-bearing prosobranchs. Pro- Perron, F. 1975. Carnivorous Calliostoma (Prosobranchia: ceedings of the Zoological Society of London 109:75-112. Trochidae) from the northeastern Pacific. The Veliger 18(1):

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Prosobranchia: Trochidae) in southern Africa. Annals of giaires. Masson et Cie, Paris, p. 462-576. the Natal Museum 28t2):283-382. Sekido, M., O. Tsukada, Y. Matsumoto. and T. Kataoka. 1976.

Hickman, C. S. 198-4a. Form and function of the radulae of On the keeping of Teremachi's slit shell, Perotrochus te- pleurotomariid gastropods. The X'eliger 27(l):29-36. remachii. Journal of the Japanese .Association of Zoological Hickman, C. S. 1984b. Pleurotomaria. pedigreed persever- Gardens and .Aquariums 18(2):49-52.

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Yochelson, and R. Robertson. 1960. Systematic descrip- and other Caribbean localities Part II Haplosclerida. tions [Archeogastropoda] In: R. C. Moore (ed.). Treatise Studies of the Fauna of Curacao and the Caribbean Islands

on invertebrate paleontology, Part I, Mollusca 1. Geolog- 62(191):1-173. ical Society of .America and University of Kansas Press, Tachibana, K., P. J. Scheuer, Y. Tsukitani, H. Kikuchi, D. Van Lawrence,' KS, n60-I331. p. Engen, J. Clardy, Y. Gopichand, and F. J. Schmitz. 1981. Komoto, S., O. McConnell, .\. Wright, F. Koehn, W. Thomp.son, Okadoic acid, a cytotoxic polyether from two marine M Lui, and K. Snader. 1987. Puupehenone, a cytotoxic sponges of the genus Halichondria. Journal of the Amer- metabolite from a deep water marine sponge. Journal of ican Chemical Society 103:2469-2471 Natural Products 50:336. Thiele, J. 1935. Handbuch der Systematischen VVeichtier- Leighton, D L 1961. Observations of the effect of diet on kunde. Part 4:1129. shell coloration in the red abalone, Haliotis rufescens Woodward, H. 1885. On Recent and fossil Pleurotomariae. Svvainson. Veliger 4:29-32, pi. 6. Geological Magazine, Decade 3, 2(101:433-439, pi. 11. Leighton, D. L. and R. A. Boolootian. 1963. Diet and growth Woodward, M. F. 1901. The anatomy of Pleurotomaria bey- in the black abalone, Haliotis cracherodii. Ecology 44(2): richii, Hilg The Quarterly Journal of Microscopical Sci- 227-238. ence 44(2):215-268, pis. 13-16.

Lindholm, VV . .\. 1927. O Pleuro!.oniariabeyrichilii\geudor{ Yonge, C M. 1973. Observation of the pleurotomarid En- (Gastropoda) v kolleklsiakh zoologicheskogo muzeia AN 5 temnotrochus adansoniana in its natural habitat Nature zametkoi o rode Pleurotomaria s. lat. Dokladv Akademii 241(5384):66-68. Nauk USSR 24:409-414. Yonge, C. M. and T. E. Thompson 1976. Living marine Matsumoto, Y., T. Kataoka, and M. Sekido. 1972. On the mollusks. William Collins Sons & Co. Ltd., London, 288

behavior of Beyrich's slit shell, Mikadotrochus beyrichi p., 16 pis. (Hilgendorf) kept in the aquarium. Venus 30(4): 147-152. THE NAUTILUS 102(3):99-101, 1988 Page 99

A New Species of Macrarene (Turbinidae: Liotiinae) from Brazil

James H. McLean Ricardo Silva Absalao Los Angeles County Museum of Renato Luiz dos Santos Cruz Natural Histor)- Departamento de Zoologia, 900 Exposition Boulevard Instituto de Biologia, CCS. Los Angeles, California 90007, USA Universidade Federal do Rio de Janeiro Ilha do Fundao, Rio de Janeiro, Brasil 21940

ABSTRACT SYSTEMATICS

The new species Macrarene digitate from the northeast Bra- Family Turbinidae Rafinesque, 1815 zilian coast represents the first record of this principally eastern Subfamily Liotiinae H. & A. Adams, 1854 Pacific genus in the western Atlantic. The species had previously been known from juvenile specimens reported as Liotia Shells of the subfamily are characterized by turbiniform admirabilis E. A. Smith, the holotype of which is not a member to discoidal profiles, nacreous interiors, fine lamellar of the subfamily Liotiinae. sculpture, intritacalx (calcified periostracum) in most genera, circular apertures, and multispiral opercula with calcareous beads. Radula like that of members of other turbinid subfamilies. INTRODUCTION Although previously treated by most authors as a full family, the Liotiinae have recently been ranked as a The species here described was first recognized as a mem- subfamily of Turbinidae by McLean (1987). ber of the Brazilian fauna by Rios (1975, 1985), who referred it to Liotia admirabilis E. A. Smith, 1890. That species was described from the oceanic island of Saint Genus Macrarene Hertlein & Strong, 1951 Helena. According to the original description. Smith's Type species (original designation): Liotia californica species has a maximum dimension of I'A mm. The 15 Dall, 1908. Recent, off Baja California, Mexico. syntypes were examined at the British Museum (Natural History) (catalogue numbers 1889.10.1.1554-68) by the Macrarene species are characterized by turbinate white senior author in 1984 and found to be similar (although shells, broad umbilici, and presence of axial ribs and not clearly referable) to the skeneiform genus Parviturbo spiral cords that form spines at their intersections. Spac-

Pilsbry & McGinty, 1945, which is not a member of the ing of the axial ribs increases in the final whorl. In some Liotiinae. species the ribs then become more closely spaced in the

The Brazilian species was clearly undescribed, but its final quarter whorl. The final lip is not thickened at true generic affinity was not readily apparent because maturity. the specimens available to Rios, which have been ex- Some Macrarene species reach relatively large sizes. amined by McLean (figure 2), were not mature and the The genus differs from Arene in lacking shell pigmen- expression of the mature lip was impossible to determine. tation and in having the spacing of the axial sculpture More recently, two larger specimens have come to light increasingly separated as the shell matures. The white- and it can now be maintained that the species has the shelled genus Liotia is smaller and retains tight spacing characters of the genus Macrarene Hertlein & Strong, of the axial sculpture. 1951. The white-shelled, new world Liotiine genera Mac- Abbreviations for institutions are as follows: LACM, rarene, Liotia, and Cyclostrema differ as a group from Los Angeles County Museum of Natural History; MORG, those of the Indo-Pacific and Australasian regions in lack- Museu Oceanografico, Universidade do Rio Grande, R.S., ing the thickened mature lips that characterize the gen- IBUFRJ, Instituto de Biologia, Universidade Federal do era Bathtjiiotina Habe, 1961, Liotina Fischer, 1885, Rio de Janeiro. Dentarene Iredale, 1929, and Austroliotia Cotton, 1948. Page 100 THE NAUTILUS, Vol. 102, No. 3

Figures 1, 2. Macrarene digitata new species. 1. Holotype, IBUFRJ 1562. x 7.1. 2. Immature specimen, MORG 18.359. x 10.0.

For further remarks on the Indo-Pacific group see McLean and axial sculpture; spines with w eblike interconnections. (1988). Axial ribs narrow across umbilical wall, forming single

There are si.\ previously described species of Macra- descending row of sharp-pointed projections along in- rene: M. californica (Dall, 1908), M. cookeana (Dall, nermost spiral cord. Operculum and radula unknown. 1908), M. diegensis McLean, 1964 (Pliocene); M. far- Type locality: Off northeast coast of Brazil (03°59'N, allonensis (A. G. Smith, 1952), M. lepidotera McLean, 49°35'W), 100 m, Brazilian Naval Research Vessel Al- 1970, and M. spectahilospina Shasky, 1970. All occur mirante Saldaiilia. station 1913, May 6, 1968, 1 speci- offshore in the tropical to temperate eastern Pacific. men.

Type material: Holotype (figure 1), IBUFRJ 1562. Macrarene digitata new species Height 5.0 mm, diameter 6.7 mm. The holot\pe is a (figures 1, 2) dead collected specimen in good condition. Parat\pe, Description: Shell small for genus, turbinate, white, LACM 2377, off Cabo San Roque, Rio Grande do Norte, interior weakly nacreous, ma.ximum diameter 6.7 mm, Brazil (04°30'S, 50°03'\\'), 146 m, Brazilian Naval Re- whorls 3.5, aperture only slightK ohliciue, final lip not search Vessel Almirante Saldanha. station 1921, May 8, thickened. Whorls circular in outline; final w horl in con- 1968 (height 4.3 mm, diameter 6.6 mm). The paratope tact with previous whorl at tips of axial ribs. Shell surface agrees with the holot)pe in size and sculptural details marked by microscopic lamellar growth increments; in- but is in subfossil condition with attached sedimentary tritacalx present. Protoconch diameter about 200 ^m. deposits on the base.

Suture deeply impressetl, first and second whorls rising Referred material: MORG 18.359, 2 immature speci- above protoconch, third whorl descending, resulting in mens [height 2.7, diameter 4.3 mm (figure 2); height 1.5, flat-topped profile for earlv whorls. F"irst teleoconch whorl diameter 2.6] from Paripueira, Alagoas, Brazil, in beach nearly smooth (except for fine lamellar sculpture), second drift collected b\ P. S. Cardoso, December, 1964; MORG with about 20 strong axial ribs and nine, nearly equal 20.620, 2 immature specimens (height 2.1, diameter 3.7 spiral cords defining deep, rectangular pits. Inter.sections mm; height 2.4, diameter 3.6 mm), Fernando de No- of axial and spiral sculpture produce sharpK- projecting ronha Island, 6 m, collected b\ L. Barcellos, January, spines that are slightly upturned adapicalK . Spiral cords 1979. of final whorl increasing to 12; axial ribs decreasing to 15. Spines produced on final whorl by interaction of spiral Etymology: From the Latin digitatus. having fingers. et at., 1988 Page 101 J. H. McLean

Remarks: Macrarene digitata is the smallest species of west coast of Mexico, and in the Gulf of California . . . ,\1V. the genus described to date. However, it is not certain The Mollusca and Brachiopoda Bulletin of the Mu- seum of Comparative Zoology, Harvard University 43: that the holot\pe is mature. Although this genus does 205-487, pis. 1-22. not form a thickened lip, maturit) in other members of Hertlein, L. G. and A. M. Strong. 1951. Eastern Pacific ex- the genus is indicated by closer spacing of the axial ele- peditions of the New York Zoological Society, XLIII. Mol- ments in the final quarter whorl of growth, as indicated lusks from the west coast ol Mexico and Central America. in the original illustration of M. spectabilospina of Shasky Zoologica, Scientific Contributions of the New York Zoo- fig. (1970: 2). The absence of such closer spacing of the logical Society 36:67-120, pis. 1-11. axial element suggests that a quarter whorl of additional McLean, J. H. 1964. New species of Recent and fossil west growth (and a substantial increase in diameter) is possible American aspidobranch gastropods. The Veliger 7:129- for M. digitata. 133. of tropical Pacific Macrarene digitata is unique in the genus in having McLean, J. H. 1970. New species eastern Gastropoda. Malacological Review 2:115-130. all elements of the spiral sculpture of similar strength, McLean, H. 1987. Angariinae and Liotiinae the primitive rather than having a strongly projecting peripheral J. — living trochaceans. .Annual Report of the Western Society carination. Such a sculptural distinction is not regarded of Malacologists 19:16. as a generic level character because generic characters McLean, J. H. 1988. Two new species of Liotiinae (Gastropo- in the Liotiinae are more reliably based on apertural da: Turbinidae) from the Philippine Islands. The Veliger morpholog\', particularly the structure of the final lip. 30:408-411. The most characteristic feature of this species is the Rios, E. C. 1975. Brazilian marine mollusks iconography. fingerlike aspect of the projecting spines. It cannot easily Museu Oceanografico, Fundagao Universidade do Rio be confused with any other member of the Liotiinae. Grande, 331 p., 91 pis. Rios, E. C. 1985. Seashells of Brazil. Museu Oceanografico,

Funda^ao Universidade do Rio Grande, 328 p., 102 pis. ACKNOWLEDGEMENTS Shasky, D. R. 1970. New gastropod taxa from tropical western America. The Veliger 13:188-195. We are grateful to Prof. E. C. Rios for the loan of the Smith, A. G. 1952. Shells from the bird guano of southeast referred material of this species. Farallon Island, California, with description of a new species of Liotia. Proceedings of the California Academy of Sci- LITERATURE CITED ences, Series 4, 27:383-387. Smith, E. A. 1890. Report on the marine Mollusca of the Dall, W. H. 1908. Reports on the dredging operations off the island of Sta. Helena. Proceedings of the Zoological Society west coast of Central America to the Galapagos, to the of London 1890:247-322, pis. 21-24. THE NAUTILUS 102(3): 102-105, 1988 Page 102

Nerita fortidentata, a New Gastropod from the Neogene of Panama, with Comments on the Fossil Record of Nerita in Tropical America

Geerat J. Vermeij Timothy M. Collins Department of Zoology Department of Geology and University of Maryland Geophysics College Park, MD 20742, USA Yale University New Haven, CT 06520, USA

ABSTRACT H,, where Ho is the distance from the horizontal plane on w hich the shell rests to the dorsal surface of the body Nerita fortidentata new species is described from the Neogene whorl. of Bocas del Toro, Panama It is most closely related to the Recent Caribbean \. fulgurans Gmelin, 1791. The fossil record and biogeography of tropical American Nerita are reviewed. SYSTEMATIC DESCRIPTION At least two lineages of Nerita present in tropical America during the Tertiary have become restricted in the Recent fauna Class Gastropoda tn the Infln-Wf^t-Pacific region Subclass Prosobranchia Order Neritacea Family Neritidae INTRODUCTION Genus Nerita Linnaeus, 1758 Neritid gastropods are prominent members of tropical Type species: Nerita peloronta 1758, Recent, tropical rocky intertidai communities. Because ihe fossilization Western Atlantic. potential of tfiese animals is poor, little is known about tlie historical development of the genus Serita Linnaeus, 1758. It was, therefore, of considerable interest to find Subgenus Theliostyla Morch, 1852 an excellently preserved specimen of an apparently hitherto unrecognized species from the Neogene of tropical Type species: Nerita albicilla Linnaeus, 1758, Recent, America. Here we describe the new species and review Indo- West-Pacific. briefly some biogeographically interesting aspects of the history of the genus Nerita in tropical America. Nerita (Theliostyla) fortidentata new species

(figures 1, 2) METHODS Diagnosis: Shell thick, moderately globose (G = 0.59), Species of Nerita are distinguished conchologically by base little excavated (E = 1.10), apex of spire barely characters of shape, apertural dentition, external sculp- raised above rest of shell. Outer lip very thick, inner edge ture, form and sculpture of the parietal callus, and the with 12 strong teeth; two teeth nearest spire \er\ large

shape and sculpture of the calcareous operculum. Two and protruding, as is third tooth from abapical end of ratios are especially helpful in describing the overall shell lip; columellar lip with tw^o strong centrally placed teeth;

form of Nerita (X'ermeij, 1973). The first is globosity, G, adapical portion of parietal region with a fold of about defined as the distance H, between the dorsal surface of the same size and strength as adapical tooth, w hich curves the body whorl and the center of the parietal callus into aperture; sculpture consisting of 21 regularK -spaced divided by the geometric mean betw een the shell's major flat-topped smooth spiral cords, w hich Dare slightly from diameter Dj and minor diameter D2: G = H,/(D,D2)'''. base to barely overhang incised interspaces about one-

The second ratio is the degree of basal excavation. The third the width of ribs; parietal callus small, its surface plane of the parietal callus typically lies at an angle to sculptured with about 10 strong ridges that bear up to the horizontal w hen the shell lies aperture-down on a 3 large granules each; holotype shows faint radial color

flat surface. The greater the angle, the greater is the pattern of alternating continuous and discontinuous pro- degree of basal excavation. An approximation of the de- socline bands of off white and grey-black; operculum

gree of basal excavation is given by the ratio E = H2/ unknown. G. J. Vermeij and T. M. Collins, 1988 Page 103

Figure 2. Nerita fortidentata new species, from Bocas del Figure 1. Nerita fortidentata new species, from Bocas del Toro, Panama. Holotype, 423644, height 19.7 mm. Ab- Toro, Panama, Holotype, USNM 423644, height 19.7 mm, UNSM apertural view. Apertural view, showing thickened outer lip and enlarged teeth on adapical portion of outer lip. Remarks: The new species clearly belongs to the subgenus Theliostyla Morch, 1852 (type N. albicilla Lin- Holotype: United States National Museum number naeus, 1758), which is characterized by granulate sculp- 423644. Major diameter 21.3 mm, minor diameter 16.9 ture on the parietal region, a barely protruding spire, mm, Hi 11.2 mm, H^ 12.3 mm, standard shell height and w ell-developed external spiral sculpture. Among the 19.7 mm (apex abraded), standard shell diameter 19.9 four Recent species of this subgenus in tropical America, mm, shell thickness at midpoint of outer lip 3.5 mm. N.fulgurans Gmelin, 1791, bears the closest resemblance

Type locality: Panama, Province of Bocas del Toro, to A', fortidentata. Measurements of 17 specimens of N. Archipelago of Bocas del Toro, Punta Robalo quadrangle. fulgurans in the Vermeij collection from the Atlantic Island of Cayo Agua, eastern side about 400 meters south coasts of Panama, Costa Rica, Venezuela, and Jamaica of Punta Nispero on the shoreline in clayey, tuffaceous, show that this species is less globose (G = 0.54 ± 0.020, quartzose, blue-grey siltstones with dense shelly horizons. range 0.50-0.57) and basally much more excavated

We have followed Woodring (1982) in referring to the (E = 1.27 ± 0.05, range 1.21-1.39) than is the new Late Miocene-Pliocene deposits of the Bocas del Toro species. Nerita funiculata Menke, 1851, the Recent east- area as the Limones Formation. The true relationships ern Pacific cognate of N. fulgurans, is also less globose between the Miocene-Pliocene of Bocas del Toro with (G = 0.51 ± 0.04, range 0.42-0.58, based on 15 speci- respect to the Costan Rican Limones Formation and the mens in the Vermeij collection from Costa Rica, Panama, Gatun Formation of the Canal Zone have not been elu- and Ecuador) and more excavated (E = 1.28 ± 0.15, cidated. Laurel B>bell of the U.S.G.S. (personal com- range 1.13-1.60) than is N. fortidentata. Both N. ful- munication) has assigned a preliminary age of Late Mio- gurans and N. funiculata have weaker and more nu- cene to Early Pliocene to the locality from which the merous denticles on the outer lip, weaker and more finely holotype of N. fortidentata was collected. This age de- granulated ridges on the parietal region, and spiral cords termination is based on the presence of the calcareous that are more numerous and more variable in size on the nannofossils Sphenolithiis abies Deflandre, in Deflandre body whorl (18-35 in N. fulgurans, usually more than and Fert, 1954 (last occurrence middle Pliocene), and 30 in N. funiculata). The spiral cords of N. fulgurans Discoaster brouweri Tan, 1927 (first occurrence middle show a tendency to bifurcate on the body whorl, whereas

Miocene). Thomas M. Cronin, U.S.G.S. (personal com- no such tendencN is seen in iV. fortidentata. The three munication), examined the ostracode faunas from several subspecies of N. ascensionensis Gmelin, 1791, from is- adjacent localities of the same formation on Cayo Agua. lands in the tropical south Atlantic (Vermeij, 1970) differ He noted a remarkable similarity between the ostracodes from iV. fortidentata by having a nearly smooth parietal from these samples and the ostracode fauna described region and by the very weak dentition on the outer lip. by van den Bold (1967) from the type Gatun Formation The West Indian A', tessellata Gmelin, 1791. differs from of the Canal Zone. On the basis of these similarities he A', fortidentata b\' having low rounded spiral cords bro- suggests a preliminary age of Late Miocene for this fau- ken irregularl) bv high and low areas correlating to the na. Harry Dowsett, also of the U.S.G.S. (personal com- characteristic black and white checkered pattern found munication), examined the planktic foraminifera from in this species, a finely granulated parietal region of an adjacent localit\ of the same formation on Cayo Agua relatively large extent, and a weakK denticulated outer and found an assemblage indicative of planktic zone lip (Russell, 1941). N17-NT8 (Late Miocene-Early Pliocene). The consensus Jung (1965) recorded Serita fulgurans from the Mid- at this stage, therefore, is that the beds from which N. dle Miocene and Upper Pliocene of Venezuela, but he fortidentata was collected are Late Miocene to Early pointed out that his specimens differed from Recent shells Pliocene in age. by having stronger apertural dentition and a less concave 1

Page 104 THE NAUTILUS, Vol. 102, No. 3

(that is, less excavated) parietal region. Later Jung (1969) .\merica, at least two have close affinities with li\ing found a similar shell in the Late Miocene Melajo Clay Indo-\^'est-Paci£ic species. Merita listrota Woodring, Member of the Springvale Formation of Trinidad. Like 1973, from the late Eocene (?) Gatuncillo Formation of the Venezuelan material, the shell from Trinidad has Panama has a finely ribbed shell w ith a peripheral keel,

only 16 ribs, but instead of having 2 centrali\ placed fine teeth on the outer lip, 7 teeth on the columellar lip, columcllar teeth, as in the Venezuelan material, the Trin- and a sparseK papillate parietal region. Woodring (1973)

idad specimen has one upper tooth which curves into noted the striking similarity' between A', listrota and the the aperture (this upper tooth is probably equivalent to recent mangrove-associated M. planospira Anton, 1839 the parietal fold found on the tvpe of \. fortidentata] (the type and only known species of the subgenus Ily- and 2 somewhat lower denticles. Jung tentati\ely re- nerita von Martens, 1887) from the tropical Indo-Pacific. ferred both lots to N. exuvioides Trechmann, 1935, a He doubted that the tw^o species were closely related, in species described on the basis of one incomplete speci- part because M. planospira has 5 rather than 7 columellar men from the Pliocene of Carriacou in the Grenadines. denticles. We consider the similarities to be so numerous Jung (1971) redescribed this latter specimen as having that an inference of close relationship seems warranted. only 12 ribs whose edges overhang the adjacent inter- If A', listrota belongs to the subgenus Ilynerita, as we

spaces. Vokes (1983) clarified the status of N. exuvioides believe it does, that subgenus may be added to the grow- when she described a very strongly ribbed shell with 1 ing list of taxa whose distributions became restricted to ribs from the Gatun Formation of Panama. This speci- the Indo- West-Pacific during the Tertiary (N'ermeij, men closely resembles the one from Carriacou and is 1986). As Vokes (1983) points out, \. (Theliostyla) exu- clearly referable to S. exuvioides. We believe that Jung's vioides may represent a second lineage that has become (1965, 1969) specimens from Venezuela and Trinidad restricted (as N. exuvia Linnaeus, 1758) to the Indo- belong neither to N. fulgurans nor to N. exuvioides, but West-Pacific. Merita exuvioides differs from the Western

instead to our new species, iV. fortidentata. This species Pacific M. exuvia chiefly by having 1 1 instead of 14 strong appears therefore to be intermediate in sculpture be- overhanging spiral cords on the body whorl. tween N. exuvioides with only 1 1-12 ribs and the Recent The other fossil species of Merita that have been de- .v. fulgurans, usually with more than 21 ribs. The re- scribed from late Eocene and younger deposits in tropical

lationship between N. fortidentata and the Late Oli- America do not easih fit with any living members of the gocene or Early Miocene N. tampaensis Dall, 1892, from genus. These are M. hadra Woodring, 1973, from the

the Tampa Formation of Florida is unclear. Merita tam- late Eocene (?) Gatuncillo Formation of Panama and N. paensis, whose granulated parietal region suggests place- oligopleura Dall and Ochsner, 1928, from the Pleistocene ment in Theliostyla, is a small species with weakly de- of the Galapagos. Merita hadra has very fine spiral threads veloped apertural dentition and highly \ariable external on the bod\' whorl and a smooth parietal region, whereas sculpture, some shells being nearly smooth whereas oth- M. oligopleura has 3 broad spiral ribs on the body whorl.

ers have fine spiral cords of varying sizes. Although the record of Merita in tropical America is The shape of xV. fortidentata suggests that this species very meager, the history of the genus points to multiple

inhabited the upper zones of rocky shores. Merita ful- instances of extinction and geographical restriction. It is gurans, its most similar living relative, is usually found too earl\ to assess the scope of these changes, but the

in areas of reduced salinity , such as the mouths of harbors record of Merita suggests that intertidal species have been (Russell, 1941) or protected embayments. Vermeij (1973) no less affected by events leading to extinction and re- showed that shells with low globosity, high basal exca- striction than have species from the better-sampled fossil vation, weak sculpture, and relatively broad apertures environments of subtidal bottoms. with weak dentition are found in middle to lower intertidal species of Merita; whereas species with a globose, ACKNOWLEDGEMENTS little excavated, strongly sculptured shell and a small aperture bordered by strong teeth are found at higher T.M.C. gratefully acknowledges a Smithsonian Tropical Research Institute short-term fellow ship, shore levels. The latter shell form is especially charac- directed by Jer- teristic of the subgenera Cyrnostyla von Martens, 1887, emy Jackson, during w hicli large collections of Late Ter- tiary fossil mollusks and Ritena Gray, 1858. Of the living and fo.ssil members were made. Collecting was made of the subgenus Theliostyla, most of which live in the more fruitful and pleasant by the expertise and company of the other members of the field part\-, A. G. and M. middle zones of the intertidal, M . fortidentata most closely J. approaches species of Ritena. Other fossils collected with Coates. We thank Laurel B\ bell, Thomas M. Cronin, and N. fortidentata, including Oliva, Olivella, Conus, Na- Harry Dowsett of the U.S.G.S. for their age determi- tica, Polinices, Strombus, Phalium, Dentalinm. and nations. Corlmla, suggest a variety of different environments, implying post-mortem transport and mixing of assem- LITERATURE CITED blages. Anton, H. E. 1839. Verzeichniss der Conchylien welche sich BIOGEOGRAPHY OF AMERICAN MERITA in der Sammlung von Hermann Eduard .^nton befinded. Halle, xvi -I- 110 p Among the fossil species of Merita that have been de- Bold, W. A. van den. 1967. Ostracoda of the Gatun For- scribed from late Eocene and younger strata in tropical mation, Panama. Micropaleontology 13(3):306-318. Page 105 G. J. Vermeij and T. M. Collins, 1988

Dall, W. H. 1892. Contributions to the Tertiary fauna of Trechmann, C. T. 1935. The geology and fossils of Carriacou, Florida; with especial reference to to the Miocene silex West Indies. The Geological Magazine 72(858):528-555, beds of Tampa and the Pliocene beds of the Caloosahatchie pis. 20-22.

River. Transactions of the Wagner Free Institute of Sci- Vermeij, G. J. 1970. The Nerita ascensionensis species com- ence 3(2):201-473, pis. 13-22. plex (Gastropoda: Prosobranchia) in the South Atlantic. Dall, W. H. and W. H. Ochsner. 1928. Tertiary and Pleis- The Veliger 13(2):135-138. in tocene Mollusca from the Galapagos Islands. Proceedings Vermeij, G. J. 1973. Morphological patterns high intertidal of the California Academy of Science 17{4):89-139, pis. gastropods: adaptive strategies and their limitations. Ma- 2-7. rine Biology 20:319-346.

its Vermeij, G. 1984. The systematic position of the neritid Gray, J. E. 1858. Observations on the genus Nerita and J. operculum. Proceedings of the Zoological Society of Lon- prosobranch gastropod Nerita polita and related species. don 26:92-94. Proceedings of the Biological Society of Washington 97(4):

I, 688-692. Gmelin, J. F. 1791. In: Linnaeus' Systema naturae, Tom. crises: Pare VI, p. 3021-3909. Vermeij, G. J. 1986. Survival during biotic the prop- Jung, P. 1965. Miocene Mollusca from the Paraguana Pen- erties and evolutionary significance of refuges. In: D. M. insula, Venezuela. Bulletins of American Paleontology Elliott (ed.). Dynamics of extinction. Wiley, New York, p. 49(223):389-652. 234-246. Jung, P. 1969. Miocene and Pliocene mollusks from Trinidad. Vokes, E. H. 1983. Nerita exuvioides Trechmann (Mollusca: Bulletins of American Paleontology 55(247):293-657. Gastropoda) from the Gatun Formation of Panama. Tu- Jung, P. 1971. Fossil molluscs from Carriacou, West Indies. lane Studies in Geology and Paleontology 17(4): 131-134. Bulletins of American Paleontology 61(269):147-262. Woodring, W. P. 1973. Geology and paleontology of Canal Linnaeus, C. 1758. Systema naturae, 10th ed. Stockholm, Zone and adjoining parts of Panama: description of Ter-

824 p. tiary Mollusks (additions to gastropods, scaphopods, pe- Martens, E. von. 1887-89. Die Gattungen Nerita und Neri- lecypods: Nuculidae to Malleidae). U.S. Geological Survey topsis. Svstematisches Conchylien-Cabinet von Martini und Professional Paper 306-E:453-539. Chemnitz 2(11):1-147. Woodring, W, P. 1982. Geology and paleontology of Canal Menke, K. T. 1851. Conchylien von Mazatlan, mit kritischen Zone and adjoining parts of Panama: description of Ter-

Bemerkungen. Zeitschrift fiir Malakozoologie, yr. 8. tiary Mollusks (Pelecypods: Propeamussidae to Cuspida- Morch, O. A. L. 1852 Catalogus conchyliorum quae reliquit riidae; additions to families covered in P.306-E; additions D. Alphonso d Aguirra & Gadea Comes de Yoldi, Copen- to gastropods; cephalopods). U.S. Geological Survey

hagen, 170 p. Professional Paper 306-F:541-759. Russel, H. D. 1941. The Recent molluscs of the family Ne- ritidae of the western .Atlantic. Bulletin of the Museum of Comparative Zoology at Harvard College 88(4):347-404. THE NAUTILUS 102{3):106-1{)9, 1988 Page 106

A New Species of Favartia from the Eastern Pacific (Gastropoda: Muricidae)

Anthony D'Altilio

Barbara ^ . Myers Department of Marine Invertebrates San Diego Natural History Museum San Diego, CA 92101, USA

ABSTRACT Type species: Mtirex hidalgoi Crosse, 1869, b>- original designation. Favartia (Murexiella) shasktji is described from Isla del Coco, Costa Rica, and compared with related species from the eastern

Pacific, This species is known onl\ from this isolated oceanic Favartia (Murexiella i shaskyi new species island (figures 1-10)

Description: Shell (figures 1, 2) broadly fusiform, spire

'/3 shell length; protoconch of holot\pe eroded, proto- INTRODUCTION conch of paratype 1 (figure 3) of 3'/2 smooth, lustrous, convex whorls, with axial buttresses arising from the te- Isla del Coco (also known as Cocos Island), one of the leoconch; teleoconch with 5-6 subangulate whorls; suture National Parks of Costa Rica, is a small, uninhabited weakly impressed; whorls gentK sloping from suture to island situated approximately 600 kilometers south south- shoulder; body whorl % shell length; aperture ovate; in- west of Puntarenas, Costa Rica, at 5°33'N latitude and ner lip erect posteriorly; outer lip thin, erect, crenulate, 87°03'W longitude. Isla Cascara and Roca Sucia are two reflecting the spiral cords; siphonal canal '3 shell length, of the many islets surrounding Cocos Island. In May, broad proximalK', tapering distalK', with narrow \entral 1985, Donald R. Shasky of Redlands, California, and opening, terminally recurved, tube-like; siphonal fascicle Kirstie L. Kaiser of La Canada, California, collected five of 3 fine distal portions of previous canals; body whorl specimens of the new species described herein, at depths with 6 broad varices, penultimate whorl with 8 varices; of 50-80 feet (15.2-24.4 m), under dead coral off these varices broader than intervarical regions; spiral sculpture two islets. In a previous paper (D'Attilio, M>ers & Shasky, of 6 cords on body whorl, all nearK equal in width, et al., 1987), a new species of Phyllonotus Swainson, packed closely together; 2 additional cords between 1833, was described from the same area. Montoya (1983, body whorl and siphonal canal, 2 major and 1 minor 1984) published a bibliography of Cocos Island molluscan cord on canal; 2 cords per whorl on spire; shoulder of faunal studies. body whorl without spiral cords; all cords terminate as spines on varices (figures 4, 5), posterior spines recurved, anterior spines long, projecting ventrally; spiral cords

SYSTEMATICS (figures 6, 7) strongly scabrous; scales prominent, fine, white, erect, closely packed, disguising the contour of Family Muricidae Rafinesque, 1815 cords; cords and scales microscopicalK grooved, incre- SubfamiK Muricopsinae Radvvin & D',\ttilio, 1971 mentally incised; adapertural sides of varices, especially Genus Favartia Jousseaume, 1880 last varix, completeK- scaled; scales arranged in 3 tiers (figure 6), with tube-like spines on tier farthest from Type species: Miirex breviculus Sowerby , 1 834, by orig- aperture; radula (figure 8) t>picall\ muricopsine, with inal designation. broad, U-shaped basal plate, and strongK projecting cusps.

Paratypes 1 and 2 with only 4 postnuclear whorls; suture Subgenus Murexiella Clench & Perez Farfante, 1945 more strongly impressed, spire angulate.

Isla Cascara, Cocos Island, Costa Rica, under Figures 1, 2. Favartia (Murexiella) shaskyi new species. 1 . Holot> pe, L'SNM 860012, Island, Costa Rica, under dead coral dead coral slab, in 24.4 m, 23.0 mm long. 2. Paratype 1, SDNHM 91873, Roca Sucia, Cocos slab, in 24.4 m, 14.0 mm long. 107 A. D'Attilio and B. W. Myers, 1988 Page Page 108 THE NAUTILUS. Vol. 102, \o. 3

Figures 3-7. Favartia (Murexiella) shaskyi new species, details of shell sculpture 3. Protoconch of parat\pe 1. 4, 5. Adapertural

(4) and abapertural (5) views of last varix of holotype. 6, 7. Microscuipture along adapertural (6) and abapertural (7) surfaces of a spiral cord of the holotype. Figure 8. Rachidian and right lateral teeth from the holotype.

Color: Base color buff to tan, with diffuse darker band Additional material examined: Tw o specimens, collec- encircling body whorl; scales white, aperture white, sition of K. Kaiser, 19.2 mm long, and 14.7 mm long, Isla phonal canal white. Cascara, Cocos Island, Costa Rica, under rocks, in 15.2- 24.4 m. Type locality: Isla Cascara, Cocos Island, Costa Rica, are pleased to name this species for under dead coral slab, in 24.4 m. Etymology: We Donald R. Shasky, M.D., who collected the three type long, 14.2 Holotype: USNM 860012 (figure 1 ), 23.0 mm specimens, and who generousK donated the holotype wide. mm and parat\pe 1.

Paratypes: Paratype 1: SDNHM 91873 (figure 2), 14.0 DISCUSSION mm long. Paratype 2: Collection of D. R. Shasky, 11.8 mm long, both from Roca Sucia, Cocos Island, Costa This new species has a distinct morphology that easily Rica, under dead coral, in 24.4 m. distinguishes it from all other eastern Pacific species of A. D'Attilio and B. W. Myers, 1988 Page 109

I. ante. the subgenus Murexiella. Favartia (Murexiella) lappa Clench, W. J. and Perez Farf 1945. The genus Mt/rei (Broderip, 1833), a closely related species, differs in hav- in the western Atlantic. Johnsonia l(17):l-58. 1869. Diagnoses Molliiscorum novorum. ing a higher spire, biconic form, and short, stubby, non- Crosse, H. Journal de Conchyiiologie 17:408-410. recurved spines. Favartia (Murexiella) vittata (Broderip, D'Attilio, A., B W. Myers, and D. R. Shasky. 1987. A new 1833) also has a more or less biconic shape, but the body species of Phyllonotus (Muricidac: Muricinae) from Isla whorl is more globose, and the spines, although recurved, del Coco, Costa Rica. The Nautilus 101(4):162-165, are short and stubby. Favartia (Murexiella) keenae (E. Jousseaume, F. 1880. Division methodique de la famille des H. Yokes, 1970) has somewhat similar sculpture, a larger Purpurides. Le Naturaliste 42:335-336. shell with a globose body whorl strongly constricted at Montoya, M. 1983. Los Moluscos marines del la Isla del Coco, the base, and a strongly impressed suture. We have care- Costa Rica. I. Lista anotada de especies. Brensia 21:325- fully studied and compared the 12 additional nominal 353. species of Murexiella, none of which are closely related Montoya, M. 1984. Marine mollusks of Cocos Island, Costa

Rica. I. Bibliographic compilation of species. Western to this new species. So- ciety of Malacologists Annual Report (1983) 16:33-44, ACKNOWLEDGEMENTS Radwin, G. E. and A. D'Attilio. 1971. Muricacean supra- specific taxonomy based on shell and radula. Echo 4:55- We are grateful to Dr. Donald R. Shasky for donating 67. Rafinesque, C. S. 1815. Analyse de la nature ou tableau du the holotype and paratype and Kirstie Kaiser for the loan univers et des corps organises. Barravecchia, Palermo, p. of her two specimens. We thank David K. Mulliner for 136-149. the photography. M. G. Harasewych, EmiK H. Yokes, Sowerby, G. B., II. 1834. Conchological illustrations. Murex. William K. Emerson and Carole M. Hertz reviewed the Sowerby, London, pis. 58-67. manuscript and made valuable suggestions. Swainson, W. 1833. The zoological illustrations, ser. 2, vol. 3,

pi. 67. LITERATURE CITED Yokes, E. H. 1970. The west American species of Murexiella (Gastropoda: Muricidae). Veliger 12(3):325-329, pi. 50. Broderip, W. 1833. Characters of new species of Mollusca and Conchifera collected by Mr. Cuming. Proceedings of the Zoological Society of London 2:173-179. THE NAUTILUS 102(3):110-114, 1988 Page 110

Conus baccatus G. B. Sowerby 111, 1877 A Panamic Fauna! Constituent

^ illiam K. Emerson Waller E. Sage III Department of Invertebrates American Museum of Natural History New York, NY 10024, USA

ABSTRACT Atlantic. \'ink (1984:356) subsequently noted certain similarities of the holotype of C. baccatus with C. selenae Specimens of a small species of Conus recent!) received from Van Mol, Tursch, and Kempf, 1967, a Brazilian taxon Golfo de Chiriqui, Republica de Panama, are confirmed to be which we consider referable to the C. jaspidcus (Jmelin, referable to Conus baccatus Sowerby , 1877, a species originally 1791, species complex. described from an unknown locality. The provenance and iden- generosit)- of Carol Skoglund and Robert tity of this taxon had remained uncertain for more than 100 Through the years until Rockel (198.5a) compared two specimens from Pa- Koch, we have examined a series of well-preserved spec- cific Panama in his collection with the holotype of Conus bac- imens from Golfo de Chiriqui. Panama that are referable catus and determined the specimens to be conspecific. We to Coijus baccatus. Our findings confirm Rockel's (1985a: this species has been con- agree with Rockel's conclusion that 29) conclusion that this taxon is a valid Pacific Panamic form of the west American fused in the past w ith the pustulate species, which for many years has been masquerading Conus perplexus Sowerby, 1857, which is a somewhat similar, in collections as a pustulose form of Conns perplexus but distinct species. For Conus perplexus, a lectotype is selected Sowerb), 1857. and the type locality is restricted to the Golfo de Panama. For A specimen apparently referable to C. baccatus was Conus baccatus. the Golfo de Chiriqui is designated the type illustrated by Kiener (1846:56, 57, pi. 83, fig. 2) as "Conus locality. albomaculatus. Reeve ", in reference to Sow.[erby] (1833,

Conus fig. 2, which was named C. bicolor h\ Sowerby in 1833 and renamed C. albomaculatus h\ Sowerb) in INTRODUCTION 1841). Kiener "s figured specimen, measuring 22 mm in height, was attributed to the Dupont Collection from an In 1877, G. B. So\verb%- III briefly described and illus- unknown locality. This specimen cannot be located in trated in color Conus baccatus from an unknown locality. the Museum National d Histoire Naturelle, Paris, and Apparently based on a single specimen in the collection the fate of the Dupont collection is not known (P. Bou- of Dr. Prevost of Alen^on, France, the figured specimen chet, in lift.. March 4, 1987). Reeve (1849:3) considered

(Sowerby, 1887:251) Kiener 's illustrations to represent the western Atlantic later was owned by J. C. Melvili species C. mindanus Hwass in Bruguiere, 1792. He also and subsec}uently by J. R. Le B. Tomlin (1937:217). The vast MeK ill-Tomlin Collection eventual!) w as deposited, stated that "C. albimaculalus [sic] [Sowerb) ] has more along with this type, in the National Museum of Wales, resemblance with very young [pustulose] specimens of 1792] Cardiff. Coomans et al. (1982:4) examined the holotype C. lithoglyphus [.sir] [Hwass in Bruguiere, ', a con- and provisionally accepted Conus baccatus as a valid clusion accepted by Coomans et al. (1979:97) and Rocke! species. They also rejected the Cjuestionable placement (1985b, nr. 22). Sowerb)- "s figured specimen of C. albo- by Walls (1979:726) of this ta.xon in the synonymy of C. maculatus has not been located. It is not in the Britisli mindanus Hwass in Bruguiere, 1792, from the western Museum (Natural History) (K, M. Way, in litt.. May 11,

Figures 1-7. Conus baccatus Sowerby, 1877. 1, 2. Holotype, NMW no. 1955.158.29; x 2 (courtesy of R. M Filmer) 3. 4.. Off x Isla Coiba, Panama. Isla Coiba, Panama, AMNH no, 221871a; x 2. 5. Off Isla Parida, Panama, AMNH no. 173698a; 5 6, 7. Off x 10. Pla\a AMNH no. 221871; x 2, Figures 8-16. Conus perplexus Sowerby, 1857, 8. 9. Lectotype, BM(NH) no. 1978118; 2. P. Skoglund; de los Angeles, Bahia Tenacalita, Jalisco, Mexico, dredged, mud bottom, Aug., 1975, AMNH no. 221873a, ex C and x "West X 5. 11, 12. Paralectotype, BM(NH) no. 1978118a; x 2 13, 14. Paralectotype, BM(NH) no. 1978118b; 2. 15. 16. Panama" (note the distorted spire), AMNH no. 212547; x 2 Figures 17. 18. Conus puncticulatus form papillosus Kiener, 1849, no. 225979; x 2. Figure 19. Conus yemanjae \aii Mol et al., Cartagena, Colombia, Iseach, 1977, ex J. M. Bijur Coll., AMNH 1967, paratype, pi. 8, fig. lb, Fortaleza (Ceara), Brazil, ex pisce; x 2. W. K. Emerson and W. E. Sage III, 1988 Page 111 Page 112 THE NAUTILUS, Vol. 102, No. 3

1987) nor is it in the National Museum of Wales (A. in 5.5-9 m, sand bottom. May, 1972, ex R. E. Hubert; 2 Trevv, in litt.. Ma\, 1987). Therefore, the taxonomic specimens, no. 210849, off Isla Parida, sand, in 5.5 m, status of C. alboviaculatus Sowerby, 1841, cannot be March, 1974, ex R. E. Hubert; 3 specimens, no. 225981, the available Isla determined from data. off de Canal de Afuera, in 55 m, ex J. Ernest; 2 Although C. baccatus was not formally named by Sow- specimens, no. 221871, off Isla Coiba, in 24-40 m, March erb\ ill until 1877, specimens were apparentl\ available 28, 1986, ex C. and P. Skoglund; 2 specimens, no. 221872, in European collections by the nii(l-I9th century, as sug- off S.E. Isla Rancheria, in 2-7.5 m, March 27, 1986, ex gested by Kiener's figured specimen from the Dupont C. and P. Skoglund; 2 specimens, no. 223269, off Isla collection. There is also a specimen of C. baccatus labeled Rancheria (7°38'N, 81°44'W), in 3-9 m, coral sand and ""Panama" in the F. A. Constable Collection (AMNH debris bottom, February 24, 1987, ex R. Koch. From

47740), which dates from the latter part of the 19th "Panama"; 1 specimen, no. 47740, ex F. A. Constable century and was received by the American Museum of Coll.; 3 specimens, no. 201566, ex T. H. and V. B. Mun- Natural History from the estate of Louise Constable in yan Coll. 1929. Remarks: Specimens of Conus baccatus superficially Abbreviations for institutions used in text; AMNH = resemble small examples of the pustulose form of Conus .\merican Museum of Natural History; BM(NH) = Brit- perplexus. There are several discrete differences in shell ish Museum (Natural History); Los Angeles County Mu- morphology which serve to distinguish the two taxa. 1, seum of Natural History = LACMNH; NMW = National Nuclear whorls: In C. baccatus, I'i whorls, nucleus short, Museum of Wales. terminating in a broad, well-defined mammillated pro- SYSTEMATIC ACCOUNT cess. In C. perplexus, 2'/2 whorls, nucleus elongate, narrow, terminating in a needle-like projection (cf. figure 5 Conus baccatus Sowerby, 1877 with figure 10). 2, Shoulder margin of bod\ whorl: In C. (figures 1-7) baccatus, oblique, twofold-angled (i.e., biangulated). In

? "Conus albomaculatus. Reeve"', Kiener, 1846:56, 57, pi. 83, C. perplexus. monoangulated (cf. figures 6, 7 with figures fig. 2 (apertura! and dorsal views; as "Conus albimaculatus 8, 9). 3, Spiral granulations; In C baccatus, rows of pus-

[sic] Sow.[erby] ), from unknown locaiit) , Dupont collec- tules more widely spaced, some rows weakK developed. tion. Not C. bicolor Sowerby, 1833: pt 24, fig. 2, renamed In C. perplexus, rows consistently spaced, nearly uni- without explanation C. albomaculatus Sowerby, 1841. formly developed (cf. figures 6, 7 with figures 15, 16). Conus baccatus Sowerby, 1877:753, 754, pi. 75, fig. 5, from 4, Color pattern: In C. baccatus, spiral bands of broken unknown locality; Tryon, 1883: ser. 1, vol. 6, p. 22, pi. 6, blotches, orange in faded specimens, reddish-broun in fig. 92 [copy of original fig.], "Habitat unknown "; Sowerby, fresh specimens. In C. perplexus, narrow spiral lines with 1887: vol. 5, p. 251, Conus pi. 29, fig. 660 [copy of original dark brown dots figures 1, 2 and 3, 4 with figures fig.], "Habitat unknown. Coll. Meiviii"; Tomlin, 1937:217, (cf. 8,

"Hab. ?, Type in coll. Tomlin"; Coomans et a/., 1982:4, 9). Additionally, the periostracum in C. baccatus is yel-

fig. 197, apertural and dorsal views of the holotype; Vink, lowish tan, whereas in C. perplexus it is a brownish tan.

1984:354-358, pi. 20, fig, 1, dorsal view of holotype; Rock- The largest specimen of C. baccatus examined (AMNH el, 1985a:29, dorsal view of holotype and dorsal and aper- 221871a; figures 6, 7) is 26.9 mm in height, compared tural views of two recently collected specimens from "Pe- to a maximum of 37.8 mm for C. perplexus (AMNH rida Island ", Pacific Panama (figured in color). 206684) from Isla Cebaco, Golfo de Montijo, Panama. Original description (in part): "Shell short, rather swol- Hanna (1963:39) reports a specimen of the latter taxon len, very minutely decussated, with regular rows of con- from Isla San Jose, Golfo de Panama, with a height of spicuous granules, whitish, with large orange blotches 41.5 mm. Both of these taxa have shells with a distinct arranged in three bands; spire acute, short, whorls con- posterior notch on the lip (cf. figure 5 with figure 10),

cave, nearly smooth, last whorl biangulated. . . . Apart whereas C. xirnenes Gray, 1839, C. nmhogani Reeve, from its somewhat stunted form, the delicacy of its mark- 1843, and C. tornatus Sowerb\, 1833, which also have ings and rows of gem-like granules, it is remarkable for been confused with C. perplexus, have a narrow anal the double angle at the top of the body whorl. Long. 23, notch (see Wolfson, 1962: figs. 6a-d; Tucker 1985: fig. lat. maj. 15 mill " [Holotype actually measures 22.2 by c; and Chancy, 1987: figs. 2, 3). All of these nominal 14.2 mm.) species have populations sympatric with those of C. bac-

catus, which is known only from Pacific Panamic waters. Type depository: Holotype in the National Museum of Conus perplexus, with which C. baccatus is most likely Wales, no. 1955.158.29, fide Trew (1982:3); here illus- to be confused, ranges from the northern part of the Gulf trated (figures 1, 2). of California, south to Mancora, Peru, 4°05'S, 81°03.5'W Type locality: Off Isla Parida, Golfo de Chiriqui, Re- (LACMNH no. 70-101). Specimens of C. perplexus in piiblica de Panama, here designated. which the interrupted spiral lines of brown dashes or dots are raised into prominent pustules are more commonly Range: Known only from Pacific Panama. found in the southern populations, from Panama south- Material examined: ,\MNH Collection: From Golfo de ward to Peru (figures 15, 16). .\ similar pustulose form Chiritpii, Pacific Panama: 6 specimens, no. 173698 and also occurs in populations of the western .\tlantic C 8 specimens, no. 211778, off Isla Parida (8''05'N, 82°20'W), puncticulatus Hwass in Bruguiere, 1792, which is a twin W. K. Emerson and W. E. Sage III, 1988 Page 113

species of C. perplexus. This form was named C. pa- their observations on the differences in the nuclear whorls pillosus Kiener, 1849 (figures 17, 18). of Conus perplexus and the specimens which were found On the basis of shell morphology, C. baccatus appears to be referable to C. baccatus. Their interest and coop- to be most closely related to members of the western eration, past and present, are greatly appreciated. We Atlantic C. jaspideus species complex, especially the pus- acknowledge with thanks the assistance of Philippe tulose, short-spired Brazilian forms which were given the Bouchet, Museum National d'Histoire Naturelle, Paris; names C. yemanjae (figure 19) and C. selenae by Van Helen DuShane. Whittier, California; James Ernest, Bal- Mol et al. (1967: pi. 8); also figured by Vink (1984: pi. boa, Repiiblica de Panama; R. M. Filmer, Chobham, 20). England; Robert A. Foster, Santa Barbara, California; Conus perplexus Sowerby (1857:20, 1858: pi. 200, fig. Barbara Good, San Diego, California; James H. McLean, 324) was briefly described in Latin, followed by the com- Los Angeles County Museum of Natural History; P. Gra- ment; "This shell perplexes me, because there is a variety ham Oliver and Alison Trew, National Museum of Wales, of C. puncticulalus [Hwass in Bruguiere, 1792, a Carib- Cardiff; Donald R. Shasky, Redlands, California; Kay C. bean twin species] which it nearly resembles. This, how- Vaught, Scottsdale, Arizona; and K. M. Way, British Mu- History), London, for the loan of speci- ever, is quite smooth, and more angular' . A single spec- seum (Natural imen was figured by Sowerby in a dorsal view, with the mens and/or photographs, or for providing information. statement: "Gulf of California, West Columbia [sic], We also are indebted to Alan Kohn, University of Wash- Cum.[ing]". Tomlin (1937:290) recorded the presence of ington for providing data on the complicated nomen- three "types" in the British Museum (Naural History), clatural history of the taxa Conus bicolor and C. albo- which were kindh' loaned to us by Ms. K. M. Way maculatus. Stephanie Crooms, AMNH, word-processed [BM(NH) 19781 18^ here illustrated, figures 8. 9, 11-14]. the manuscript. The original labels confirm these specimens as syntypes, We owe a special debt of gratitude to Henry W. Cha- although none of them can be referred with certainty to ney, Santa Barbara Museum of Natural History; James

Sowerby s figured specimen. The drawing may represent H. McLean; and James Nybakken, Moss Landing Marine a composite. The three syntypes are somewhat faded, Laboratories, for critically reading drafts of the manu- but are otherwise well-preserved. We have selected as script. the lectotype (figures 8, 9) the syntype that most closely resembles the original figure. The lectot\pe is slightly LITERATURE CITED larger in height (26.8 vs. 26.1 mm) than the specimen depicted in the original illustration. Born, I. von. 1778. Index rerum naturalium Musei Cesarei the lot of The original labels accompanying syntype Vindobonensis. Pars prima, Testacea. Vienna, xlii + 458 list of California and West Conus perplexus the Gulf p., 1 pl. Columbia [.sic], which are the same localities cited by Bruguiere, J,-G. 1792. Cone. In: Encyciopedie methodique. Sowerby (1858, caption to Conus, pi. 14). Hugh Cuming Historie Naturelle des Vers 1:586-757, illus. study of two similar is not believed to have collected in Mexican waters on Chaney, H. W. 1987. A comparative mahogani. The his voyage to the west coast of South America, 1828-30. Panamic cones: Conus ximenes and Conus Veliger 29(4):428-436, 6 figs. He is known, however, to have made extensive collections Coomans, H. E., R. G. Moolenbeek, and E. Wils. 1979. Al- in the "Gulf of Panama, the Pearl Islands, and the Gulf phabetical revision of the (sub)species in Recent Conidae of Chiriqui", as well as visits to Costa Rica, Nicaragua, 2. adansoni io atbuquerquei. Basteria 4;3(5-6):81-105, figs. the and Honduras (Dance, 1986:114). We here restrict 26-50. Golfo Panama, as type locality of C. perplexus to the de Coomans, H. E., R. G. Moolenbeek, and E. Wils. 1982. Al- the Isthmus of Panama was part of Colombia at that phabetical revision of the (sub)species in Recent Conidae.

time. 5. baccatus to byssinus, including Conus brettinghami In summary, Conus baccatus G. B. Sowerby III, 1877, nomen novum. Basteria 46(l-4):3-67, figs. 172-292. of shell collecting. E. Brill, is recognized as a Panamic faunal constituent and as a Dance, S. P. 1986. A history J. twin species of C. jaspideus Gmelin, 1791, from the Leiden, xv -I- 265 pp., illus. Gmelin, F. 1791. Systema naturae per regna tria naturae, western Atlantic, especially the pustulose form named J. 13th ed. 1(6), cl. 6, Vermes. Leipzig, p. .3021-3910. C. yemanjae Van Mol et al., 1967. Conus baccatus is Gray, E. 18.39. Molluscous animals and their shells. In: F. compared with pustulose specimens of the sympatric C. J. W. Beechey. The zoology of Capt Beechey's Voyage . . . perplexus G. B. Sowerby II, 1857, with which it has been to the Pacific and Behring's Straits in His Majesty's ship of C. punc- confused, as well as with pustulose specimens -I- 33-44. Blossom. . . . London, p. i-xii 103-155, pis. ticulalus Hwass in Bruguiere, 1792, a Caribbean twin Hanna, G D. 1963. West American mollusks of the genus species of C. perplexus. A lectotype is selected and the Conus— II. Occasional Papers of the California Academy

t\pe locality is restricted for C. perplexus. A type locality of Sciences 35:103 p., 4 text figs., 11 pis. Species general et ico- is designated for C. baccatus. Kiener, L.-C. 1846[-48], 1849[-.50]. nographie des coquilles vivantes. Vol. 2, Conus. Paris, p. ACKNOWLEDGEMENTS 1-379, 111 pis. [Conus alhomaculatus, 1846, pp. 56, 57, pl. 83, fig. 2; C. papillosus. 1849, p. 271, pl; 72, fig. 4] 1967. Resultats We are indebted to Carol Skoglund and Bob Koch, Phoe- Mol, J.-J. Van, B. Tursch, and M. Kempf. nix, Arizona, for sending us comparative material, for scientifiques des campagnes de la 'Calypso' 32. Campagne '

Page 114 THE NAUTILUS. \ol. 102, No 3

de la Calypso au large des cotes atlantiques de I'Amerique Dr Prevost. Proceedings of the Zoological Society of Lon- de Sud (1961-1962) 16. Mollusques Prosobranches; les don, for 1876, p. 752-755, pi. 75. [Issued Jan. 4, 1877] Conidae du Bresil. Annales de I'lnstitute Oceanographique Sowerby, G.B., III. 1887. /n.G. B. Sowerby II (ed.). Thesaurus Monaco 45(2):233-254, 6 pis. conchyliorum. Second supplement to Monograph of the

Reeve, L. 1843. Conchologia iconica, \ ol 1 London Conus. genus Conus, Vol. 5, pt. 44 London, p 249-279, pis. 507- London, 47 pis., with text. 512, Conus pis. 29-36 [Conus baccatus, p. 251, pi. 507,

Reeve, L. [184S-]1849. Conchologia iconica, \'ol. .5. Conus, fig. 660, Conus pi. 29. Issued July. 1887]

te.xt. le 1937. of Recent fossil cones. emendations, London, 7 p., 9 pis , with Tomlin, J R B. Catalogue and Rtickel, D. 1985a. Conus Imccatus Sowerby III, 1877, a for- Proceedings of the Malacological Societ) of London 22(4- gotten valid species from tropical west-America. La Con- 5):205-330.

chiglia, Rome, yr. 17, nos. 192-193, p. 29, 8 figs. Trew, A. 1982. The .Melvill-Tomlin Collection, part 10, Co- Rockel, D. 198.5b. Die Familie Conidae, nr. 22 [Contis lito- nacea (Conidae). Handlists of the Molluscan Collections

glyphus Hwass in Bruguiere, 1792]. Privately issued, 1 p., in the Department of Zoology, National Museum of Wales, 4 figs Cardiff, Ser. 1:1-28 + i-ix. Shaw, H. O. N. 1909. On the dates of issue of Sowerby s Tryon, G. W., Jr. 1883[-84]. Manual of concholog>, Ser. 1, "Conchological Illustrations." from the cop\ presented to Vol. 6, family Conidae, pp. 1-1.50, 31 pis. [Part 21, pp. 1- the Radcliffe Library. Oxford. Proceedings of the Mala- 64, Dec. 27, 1883]

cological SocietN of London 8(6):333-340. Tucker, J. K. 1985. Conus ximenes Gra\, 1839, Conus ma- Life Sowerby, G.B., I. 1833[-40], 1841. Conchological illustrations, hogany [sic] Reeve, 1843 and friends. Shells and Sea

Conus, pt. 24, fig. 2 [published March 29, 1833]; list for 17(8):201-202, figs, a-e, 1-5. Bayside, California.

Conus section, p. 1-4 [published in 1841]. London. [See Vink, D. L. N. 1984. On the identity of Conus anaglypticus, Shaw, 1909:340] Conus baccatus, Conus selenae and Conus yemanjae.

Sowerby, G. B., II. 1857-58. Thesaurus conchyliorum. Vol. Journal of Concholog\. London 31(6):353-.358, pi. 20. of living 3, pts. 17, 18. London, pp. 1-24, pis. 187-195, 196-210. Walls, J. G. 1979 Cone shells, a synopsis the Conidae. [Description of Conus perplexus, 1857, p. 20; 1858, pt. 18, T.F.H. Publications, Inc., Neptune Cit\. NJ, 1011 p, illus.

pis. 196-210; figure of C. perplexus, 1858, pi. 200, fig. Wolfson, F. H. 1962. Comparison of two similar species of 324, Conus pi 14] Conus (Gastropoda) from the Gulf of California. Part 1: Sowerby, G. B., III. 1877. Descriptions of six new species of a statistical analysis of some shell characters. The Veliger shells from the collections of the Marchioness Paulucci and 5(l):23-28, 6 figs. THE NAUTILUS 102(3):115-122, 1988 Page 115

References to Molluscan Taxa Introduced by Linnaeus in the Systema Naturae (1758, 1767)

Kenneth J. Boss Miiseuiii of Comparative Zoology Harvard liniversity Cambridge, MA 02138, USA

ABSTRACT tina: Teredo, Vermes: MoUusca: Clio, Limax, Doris, Te- thijs, Scyllaea, and Sepia; and Vermes: Testacea: (all Bibliographic documentation is provided for the more than 90 genera with the exception of Lepas and Serpula though published resources utilized by Linnaeus as citations for mol- references to the two molluscan Serpula are included)], lusks in the 10th and 12th editions of his Systema Naturae. the total exceeds 90. Dance (1967) also provided a short

list of volumes personally owned by Linnaeus which were INTRODUCTION utilized in various editions of the Systema. Unknown to any of us were the studies of Professor Linnaeus's 10th edition of the Systema Naturae of 1758 John L. Heller, Emeritus, of the Department of Classics

is the cornerstone of our system of binominal nomencla- of the University of Illinois on various botanical works ture in zoology. Many species of mollusks were described of Linnaeus (see Heller, 1983, for a collection of these therein, and these have been the subject of individual articles). He provided extensive bibliographic references investigations such as revisions of particular taxa as well to similarly abbreviated citations by Linnaeus in some as studies of the specimens themselves (Hanley, 1855; of his botanical works such as the Species Plantarum Dodge, 1952-59; Dance, 1967). and the Hortus Cliffortianus. He also explicated certain Linnaeus also attempted to apply the binominal meth- particularly equivocal abbreviations used by Linnaeus in od to the references that accompanied his treatment of referring to various insects (Heller, 1961) and planned the species. If he cited a reference that previously men- to finish a complete bibliographical guide to zoological tioned or figured the species under discussion, he usually works cited by Linnaeus under the proposed title Bib- gave an abbreviated citation, which frequently embod- liotheca Zoologica Limmeana (Heller, 1968, 1979, 1983); ied a shortened name of the author and that of the in- this work is currently in press (Heller, personal com- dividual work. Many of these are readily apparent to munication) but will contain more than a malacologist someone with a knowledge of the pre-Linnaean litera- need know. ture; thus, "Gault. test." may be easily recognized as Although I am separately preparing a more extensive being a reference to Gaultieri's Index Testarum Con- treatment of the pre-Linnaean malacological citations chyliorum of 1742; others are more enigmatic. including comments on subsequent editions, translations

Some years ago I thought that it might be interesting into other languages as well as annotations of the holdings to decipher these puzzling references in Linnaeus's Sijs- of the Harvard College Library system (Boss, in prepa-

tema, both the 10th (1758) and 12th (1767) editions (the ration), I present below the bibliographic abbreviations

nth is a reprint of the 10th), much like Wheeler (1979) utilized by Linnaeus in ihe Systema Naturae (1758, 1767) has done for fishes. However, in correspondence with and the full citation for each reference. Of all the ref-

the late Professor A. Myra Keen, I found that both she erences, only one remains undeciphered: Linnaeus (1758:

and Dr. S. S. Berry also had an interest in this problem. 708, 1767:1161) referred to "Tessin. epist. 1 n. 28 Cym- Professor Keen sent me a short draft of some 30 titles bium" under Argonauta argo. This is presumed to have that she had connected with Linnaeus's abbreviations, been a letter of Count Carl Gustaf Tessin, on whose

and I supplied her with several that she had been unable collection Linnaeus worked. Despite a thorough check- to find. Eventually, she published a narrative account of ing of apposite sources (i.e., items in the Literature Cited certain authors whom Linnaeus had cited (see Keen, as well as subsequent editions of the Systema and several

1983a, b). In these papers, she noted that Linnaeus used classical cephalopod authorities such as Chemnitz, Ke- about 40 different references in his sections on mollusks. ferstein, Conrad, Naef, Robson, Tryon, and Ferussac- However, when the different citations are tallied for both Orbigny), this reference could not be located. soft bodied and shelled mollusks [including therefore There are several bibliographic sources for pre-Lin- Linnaeus's categories with their genera, Vermes: Intes- naean works, and sometimes there are discrepancies in Page 116 THE NAUTILUS, Vol. 102, No. 3

citations of the sources. These may arise due to different Chin. Lagerstr. (see Odhelius, 1754) bibliographic methods, through error, or due to the vari- Column, aqu. or aquat. (see Colonna, 1606) ability in the copies of pre-Linnaean works, which were Column, phytob. (see Colonna, 1592)

not always issued in uniform copies I have relied prin- (Column, purp. (see Colonna, 1616)

cipally on the Catalogue of the Books . . . British Mu- Crew. mus. (see Grew, 1681) seum .... Soulsby (1933), Engelmann (1846), Heimann E.N.C. (see Francus de Frankenau, 1727) (1957), Nissen (1969), and the resources of \arious Har- Edw. av. (see Edwards, 1758-64) vard libraries. The birth and death dates of authors have Ellis cor. or corall. (see Ellis, 1755) been included, mainly as listed in the British Museum En. svec. (see Linnaeus, 1746) Catalogue; these do not always concur with such dates Faun. svec. or En. svec. (see Linnaeus, 1746) given by other authors, principally Dance (1986) or Nis- Frisch ins. (see Frisch, 1730)

sen (1969). I have utilized my own method of punctua- Gesn. aqu. or aquat. (see Gesner, 1551-87) tion and have spelled out such words as "engraved" and Ginam. adr. (see Ginnani, 1755-57)

"portraits" when describing the texts; I have followed Ginan. or Ginan. adr. (see Ginanni, 1755-57) standard procedure for the usage of brackets, etc., as Gmelin act. petrop. vol. 3, p. 246 (see Gmelin, 1729)

noted in the Catalogue of the Books . . . British Museum. Gnalt. test, (see Gualtieri, 1742) Further, the names of publishers or printers were added Grev. mus. or Grew. mus. (see Grew, 1681)

when 1 knew them. With few exceptions, which are listed Gron. lap. (see Gronovius, 1740) as "not seen", examination oi the original publication Gualt. test, (see Gualtieri, 1742) was possible. Hasselq. or Hasselqv. itin. (see Hasselquist, 1757) Hasselqv. act. ups. 1750. p. 33. (see Hasselquist, 1750) Hevde .Ant. Anatome mvtulorum 1683 oct (see Heide, ABBREVIATED CITATIONS UTILIZED 1683) BY LINNAEUS FOR MOLLUSKS Imperat. nat. (see Imperato, 1599)

It. gotl. (see Linnaeus, 1747)

Act. angl. 301, p. 2051 (see Breyn, 1705) It. oel. or oeland (see Linnaeus, 1745)

Act. helv. 4. p. 212, t.9, f.21, 22 (see Hofer, 1760) It. wgot. or wgoth. or wogth. (see Linnaeus, 1747)

Act. helv. 5. p. 283, n.4, t.3, f.25, 26 or n.5 t.3 f. 27, 28 Johnst. aquat. or exsangu. (see Jonstonus, 1650-53) (see Schlotterbeccius, 1762) Jonst. aquat. (see Jonstonus, 1650-53)

.\ct. paris. 1710, p. 463 (see Reaumur, 1710) Kaehl. or Kaehler act. Stockhb. 1754. p. 144, t.3, f.A-E

Act. paris. 1711, t.3, f.4,5 (see Reaumur, 1711a) or A-F (see Kahler, 1754)

Act. paris. 1711, p. 199, t.6, f.5.7. or 9 (see Reaumur, Kirch, mus. (see Buonanni, 1709) 1711b) Klein or Klen. ostr. (see Klein, 1753)

Act. paris. 1712, p. 163 (see Reaumur, 1712) Kratz. or Kratzenst. Regenf. (see Regenfuss, 1758)

Act. Petropol. 7, p. 321, t.U, 12 (see Koelreuter, 1761) Labat. itin. (see Labat, 1722)

Act. Upsal. 2. p. 560, t.l52, f.4. idem or f.a. (see Bromell, Lederm. micr. (see Ledermueller, 1760-61) 1729) Lewenh. arcan. (see Leeuwenhoek, 1695) Adans. sen. or seneg. (see Adanson, 1757) List. angl. (see Lister, 1678) Aldr. exs. or exsang. or exangu. (see Aldrovandi, 1606) List. angl. app. [appendices] (see Lister, 1685, under Lis-

Aldr. ins. (see Aldrovandi, 1602) ter, 1678) Amoen. acad. (see Amoenitates Academiae) List, conch, or hist, (see Lister, 1685-92[-97])

Amoen. acad. 1, p.325 (see Balk, 1746) List. exer. 2 or exerc. 2 or exercit. 2 (see Lister, 1695) Amoen. acad. 3: 256 (see Odhelius, 1754) List, exercit. anat. 1 (see Lister, 1694) Argenv. conch, (see Dezallier d'Argenville, 1742) List, exercit. anat. 2 (see Lister, 1695)

Barr. ic. or rar. or Barrel, icon, or rar. (see Barrelierus, M.L.U. or Mus. L. U. (see Linnaeus, 1764) 1714) Mus. Ad. Fr. (see Linnaeus, 1754) Baster subs, (see Raster, 1759-65) Mus. Tess. or Tessin (see Linnaeus, 1753) Bell. aqu. or aquat. or Rellon. aqu. (see Belon, 1553) Needham microsc. (see Needham, 1745)

Blank, ins. (see Blankaart, 1688) Olear. mus. (see Olearius, 1674) Bocc. observ. (see Boccone, 1674) Osb. iter, or Osbeck. iter, (see Osbeck, 1757) Bohads. mar. (see Bohadsch, 1761) Pet. or Petiv. amb. or amboin. (see Petiver, 1713) Ronan. kirch. (see Buonanni, 1709) Pet. or Petiv. gaz. (see Petiver, 1764) Bonan. recr. or recreat. or Bonann. recr. (see Buonanni, Pet. mus. (see Petiver, 1695) 1681, 1684) Plane, conch, (see Plancus, 1739) Brad, natur. or Bradl. nat. or Bradl. natur. (see Bradley, Reaum. act. paris. 1712, p. 163 (see Reaumur, 1712) 1721) Regenf. conch, (see Regenfuss, 1758) Breyn. polyth. (see Breynius, 1732) Roes. ins. or in.sect. (see Roesel von Rosenhof, 1746-61) Brown, jam. (see Browne, 1756) Rond. aqu. or aquat. (see Rondeletius, 1554-55) Bruckm. cent. 2 epist. (see Brueckmann, 1743) Rond. or Rondel, pise, or test, (see Rondeletius, 1554- Calceol. mus. (see Cerutus and Chiocco, 1622) 55) K. J. Boss, 1988 Page 117

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a" curiosi delle opere della natura, &c. Roma, Varese, 4 Frisch, Johann Leonhard [1666-1743]. 1720-38. Beschrei-

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engraved half-titles, 4to. lichen Anmerckungen . . . von diesem . . . inlandischen

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Uppsala, p. 240-264. I wish to thank Dr. Elizabeth A. Shaw% Bibliographer Heller, John L. 1983. Studies in Linnaean method and no- Mrs. and Research Taxonomist in the Gray Herbarium, menclature. Marburger Schriften zur Medizingeschichte, Geraldine C. Kaye of the Fariow Herbarium Library, (Verlag Peter Lang, Frankfurt am Main, Bern, New York.

Ms. Barbara Callahan of the Harvard University Her- 7(IX):328 p. barium, and Eva Jonas, Dana Fisher, Roxane Coombs, Keen. Angeline Mvra. 1983a. (January). On Linnaeus" book- and Mina Brand of the library of the Museum of Com- shelf. The Festivus (San Diego Shell Club) 15(1):5-15. 24 figs. parative Zoology. Mr. Richard I. Johnson also made avail- (August Linnaeus' book- able rare malacological volumes, and valuable criticisms Keen. Angeline Myra. 1983b 30). On shelf [abstract]. .Annual Report of the Western Society of were offered by Mr. Richard E. Petit and two anonymous Malacologists 15:15. reviewers.

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Watson and Viney, Ld., Vol. 1. A-D (1903), p. viii, 1-500; synonymis, locis . . . Editio duodecima, reformata. Impen- Page 122 THE NAUTILUS, Vol. 102, No. 3

sis Direct. Laurentii Salvii: Holmiae, Tom. 1 Regnum (and publications more immediately relating thereto) pre- Animale. Pars II. Classis V-VI, Nomina Generica. Nomina served in the libraries of the British Museum (Bloomsbury) specierum propria: Nomina trivialia. Nomina trivialia and the British Museum (Natural History) (South Ken- Papiliomim and Phalaenarum Synonyma Termini Artis. sington). Second Edition Trustees of the British Museum,

Appendi.x Svnonvniorum. Addenda. Errata, p. 533-1327 London, p. xi, 246; .\ddenda and Corrigenda, 68 p., 7 [36]. plates. Nissen, Glaus. 1969. Die Zoologische Buchillustration. Ihre Wheeler, A. 1979. The sources of Linnaeus knowledge of

Bibliographie und Geschichte. Band I. Bibliographie. An- fishes. Svenska Linnesallskapets arsskrift. argang 1978. ton Hiersemann, Stuttgart, p. 666. Yearbook of the Swedish Linnaeus Society. Commemo- Soulsby, B. H. 1933. A catalogue of the works of Linnaeus rative Volume. Uppsala, p. 156-211. THE NAUTILUS 102(3):123-124, 1988 Page 123

Frederick Benjamin Isely: Biographical Sketch and Malacological Contributions

Mark E. Gordon Department of Zoology University of Arkansas Fayettevi'lle, AR 72701, USA

Frederick B. Isely was known primarily for his pioneer- endeavors were pursued during the summers of 1929 ing research in orthopteran ecology (mainly Acrididae and 1931 at the University of Chicago and University of and Tettigoniidae) and as an educator/administrator. Colorado, respectively. However, he contributed significantly to the basic knowl- His teaching career began with an appointment at edge of unionacean biology and ecology. Isely began his Franklin Rural School, Brown County, Kansas (1894- studies with freshwater mussels in the Chikaskia River 95). Following his graduation from Fairmount College, of north-central Oklahoma during 1906. These initial observations soon expanded to include aspects of unionacean ecology, growth, migration, and distributional survey of mussels within the Arkansas and Red river basins of eastern Oklahoma. In these endeavors, he was finan- cially aided through an appointment as a scientific assistant with the U.S. Bureau of Fisheries during the summers of 1910-13. As a result, five manuscripts were published between 1911 and 1931. Isely's innovative concepts for ecological investigations are reflected by his discussion in the 1914 growth paper of the use and value of replicate samples and of the problems encountered using mark/recapture techniques. One such problem was predation by small bo\s interested in collecting the brass tags used to mark the mussels. The distributional survey, although funded by and conducted for the U.S. Bureau of Fisheries, was delayed in publication due to govern-

ment reassessment of funding priorities—World War I. This manuscript was published independently by Isely after the war. His last malacological publication resulted from the recovery of one of the "lost" growth study specimens 15 years following the completion of that in- vestigation. Further evidence for the value of replication in research. Frederick was born June 20, 1873 of Swiss parents (Christian Isely and Elise Dubach) at Spring Grove farm near Fairview, Brown County, Kansas. His early edu- cation was gained locally, culminating with his graduation in 1894 from Hiawatha Academy (Hiawatha, Kan- sas). He entered Fairmount College (now Wichita State

University) in its founding year 1895. He was quite active in college activities (e.g., debate team, class president

1895-99) and sports (e.g., track, football) and was award-

in first graduating class of the ed a B.S. 1899, the 4-year Figure 1. Frederick B. Isely during his Trinity University college. In 1909, he earned an M.S. from the University years. Reproduced from a photograph given to the Department of Chicago and continued that summer with further study of Biology, Trinity University by Mrs. F. B. Isely (courtesy of at the Marine Biological Laboratory. Later academic H. D. Murray). Page 124 THE NAUTILUS, Vol. 102, No. 3

he assumed the principalship of Central School, Hia- quately labeled lots may also be referable to Isely. Thirty- watha (1899-1901) followed by stints as teacher of bi- nine specimens including seven species represent vouch- ology at Wichita High School (1901-06) and University ers for the 1914 growth paper. Fourteen detached brass Preparatory School, Tonka wa, Oklahoma (1906-12). tags are also present. The remainder of his collection is

Equipped \s ith his M.S., he served as professor of biolog\' from the 1925 distribution sur\e\. Little else of Isely's at Central College, Fayette, Missouri (1912-20) with so- malacological material, other than some papers in the journs during the summers of 1915-17 as instructor of archives of the Biology Department at Trinity University biology at the L'niversity of Missouri. His administrative (H. D. Murra\, personal communication), remain. career was initiated with an appointment as dean and A small malacological collection of his brother, Dwight

professor of biology at Ckilver-Stockton College, Canton, IseK , was redisco\ered at Wichita State L'niversity (Met- Missouri (1920-22) followed by his assumption of the calf and Distler, 1984). Further information pertaining same roles at Texas Women's College, Fort Worth (1922- to D. Isely may be obtained in Miner (1976) and Metcalf 31). He returned to a strictK teaching position as pro- and Distler (1984).

fessor of biology at Trinity Universit\, Wa.xahachie and I thank the following people for their kind assistance later San Antonio, Texas (1931-47) where he developed in providing data for this note: Mrs. Katherine Isely the majority of his orthopteran research. He also served McGuire and Drs. Douglas G. Alexander, Harle\ P. as department chairman from 1931 to 1946, retiring from Brown, Donald A. Distler, Duane Isely, Charles G. Lin- active teaching in 1946. In recognition of his achieve- coln (deceased), Harold D. Murra\-, Robert W. Pennak, ments. Trinity University awarded him the Sc.D. honoris William D. Shepard, and Gary D. Snell. causa upon his retirement. Isely married Mary E. Nickerson of Clearwater, Kan- LITERATURE CITED sas, on May 8, 1901. They had four children and he

appears to have influenced his famiK , as he did his stu- Alexander, G. 1949. Frederick B. Isely (1873-1947), Ento- dents; biologists continue in the famih to date. In spite mology News 60:29-30. of a heavy teaching load, administrative duties, and re- Geiser, S. W. 1949. Proceedings of the forty-fifth annual meeting of the American Society of Zoologists: 3. Memorial search, he was active in community organizations resolutions, Frederick B. Isely 1873-1947. Anatomical throughout his life and still managed time for personal Record 103:233-234. pleasures {e.g.. camping, gardening, keeping up with the Metcalf, A. L. and D. A. Distler. 1984. Gastropods collected latest episodes of Palooka and Little Orphan Annie). Joe from eastern Oklahoma by Dwight Isely in 1911. The Isely was a well-loved and enthusiastic instructor in ad- Nautilus 98:135-137. dition to being an active and respected scientist. He Miner, F. 1976, Dwight Isely 1887-1974 Journal of Economic received grants-in-aid for his orthopteran research from Entomology 69:289-290. the National Research Council (1935, 1936) and the Shepard, W. D. 1982. Rediscovery of a portion of the Isely American Philosophical Society (1937, 1941). He was a unionid collection. The Nautilus 96:8. fellow and founding member of the Oklahoma Academy of Science (secretar\ 1901-12) and the North Texas Bi- MALACOLOGICAL BIBLIOGRAPHY OF ological Society (president 1924-27); a fellow of AAAS FREDERICK B. ISELY and the Texas Academy of Science (president 1938); and a member of the Society of American Zoologists, Eco- 1911. Preliminary note of the ecolog> of the early juvenile life 20:77-80. logical Society of America, Entomological Society of of the Unionidae. Biological Bulletin Experimental stud> of the growth and migration of fresh- America, Kansas Academy of Science, Texas Entomo- 1914. water mussels. U.S. Bureau of Fisheries Document no. logical Society, and Texas State Teachers Association. 792, 26 p. Frederick B. Isely died December 30, 1947 still active 1914. Mussel streams of eastern Oklahoma. U.S. Bureau of in research. Additional information may be acquired Fisheries Economic Circular no. 9, 6 p from two memoria (Alexander, 1949; Geiser, 1949). 1925. The fresh-water mussel fauna of eastern Oklahoma. Pro- Recently, a remnant of Isely s malacological collection ceedings of the Oklahoma Academy of Science 4:43- was rediscovered at the Stovall Museum, University of 118. Oklahoma (Shepard, 1982). About 450 specimens of 1931. A fifteen vear growth record in fresh-water mussels. unionids, representing 37 species remain. Some inade- Ecology 12:616-619, THE NAUTILUS 102(3): 125-126, 1988 Page 125

Rediscovery of Planorhella magnifica (Pilsbry) in Southeastern North Carohna

William F. Adams Andrew G. Gerberich Environmental Resources Branch Division of Fishes U.S. Armv Corps of Engineers National Museum of Natural History P.O. Bo.x 1890 Smithsonian Institution Wilmington, NC 28402, USA Washington, DC 20560, USA

Early in this century, Planorbis magnificus was de- erably in size. The maximum shell diameters of the spec- scribed by Pilsbry (1903) from the lower Cape Fear River imens collected alive are 35.5, 21.5, and 16.3 mm. Vouch- region of North Carolina. The type localitv for the species er specimens from our collection have been deposited in was given simply as "lower Cape Fear River". Bartsch the collections of the North Carolina State Museum of (1908) correctly surmised that the species is an inhabitant Natural History (NCSM P468-P471) and the National of lentic environments and found it living in Greenfield Museum of Natural History (USNM 857935). Because of Lake, a manmade impoundment lying to the south of restricted access, additional trips to Orton Pond have not what were then the city limits of Wilmington, NC, as been made. well as in Orton Pond, then called Sprunt's Pond, ap- Live specimens of Planorhella magnifica were found proximately 16 km south of Wilmington. Baker (1945) on the stems and undersides of the floating leaves of figured and studied portions of the anatomy of specimens Spatterdock, Niiphar liiteum (Sibthorp & Smith, 1809), collected b\' Bartsch and concluded that P. magnificus and Fragrant Waterlily, Nymphaea odorata (Aiton, 1789). should be reassigned to Planorhella. This ta.xon repre- Water depth where living specimens were collected was sents the largest known planorbid. approximately 1 meter, and the bottom substrate was Planorhella magnifica has not been reported since the organic. Orton Pond closely matches Bartsch's (1908) accounts b\ Pilsbry and Bartsch and has been considered description of habitat suitable to P. magnifica in Green- extinct by some (Opler, 1976; Imlay, 1977; Palmer, 1985). field Lake.

Fuller (1977) hypothesized that the species may still sur- Like Greenfield Lake, Orton Pond is a manmade im- vive in Orton Pond, a manmade pond similar to Green- poundment. Both were created early in the last century field Lake in age and physiography, based on an obser- to serve as a fresh-water source for rice agriculture. Orton vation of large planorbid egg masses there by J. P. E. Pond exemplifies a type of lentic waterbody unique in Morrison. southeastern North Carolina. Although blackwater lakes The Greenfield Lake watershed has been almost totally and ponds in the region typically are acidic, Orton Pond's developed since Planorhella magnifica was described, waters have a circumneutral pH, ranging from 6.2 and now undergoes intensive management for control of through 7.9 (Smock and Lenat, 1978). We suspect that nuisance algae. Management measures include appli- Orton Pond may have a direct connection to waters of cation of algicides during the growing season and oc- the Castle Hayne aquifer, an Eocene limestone deposit. casional drawdown during winter months. Fuller (1977) The molluscan fauna of Lake Waccammaw (Columbus mentioned fruitless attempts by himself and others to County, NC), the only other sizable circumneutral-pH find Planorhella magnifica in Greenfield Lake. Our in- waterbody in the region, has been investigated frequent- vestigations of that lake indicate that it no longer provides ly (Fuller et al, 1976; Porter, 1985), but Planorhella a suitable habitat for this species. magnifica has not been found in that system. Lake Wac- During July 1986, Planorhella magnifica was seren- camavv differs from Orton Pond in being much larger, dipitously rediscovered in Orton Pond, Brunswick Coun- a natural waterbody, and is not a blackwater system. ty, NC, while collecting water samples. Three living The owners of Orton Pond undertake little manage- specimens were obtained in 20 minutes. Four empty ment of the pond and manage the lands that surround shells were also collected from the drift line on the shore. it for timber and wildlife. With a continuation of this During another water-sampling trip in January 1987, an type of management, the near future for Planorhella attempt was made to collect live specimens for captive magnifica seems secure. However, not all of the pond s propagation. This effort was unsuccessful as much of the watershed is protected in this way. Therefore, potentially aquatic vegetation had died back. However, two more adverse developments could occur quite some distance empty shells were found washed up on the shore. from the pond. Our samples of Planorhella magnifica vary consid- Because of the apparent uniqueness of the waters of Page 126 THE NAUTILUS, Vol. 102, No. 3

LITERATURE CITED

Baker, F. C. 1945. The molluscan family Planorbidae. Uni-

versity of Illinois Press, Urbana, p. 1-530. Bartsch, P. 1908. Notes on the fresh-water mollusk Planorbis magnificus and descriptions of two new forms of the same genus from the southern states. Proceedings of the United States National Museum 33:697-700.

Fuller, S. L. H. 1977 Freshwater and terrestrial mollusks. /n:

Cooper. J. E . S S Robinson, and J B Funderburg (eds. ). Endangered and threatened plants and animals of North Carolina. North Carolina State Museum of Natural His-

tory, Raleigh, p. 143-194.

Fuller, S. L. H., M. J. Imlay, and J. D. Williams. 1976. En- dangered or threatened fresh-water mussels (Mollusca: Bivalvia; Unionidae) of the Waccamaw River basin of the Carolinas. Association of Southeastern Biologists Bulletin 23:60. Imlay, M. 1977. for survival Figure 1 . Shells of Planorbella magnifica collected alive from J. Competing Water Spectrum Orton Pond, Brunswick County, North Carolina, July 1986 9:7-14. (apprcximately natural size). Opler, P. A. 1976. The parade of passing species: a survey of extinctions in the U.S. Science Teacher 43(9):30-34.

Palmer, S, 1985. Some extinct molluscs of the U.S.A. .Atala Orton Pond and absence of Planorbella magnifica in 13(l):l-7.

Greenfield Lake and Lake W'accamaw, it appears that Pilsbry, H. A. 1903. The greatest American Planorlns. The 17:75-76. the species is extremely restricted in distribution and can Nautilus Porter, H. 1985. Rare and endangered fauna of Lake Wac- be found today only in Orton Pond. We believe that any J. camaw, North Carolina watershed s\ stem, \'ols. 1 and 2. changes in the quality of the surface waters in Orton North Carolina Endangered Species Restoration: Job Title Pond's drainage basin or in its underlying groundwater No. VI-7. North Carolina Wildlife Resources Commission, regime may have a deleterious effect on P. magnifica. Raleigh, p. 1-358. Due to the uncertainty of the future of the pond, an in- Smock, L. A. and D. R. Lenat. 1978. Preliminary stud\ of depth study of the autecology of the species is needed the effects of salt water intrusion on the macrobenthic so that potential conservation measures can be developed community of Orton Pond, North Carolina. U.S. Army and implemented should they prove necessary. Engineer District, Wilmington, p. 1-48. THE NAUTILUS 102(3):127-128, 1988 Page 127

Bite by Octopus joubini: A Case Report

Donald M. McKinstry Division of Science, Engineering, and Technology The Pennsylvania State University at Erie The Behrend College Erie, PA 16563-1200, USA

While octopuses are capable of delivering toxic bites, shiny, hard circular scar, 1 mm in diameter, was still human case reports are scarce (Halstead, 1978). This evident by 1 year. While in excellent health, Patrick is report describes a bite by Joubin's octopus (Octopus jou- allergic to hymenopteran venoms and to penicillin. How- bini Robson, 1929) in a 12-year-old boy. The incident ever, this was his first exposure to octopus venom, thus, occurred on Sanibel Island, Florida, on January 2, 1987 an allergic reaction to explain the bite effects was dis- at about 0900. A storm had deposited a variety of marine counted. Secondary bacterial infection could have con- animals on shore from the Gulf of Mexico. Not realizing tributed to the effects experienced, however, signs of that octopuses are venomous, the boy, Patrick Reed infection, e.g., pus, lymphangitis, and fever, were not McKinstry, picked up a small specimen and was prompt- observed. ly bitten on the dorsal surface of the right thumb prox- The octopus died after about 2 hours in a pail of sea imal to the interphalangeal joint. He pulled the octopus water and was preserved in 70% isopropyl alcohol. It was away from his thumb within 10 seconds. The actual bite, identified as a nearly reproductively mature male spec- described as a "sharp pinch", was quickly followed by imen of O. joubini by Dr. Ronald Toll. This species, the a burning sensation of greater intensit% than a bee-sting. smallest of the western Atlantic octopuses, ranges from A tiny red spot surrounded by a white circular area 6 southern Florida to the Bahamas and the West Indies. mm in diameter was then noticed. Bleeding was minimal, Specimens are often washed ashore by storms on the Gulf a few small drops. Over the next se\ eral hours the thumb coast of Florida (Meinkoth, 1981). The dorsal surface of

became moderately swollen and felt sore, stiff, and, at the specimen is deep tan, while the arms and ventral

times, somewhat numb. These symptoms, to a lesser de- surface are light tan. The mantle is 3.2 cm in length. I

gree, extended to the wrist. Medical attention was not have donated this specimen to Dr. Toll s collection. sought because only local effects, which gradually di- Two accounts of envenomation following the bite of minished by day one, were noticed. By day three, the O. joubini were found. In the first (Anonymous, 1965), thumb was slightly swollen, reddened, and sore, and a a physician was bitten on the finger while handling a

tiny hole was noticed at the site. Over the next several specimen washed ashore on Sanibel Island, Florida. He days, the thumb became itchy and was treated with 3% experienced a "sharp" bite with a little bleeding, quickly hydrogen peroxide solution. This was the first treatment followed by '"the most excruciating pain which rapidly of the bite. By day eight, the swelling and redness had became almost intolerable." Within a few hours, the disappeared but some soreness in the metacarpopha- finger became very swollen, hard, and deep red. By the langeal joint was experienced. The hole had enlarged to next day, the inflammation and pain had extended to 2 mm in diameter and displayed a slightly irregular the hand. Later, some white discharge was noted from border. During the period up to day 12, the hole became the bite wound. The swelling and hardness gradually darker and a red circular area of up to 7 mm in diameter subsided over a period of 2 months. The only systemic

developed around it. Itching continued but soreness in effect was a mildly elevated temperature. He noted that the joint area decreased. On day 12, a small amount of "there was never any evidence of bacterial invasion or ' clear yellow fluid oozed from the hole. The area was infection, only an extreme degree of cellulitis. then treated with hydrogen peroxide solution and anti- The second case (Wittich, 1968), occurred on a re- biotic ointment. The signs and symptoms gradually di- search vessel off Egmont Key, Florida. A Florida Board minished, and by day 32 the hole was replaced by a of Conservation worker was bitten on the back of the slightly indented circular scar 1 mm in diameter, sur- hand while sorting marine specimens. In this case, '"a rounded by a slightly darkened firm area 6 mm in di- sharp, piercing sensation " was felt, and a "severe pain" ameter. By da\- 100, onK' the indented scar was evident extended to the upper arm. The bite wound bled slightK

and the wound was considered completeK healed. .A and almost instantK became surrounded by a "'pure white Page 128 THE NAUTILUS, Vol. 102, No. 3

welt of about 25 mm diameter." In the following hour, Sewanee, TN; Sherman A. Minton, Jr , Indiana Univer-

the victim experienced pain and sw elling around the bite, sit\ , School of Medicine, Indianapolis, IN; Harry N. Cun- accompanied by nausea, headache, and fever. Improve- ningham, Jr., Kath\' Mauro, and Robert Rose, all of The ment gradually occurred, but after 3 days the area was Pennsylvania State University at Erie, The Behrend Col-

still sensitive and swollen. The swelling persisted, and lege, Erie, PA. A special thanks is due Patrick R. Mc- some itching and a serous discharge from the wound Kinstrv for his account of the octopus bite.

occurred. After 1 month, a 2 mm wound surrounded by a 6 mm red periphery was present. The wound then LITERATURE CITED healed quickK.

In the case I report, the bite effects were not as severe .\nonymous. 1965. Florida octopus bite. Sea Secrets 9(7):5. as those noted above even though the victim was a child Halstead, B. W 1978. Poisonous and venomous marine animals of the world The Darwin Press, Princeton, NJ, with less tissue mass to dilute the venom. However, the 1043 amount of venom delivered could have been minimal. p. Meinkoth, N. A. 1981. The Audubon Society field guide to

North American seashore creatures. .\if red .\. Knopf, New

ACKNOWLEDGEMENTS York, NY, 799 p. Wittich, A. C. 1968. Account of an octopus bite. Florida following for their helpful assistance I am grateful to the Academy of Science Quarterly Journal 29l4):265-266. and suggestions: Ronald Toll, The L'niversity of the South, THE NAUTILUS 102(3):129, 1988 Page 129

Occurrence of Mites in Mexican Land Snails

0§car J. Polaco Wolfgang Mendl Departamento de Prehistoria Lab. Acarologia Institute Nacional de Antropologia Departamento Zoologia e Historia Escuela Nacional de Moneda #16 Ciencias Biologieas Mexico 06060, Mexico Apartado Postal 42186 Mexico 11340, Mexico

To date there has only been a single report (Baker, 1945) with the exception of a single specimen of Helix aspersa of mites infecting the mantle cavity of a land snail from (MiJller, 1774), an introduced species collected in Mexico Mexico (Helix pomatia Linne, 1758, an introduced City, which was infected with two specimens of E. vox- species). The mite was identified as Riccardoella oude- encollinus. As no mites were found in any Mexican land mansi (Thor, 1932), however, the mite shown by Baker snails from the above families, the family Bulimulidae

(1945) is similar to the figure published by Fain and Van remains the only new record of a host for mites. Goethem (1986) for Riccardoella limacum (Schrank, The examined snails are deposited in the Mollusk Col- 1776) (Prostigmata: Ereynetidae). This mite has been lection of the Departamento de Prehistoria, I.N.A.H., studied by several European authors (Turk & Phillips, and the mites in the Acarology Collection of the Labo- 1946; Baker, 1970) and reported as cosmopolitan. ratorio de Acarologia of the E.N.C.B. In this note, we report findings of mites parasitizing We acknowledge two anonymous reviewers for their the mantle cavities of two species of land snails of the critical comments to improve this paper. famiK Bulimulidae and one species referable to the family Helicidae. The bulimulid Bidimulus unicolor (Sow- erby, 1833), collected in Tomas Garrido, in the state of LITERATURE CITED Quintana Roo, was parasitized by Riccardoella limacum. with one protonymph and one deutonvmph collected Baker, E. W. 1945. Five mites of the family Ereynetidae from nearly 50 specimens of the snail. Of 79 specimens from Mexico. Journal of the Washington Academy of Sci- of Rhabdotus alternatus (Say, 1830), another bulimulid ences .35:16-19. collected in Xicotencatl in the state of Tamaulipas, that Baker, R. A. 1970, The food of Riccardoella limacuni (Schrank) (Acari: Trombidiformes) and its relationships with pul- were examined, one contained three mites of the genus monate mollusks. Journal of Natural History 4:521-530. Boydaia (Ereynetidae) and another contained a single Fain, A. and L, VanGoethem. 1986. Les acariens du genre specimen of Eupodes voxencollinus (Sig Thor, 1934). J. Riccardoella Berlese, 1923, parasites du poumon de Mol- This is the first report of the last two mite species on lusques gasteropodes pulmones terrestres. Acarologia 27(2): land snails. 125-140.

Examination of other land snail species of the families Turk, F. A. and S. M. Phillips. 1946. A monograph of the Helicidae, Achatinidae, Succinidae, Oleanicidae, and slug mite Riccardoella limacum (Schrank). Proceedings of Polygyridae revealed no evidence of parasitism by mites, the Zoological Society of London 115:448-472. THE NAUTILUS 102{3):130, 1988 Page 130

Axelella, New Name for Olssonella Petit, 1970, a Preoccupied Taxon (Mollusca: Cancellariacea)

Richard E. Petit Research Associate Department of Invertebrate Zoology National Museum of Natural History Smithsonian Institution Washington, DC 20560, USA

The cancellariid genus Olssonella Petit, 1970, was pro- Museum National d'Histoire Naturelle, Paris, for calling posed for a rather compact and well-defined group of the prior use of Olssonella to m\ attention. species from the Later Tertiary and Recent faunas of the Americas. An Eastern .Atlantic species was subse- LITERATURE CITED quently placed in this genus by Bouchet and Waren (1985;263), and the morpholog\ of the type species was Bouchet, P. and A. Waren. 1985. Revision of the Northeast published b\ Harasew\ch and Petit (1984). The ta.xon Atlantic bathval and ab\ssal Neogastropoda excluding Turridae (Mollusca: Gastropoda). Bollettino Malacologico, has been used b\ a number oi other authors since it was Supplemento 1:121-296 proposed. Unfortunately, it has just been noted that Dall, W. H. 1888. Mollusks. In: Alexander Agassiz, Three Olssonella is preoccupied, and a replacement name is Cruises of the United States Coast and Geodetic Survey here proposed. Steamer Blake 2:62-75. Glibert, M. and L. Van de Poel. 1967, Les Bivalvia fossiles Axelella, new name for Olssonella Petit, 1970:83. du Cenozoique etranger des collections de I'lnstitut Royal Not Olssonella Glibert & \'an de Poel, 1967:121. des Sciences Naturelles de Belgique. \". Oligodontina. In- stitut Royal des Sciences Naturelles de Belgique, Memoires Type species, by original designation of Olssonella (2nd ser.') 83:1-152. Petit, 1970, Cancellaria smithii Dall, 1888, Recent, Harasew ych, M. G. and R. E. Petit. 1984 Notes on the mor- Western .\tlantic. phology of Olssonella smithii (Gastropoda: Cancellari- late A.xel \. Olsson, as did This new name honors the idae). The Nautilus 98(l):37-44. the name it replaces, and the name which makes the Petit, R. E. 1970. Notes on Cancellariidae (Mollusca: Gastrop-

replacement necessary. oda) — II. Tulane Studies in Geologv and Paleontology 8(2): Appreciation is expressed to Dr. Philippe Bouchet, 83-88. INSTRUCTIONS TO AUTHORS

THE NAUTILUS publishes papers on all aspects of the All line drawings must be in black, high quality ink, biology and s\stematics of mollusks. Manuscripts de- clearly detailed and completely labeled. Photographs scribing original, unpublished research as well as re\ iew must be on glossy, high contrast paper. All figures are ... articles will be considered. Brief articles, not exceeding to be consecutively numbered (figs. 1, 2, 3, , NOT

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THE NAUTILUS

Volume 102, Number 4 December 21, 1988 ISSN 0028-1344

A quarterly devoted to malacology.

Marine Biological Laboratory LIBRARY

JAN 3 1989

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Mr. Richard F. Johnson Dr. Ruth D. Turner Department of Mollusks Department of Mollusks Museum of C^omparative Zoology Museum of Comparative Zoolog\ Harvard University Harvard University C:ambridge, MA 02138 Cambridge, MA 02138 THEt^NAUTILUS Volume 102, Number 4 December 21, 1988 ISSN 0028-1344 CONTENTS

Nidia S. Romer A new species of Alvania (Rissoidae) from the West Indian Donald R. Moore region 131

Charles N. D'Asaro

THE NAUTILUS 102(4):131-133, 1988 Page 131

A New Species of Alvania (Rissoidae) from the West Indian Region

Nidia S. Ronier Donald R. Moore

Rosenstit'l School of Marine and Atmospheric Science l'niversit> of Miami 4600 Bickenbacker Causeway Miami, FL 33149, USA

ABSTRACT ABBREVIATIONS

Alvania (Alvania) colombiana new species was found from 45 USNM = National Museum of Natural History, Smith- to 261 m (24 to 143 fms) depths off the Caribbean coast of sonian Institution. Mexico, Puerto Colombia to South Florida, the Gulf of and FSBS I = Florida Department of Natural Resources, Rissoa xanthias (Wat- Rico. Alvania colombiana differs from Marine Research Laboratory, St Petersburg. in having a protoconch with a single spiral zig-zag son, 1886), MCZ = Museum of Comparative Zoology, Harvard Uni- line of tiny nodules just above the suture, and by its smaller versity. shell size. Alvania colombiana also differs from Rissoa precip- UMML = University of Miami Marine Laboratory. itata (Dall, 1889), by its protoconch and size. CNMS = Colombian Natural Museum of Science, Na- tional University, Bogota INTRODUCTION

Exploration of the deep sea a century ago was the equiv- MATERIALS AND METHODS alent of the space age today. The famous "Challenger" All the specimens to be photographed in the SEM were expedition sampled two offshore stations in the tropical cleaned in 100% sodium hypochlorite for 2 minutes, then western Atlantic in which small mollusks were numerous. rinsed in distilled water twice. If the specimens still These were described by Watson (1886). Among the showed residual material (e.g., sand grains), they were Rissoidae were two species, Rissoa pyrrhias (Watson, sonicated for 60 seconds. Finally, the specimens were air 1886) and Rissoa xanthias (Watson, 1886), that were dried and put on a SEM stub with double-sided Scotch similar in size and shape. Dall (1889), reporting on the tape. "Blake" material, described two more species of Rissoa, Rissoa acuticostata and Rissoa precipitata. Rissoa acu- Dall ticostata was similar to Rissoa xanthias. and finally Alvania (Alvania) colombiana new species (1927) acknowledged that with more specimens it was (figures 1, 2) impossible to separate the material into two species. Ris- soa precipitata, on the other hand, is known principally Description: Shell 1.0 to 1.3 mm in length. Protoconch from the original description and illustration. multispiral, glossy, light brown, with 3'/2 whorls that are

There is another small western Atlantic rissoid which sculptured with 1 spiral zig-zag line of tiny nodules just at first glance appears similar to the illustration of Rissoa above the suture and 1 undulating line of the same nod- precipitata. However, there are a number of features ules in the middle of the whorl. Nodules irregularly ar- that differentiate them. Size is one: the small species has ranged over entire protoconch. Teleoconch of approxi- an average adult size of 1.2 mm while Rissoa precipitata mately 2 whorls, with numerous axial ribs. Adult shell (holotype) has an adult size of 3.92 mm. Protoconch is translucent, very light brown in color. the other: the small species has a projecting protoconch Axial ribs slightly curved, 26 ribs on the body whorl while Rissoa precipitata has a depressed protoconch. One fading out on the base of the shell. Body whorl with a of us (D.R.M) had obtained specimens of the small species narrow spiral groove just below the suture and another during the MAFLA (Mississippi, Alabama, and Florida) 7 to 9 grooves between the periphery of the final whorl study (1974 to 1975) in the eastern Gulf of Mexico. This and the base of the shell. Varix terminal with a narrow small species was first reported by one of us (N.S.R.) as extension forming outer lip of semilunate aperture. Some

Alvania sp. 1 (see literature cited; Rodriguez, 1983). specimens weakly umbilicate. Page 132 THE NAUTILUS, Vol. 102, No. 4

Figures I, 2. Alvania colomhiana new species. I. Hololype, USNM miriiber 8.59339, 1.2S mm in length. 2. Protoconch, 280 x. Figures 3, 4. Rissoa xanthias (Watson, 1886), from off Miami, in the junior author's collection, 2.26 mm in length. 4. Protoconch, 220 X.

Type locality: Off the west coast of Florida at 29°35'N, 859340; 2 from off the west coast of Florida at 28°24'N,

87°20'06"W, depth 107.3 m. 85°15'06"W, depth 164.6 m, FSBC I number 33113; and off the west coast of Florida at 29°49'30"N, 86°25'30"W, Holotype: USNM number 859339. Length, 1.28 mm. depth 82.3 m, FSBC 1 number 33114; 2 from off Puerto W'idtii, U.78 mm. Rico at 17°53'24"N, 66°35'10"W, depth 221 m, MCZ Paratypes: One from off the west coast of Florida at number 297220; and off Miami at 25°47'N, 80°01'30"W, 29°35'N, 87°20'06"W, depth 107.3 m, USNM number depth 137 m, MCZ number 297219; 2 from off Miami N. S. Romer and D. R. Moore 1988 Page 133

Alvania colombiana has been confused with Rissoa

precipitata (Dall, 1889) (figure 5). This was due to the fact that specimens of Rissoa precipitata were not available for comparison. The holotypc and only known spec-

imen, MCZ 7470, is a thin shell v\itli no terminal varix. In fact, the sculpture fades away almost completely on the last half whorl (except for the sub-sutural tubercles).

The protoconch is of the form seen in gastropods without a planktonic larval stage: large, rounded, unsculptured, and with a smooth transition from protoconch to teleo-

conch. The type locality is Yucatan Strait at a depth of

640 fathoms (not 670 as is given by Dall, 1889:280).

Another similar species is Rissoa xanthias (Watson,

1886) (figures 3 and 4). It, however, is about twice the size, has just over half as many axial ribs, and has a different and very distinctive protoconch. In this species the protoconch has 3'/2 whorls, in which the sutures are deep and ornamented with a line of spirally arranged

vertical pustules. Just above the suture there is a wide canal whose edges are formed by an undulating line with vertical pustules equally spaced and directed downward. Each canal also has in the middle dots and small nodules

randomly distributed. Watson reported it from Chal- lenger Sta. 24 off Culebra Island, 18°38'30"N, 65°05'30"W, Figure 5. Drawing of Alvania precipitata (Dall, 1889), ho- depth 715 m, and from Sta. 122, 9°10'S, 34°49'W to lotype, MCZ 7470, 3.92 mm in length. 34''53'W, depth 640 m.

Finally, Rissoa pyrrhias (Watson, 1886) is another sim-

ilar species. It is slightly larger than Rissoa xanthias, and at 25°47'N, 80°01'30"W, depth 137 m, UMML number its spiral grooves are not confined to the base of the shell. 8349; and off Miami at 25°46'30"N, 80°00'08"W, depth According to Watson (1886), the protoconch has 2'/2 whorls 76.81 m, UMML number 8350; 2 from off the Caribbean which are scored with coarse but feeble spiral threads. coast of Colombia at 09°52'35"N, 75°47'25"W, depth 72 m, CNMS number ICN-MHN(MO)522; and off Miami at 25°46'30"N, 80°00'08"W, depth 76.81 m, CNMS num- ACKNOWLEDGEMENTS ber ICN-MHN(MO)523. We are very thankful to Dr. Patricia Blackwelder and Distribution: This species was found in 31 samples from her assistant team at the SEM lab for their collaboration off the Caribbean coast of Colombia collected between in the use of and picture taking on the SEM. We also 1979 and 1981. The specimens were found at depths thank Dr. Kenneth J. Boss for the loan of the holotype ranging from 45 m to 261 m, but were most abundant of Rissoa precipitata. at depths between 65 m and 160 m. Other specimens of this species were found from off the west coast of Florida, LITERATURE CITED off southern Puerto Rico, and the Straits of Florida east of Miami. The bottom in almost all cases consisted of Dall, W. H. 1889. Reports on the results of dredging ... in sandy mud. The species is probabK' found in these depths the Gulf of Mexico (1877-78) and in the Caribbean Sea throughout the Caribbean, the southern and eastern Gulf (1879-80) by the U.S. Coast Survey Steamer "Blake" . . . of Mexico, and the Bahamas. All the specimens were 29. Report on the Moilusca, part II. Gastropoda and Sca- found dead. phopoda Bulletin of the Museum of Comparative Zoology 18:1-492. pis. 10-40. Discussion: Alvania colombiana appears to be a com- Dall, W. H. 1927 Small shells from dredgings off the southeast mon widespread species throughout the Caribbean and coast of the United States b\ the United States Fisheries adjacent areas. This is a very small compact species that Steamer "Albatross" in 1885 and 1886. Proceedings of the should not be confused with any other in its range. Some L'nited States National Museum 70(2667);1-134. Rodriguez, (Romer) N. S. 1983. Micromoluscos recientes de features, however, are lacking or lost on the protoconch la plataforma Caribe Colombiana entre Bocas de Ceniza of some specimens. These include the fine nodules dis- e Isla Fuerte. Thesis, Universidad Jorge Tadeo Lozano, tributed all over the protoconch, and the undulating line 247 p. of nodules on the periphery. These features are seen only Watson, R. B. 1886. Report on the Scaphopoda and Gaster- under high magnification, so are difficult to observe for opoda collected by H. M. S. Challenger during the years teleoconch sculp- routine identification. Details of the 1873-1876. Voyage of H. M. S. Challenger, Zoology 15: ture, however, should be easy to make out under low 1-675; appendix A, 677-680; geographical distribution, power, even in a somewhat worn specimen. 691-722; index, 723-756; pis. 1-50. —

THE NAUTILUS 102(4):134-148, 1988 Page 134

Micromorphology of Neogastropod Egg Capsules

Charles N. D'Asaro Department of Biology L iii\ersit\ of West Florida Pensacola, FL 32514, USA

ABSTRACT formed in the oviduct and ventral pedal gland. Nor is it

known if a common capsular microstructurc exists in Egg capsule micromorphology of eight species of neogastropods various neogastropod taxa. {Chicoreus florifer dileclu.s. PhijHonotus pomum. Cantharus Macromorphology of the often species-specific inuUangiilus, C. cancellariti.s. Conus floridanus floridensis, C. egg ju.spidcus stearnsi. Cranulina ovuliforniis. and Marginella au- capsules and egg masses produced by neogastropods has reocincta) was studied with light microscopy hy examining been studied more frequently than any other aspect of very thick, toluidine blue stained sections Laminae exposed neogastropod egg encapsulation. This was illustrated in hy fracturing the sections, and their reactions to the stain, the review of prosobranch reproduction by Webber provided the characters used to describe micromorphology. (1977), in a more recent review of egg encapsulation for results that buccinaceans The showed muricaceans and have all mollusks by Pechenik (1986), and in descriptive re- complex but microstructurally similar egg capsules, while co- ports on the external structure of neogastropod egg cap- nids and marginellids have egg capsules with taxonomically sules by Bandel (1975. 1976a,b,c, 1982) and D'Asaro distinct rnicrostructural characteristics. In the Muricacea and (1986a). Buccinacea, four structural laminae of similar origin and func- In early reports on neogastropod capsule-wall micro- tion are usually present in the capsule wall, the second from the outermost having the most complex pattern of fibers and morphology (e.g., .\nkel, 1937; Hancock, 1956), three or the greatest thickness. The third outermost lamina is continuous four laminae were described, including specific patterns with one or more components sealing the escape aperture. In of fibers. However, it was not established if these are some buccinaceans, only a trace of the third lamina exists in structural relationships common to the species studied the wall. Four layers, including one or two mucoid plugs, close or to higher taxa. Fretter (1941) provided for Nttcella capsules differ in that they the escape aperture. Conid egg lapillus (Linne, 1758), Ocenebra erinacea (Linne, 1758), include only three structural laminae in the capsule wall and Nassarius reticulatus (Linne, 1758), and Buccinum un- three layers, including a riuicoid plug, sealing the escape ap- datiim Linne, 1758 the most frequently cited descrip- erture. Microstructurc and fiber pattern in the middle lamina tions of how these neogastropods construct, in the oviduct are probably unique to this family. Egg capsules of marginellids and ventral pedal gland, three or four lay ered capsules. have a distinctive thin, dense wall not separated into laminae and lack an escape aperture closed by a mucoid plug. There To characterize the laminated egg capsule of Uro- is a preformed suture that fractures at hatching in the wall of salpinx cinerea follyensis B. Baker, 1951, Tamarin and most marginellid egg capsules. Carriker (1967) published the first comprehensive study light Ke\i words: Reproduction; egg capsules; Neogastropoda; Chi- employing microscopy — including polarized light coreus. Phyllonotiis. Cantharus, Conns, Cranulina: Margi- and electron microscopy. They described a capsule with ncllti four laminae composed of asymmetrical protein-like molecules bound into collagen-like matrices. They also noted that the great structural complexity of the capsular INTRODUCTION wall could only be partialh explained b\ Fretter's (1941) description of the formative process. Bayne (1968) con-

Encapsulation of early ontogenetic stages is typical of ducted a histochemical study of several gastropod egg higher gastropods, especially neogastropods. The highly capsules, including those of Nucella lapillus, and found refractory, layered envelopes of carbohydrate and pro- mucopolysaccharides in at least three capsular laminae tein are structurally and chemicalK complex (Hunt, 1971; of that species. He was one of the first to use toluidine Flower, 1973; Goldsmith et ai, 1978; Gruber, 1982; Sul- blue for this purpose. Flower et al. (1969) and Flower livan and Maugel, 1984). Encapsulation and the for- (1973) described the origin and the ultrastructure of neo- mation of egg masses provides protection and has con- gastropod capsular proteins. In another histochemical siderable survival value (Tamarin and C^arriker, 1967; study, (Jruber (1982) described six structural layers in Pechenik, 1979, 1983; Abe, 1983; Lord, 1986). Although the capsule wall of Eupleura caudata etterae B. B Baker, there are thousands of neogastropod species, it still has 1951, and three additional layers associated with tlie not been clearly established how the egg capsules are escape aperture and la\ ers of albumen. C. N. D'Asaro 1988 Page 135

Table 1. Comparative micromorphology of muricacean egg capsules stained in most cases with toluidine blue (meta = metachromatic, mp = mucoid plug, nr = not reported, ortho = orthochromatic, pre = present, tt = two types of albumen cells in the oviduct, z = zone). Page 136 THE NAUTILUS, Vol. 102, No. 4

tural lamina was first isolated and defined in a fractured or delaminated section, then the lamina could be rapidly

and comparably traced in whole, stained sections. If a capsule fractured and delaminated repeatedK' in a consistent pattern, then the parts were considered distinct structural laminae produced by different types of cells or produced at different times. Those less than a few micrometers wide could be recognized when they separated from the sections and folded to one side. Very thick sections also allowed microsculpture on the outer

laminar surface to be examined {e.g., figure 3). Following Tamarin and Carriker's (1967) method, structural laminae were identified from outermost to innermost as Ll- L4. Diagrams of median sections were prepared with a drawing tube mounted on a compwund microscope, while photomicrography was used to record microstructure at selected positions on a section Measurements of the capsule wall, structural laminae, and the sealed escape aperture were taken from the middle regions of each (tables 1-3). These dimensions should be considered approximations, as Sullivan and Mangel (1984) suggested, because of possible shrinkage introduced by the method and the wide range in width occurring at approximately the same position on different capsules of the same species.

Figure 1. Schematic drawing of a longitudinally sectioned egg capsule of Cbicurciis florifer dilectiis. Figure 2. RESULTS

Drawing, like figure 1, of a PhijUonoius pomum egg capsule. Chicoreus florifer dilectus (A. Adams, 1855) Numbers with arrowheads indicate positions at which photo- (figures 1, 3-8; table 1; USNM 860426) graphs for figures 3-12 were taken. Empty spaces in figures 1 2 are artifacts produced during dehydration. A = adhesive, and Egg capsules of this muricid are distinctly vasiform in DA = dense albumen, E = embryo, IMP = inner mucoid plug, section (figure 1). Macromorpholog\ was described by LI = metachromatic first lamina — solid line, L2 = second D'Asaro (1970:420, fig. 3). lamina — light stipple, L3 = metachromatic third lamina—solid The capsules have a thick, highK fibrous wall in which line, L4 = metachromatic fourth lamina —broken line, OMP four structurally distinct components can be identified: = outer mucoid plug, T.'\ = thin albumen. L2 is metachromatic thin, fineK fibered, outer lamina; in figure 1 and partially orthochromatic in figure 2. LI, a metachromatic L2, a metachromatic lamina, with layered, coarse fibers forming the bulk of the wall; L3, the thinnest meta- turcd and partially delaminated by varying the speed at chromatic lamina, closely applied to the innermost layer;

\\ liicli the microtome blade cut a section. and L4, a metachromatic lamina surrounding the al-

To demonstrate mucopolysaccharides, sections were bumen (figures 3, 4; table 1). LI is so transparent that

stained with toluidine blue for 1.5-2.0 minutes according the coarser L2 fibers can be seen through it. Except on

to the method of Gurr (1962:440). They were then rap- the stalk and basal plate, LI is folded in a corrugated id!) delnclrated in ethyl alcohol, cleared in toluene, and maruier (figures 3, 4, os). L2 may have fibers directed at mounted in methyl methacrylate. right angles or parallel to the long capsule axis and ar- Egg capsule microstructure was examined using light ranged with regional differences in two or three fused microscopy. Observation of the preliminary sections sug- layers. When three fibrous components are present, the gested that toluidine blue metachromasia would produce middle one has the coarser fibers arranged parallel to the a range in color from pur[)le through heliotrope (reddish long axis (figure 3, L2). Where the wall was folded during purple) but rarely red. Also, some layers would be or- formation, L2 fibers separate, forming large \acuoles

thochromatic (blue) or colorless. In the intact capsule (figure 4, v). These vacuoles are not stained and are wall, structural laminae were difficult to trace by follow- probably filled with a nonmucoid liquid. L2 fibers fuse ing metachromatic la\ers alone. However, when a struc- with a more or less homogeneous inner component in

Figures 3-8. Sections of the Chicoreus florifer dilectus egg capsule. 3. Wall showing laminae 4. Wall just below the apical plate. 5. .Albumen fibers. 6. Mucoid plugs in the escape aperture at the junction with 1.2 7. Wall in the stalk 8. Basal plate and adhesive. = Positions at which photographs were taken are indicated on figure 1. a = adhesive, da = dense albinnen, f = fracture, fl,3 I

C. N. D'Asaro 1988 Page 137

- * 'fly J -^ V w. .. V *;^ A*" I .

•ts %T:y ta m

50 u m • '— fragment of L3, fiL2 licmmm luuus component of L2, imp = inner mucoid plug, isz = intensely stained zone, LI = metachromatic first lamina, L2 = metachromatic second lamina, L3 = metachromatic third lamina, L4 = metachromatic fourth lamina, os = outer surface of LI, omp = outer mucoid plug, sL2 = outer portion of L2 in the stalk, ta = thin albumen. Page 138 THE NAUTILUS, Vol. 102, No. 4

Figures 9-12. Sections of the Phyllonotuti pnmum egg capsule. 9. Wall showing all laminae. 10. Escape aperture toward the base. 11. Escape aperture toward the ape.x 12. Stalk and basal plate. Positions at which photographs were taken are indicated on = figure 2. a = albumen, cf = circular fibers, e = embryo, hL2 = homogeneous component of L2, imp = inner mucoid plug, Li metachromatic first lamina, L2 = partially orthochromatic second lamina, L3 = metachromatic third lamina, L4 = metachromatic fourth lamina, omp = outer mucoid plug, os = outer surface of LI. sL2 = outer portion of L2 in the stalk, r = ridge. contact with L3 (figures 3, 4, hL2). L3 stains less inten- The apical escape aperture is closed by a structuralh sively than L4. In the apical plate, L3 appears to fuse complex barrier composed of four la\ers (figures 1. 6). with the inner mucoid plug and may have similar com- LI iscontiguouswith the outer layer. The metachromatic position, .^s described by Ankel (1929: fig. 1) for Nucella (purple) second layer or outer mucoid plug is larger than lapilhis and Franc (1940: fig. 2) for Ocinelmna aciculata the aperture and interdigitates with but is clearly sepa- (Lamarck, 1822), L4 completely surrounds the albumen rated from the homogeneous iimer component of L2 and probably contains it during assembly of the capsule (figure 6, omp, hL2). Fractures across the w idth of the while other components are more fluid. Hereafter, L4 outer mucoid plug apparently caused during sectioning will be called the albinnen retaining layer. occurred (figure 6, f). The third layer is a lenticular, Two structurally distinct types of albumen occur: an heliotrope mucoid plug, larger than the aperture, w hich outer layer including metachromatic fibers with random- sometimes fractures during sectioning. L3 is fu.sed to the ly attached granules in which all embryos arc situated lenticular or inner plug in an intensely stained zone (fig-

(figures I, TA; 5, ta) and, an inner core including denser ure 6, isz). L4 is the innermost layer closing the escape strongly metachromatic fibers (figures I, DA; 5, da). Fi- aperture and is fused to L3 in the same intenseK stained bers of dense albumen ma\ fuse in a continuous layer zone. where the albuminous components meet; however, the The stalk and basal plate are formed from the capsule fused layer did not delaminate during sectioning. The wall and an adhesive material (figures 1, A; 7, 8, a). The thin outer albumen does not envelop the denser core outer portion of both includes LI and up to the longi- basally (figure 1). tudinal, fibrous component of L2 (figure 7, sL2), while —

C. N. D'Asaro 1988 Page 139

the base of the capsular lumen is formed from the remaining parts of L2 as well as L3 and L4. Mucoid adhesi\e, which has orthochromatic and metachromatic zones and occasional L2 fibers, fills the center of the stalk

(figures 7, 8, a).

Phyllonotus pomtim (Gmelin, 1791)

(figures 2, 9-12; table 1; USNM 860425)

The somewhat tongue-shaped capsules and the rather massive, communal egg masses were described by Perr\ and Schwengel (1955: fig. 338), D'Asaro (1970:422, fig. 3), Radwin and Chamberlin (1973:107, fig. 1), Bandel

(1976a:10, fig. 4), and Moore and Sander (1978:253, fig.

2). Unlike Chicoreus florifer egg capsules, those of Phyl- lonotus pomum may have two points of attachment w ith a supporting substratum that nearly always is a conspecific capsule. Sections of such capsules are very variable in outline. Phyllonotus pomum capsules have a distinctly fibrous wall that is only partially metachromatic (table 1). Four structural parts e.\ist: LI, a thin, metachromatic outer lamina; L2, a thick, coarsely fibered and partially orthochromatic lamina; L3, a strongly metachromatic component; and L4, a metachromatic albumen retaining layer (figures, 2, 9). LI is as transparent as the homologous lamina in Chicoreus florifer dilectus and is corrugated on or near folds in the wall (figures 10, 11, os). L2 remains almost unstained. It has a core of longitudinal fibers bordered in most areas by a few circular fibers (figure 9, L2, cf). Each side of the fibrous zone may be bordered by a narrow, faintly orthochromatic, homogeneous zone Figure 13. Schematic drawing of a longitudinally sectioned (figure 9, hL2). L3 has no obvious fibers and appears egg capsule of Canlharus multanguliis. Figure 14. Drawing, fused to the outer mucoid plug of the escape aperture, like figure 13, of a Cantharus cancellarius egg capsule. Empty spaces in figures 13 and 14 are artifacts produced during de- as noted later. L4 has distinct circular fibers arranged in hydration. Numbers with arrowheads indicate positions at w hich a single layer. photographs for figures 15-23 were taken. A = adhesive, .\D Albumen, including fibrous and amorphous material, = apical depression, BM = basal mucoid material, D.A = dense is present in sections only near the retaining layer (figure albumen, E = embryo, LI = orthochromatic first lamina 10, a). This distribution is an artifact, as noted in Can- solid line, L2 = second lamina— light stipple, L4 = metachro- tharus cancellarius, caused by loss of the more fluid matic fourth lamina—broken Une, MP = mucoid plug, TA = contents during dehydration. Embryos are distributed in thin albumen both components of the albumen. Positioned on one side of the capsule, the sealed escape aperture includes four layers (figures 10; 11, LI, omp, sL2). Where the outer half of L2 was in contact with the imp, L4). The outer component is formed from LI. The substratum, a homogeneous zone having some fibers ex- second outermost layer, including intensely stained tending through it toward the substratum is present. The metachromatic regions, is continuous with but distinct inner half of L2, L3, and L4 form the floor of the capsular from the homogeneous proximal part of L2, and appears lumen. No adhesive layer was observed on capsules that to be fused or continuous with L3. This pattern is dif- had been attached to those of conspecifics. ferent from that of Chicoreus florifer dilectus. A third with small ves- layer, which is metachromatic, mucoid Cantharus multangulus (Philippi, 1848) icles, and exceedingly fragile, bulges into the capsular (figures 13, 15-19; table 2; USNM 947143) lumen, where it is bordered by L4, the albumen retaining layer. The more apical and lateral borders of the escape Macromorphology of the egg capsule was described by aperture, formed from L2, have an inward projecting Perrv and Schwengel (1955: fig. 340), Radwin and Cham- ridge (figure 11, r). berlin (1973:110, fig. 6), and D'Asaro (1986a:85, fig. 3).

The basal plate is formed from LI and the outer half Median sections through the ribbed and spined capsule of L2. A separation occurs in a dense reticulum or vac- are roughK vasiform in outline and have a rounded, uolated zone near the midline of the lamina (figure 12, apical depression partly covered by a transverse ridge Page 140 THE NAUTILUS, Vol. 102, No. 4

Figures 15-19. Sections of the Canlhartis mullai^gulus egg capsule 15. Wall showing laminae 16. Escape aperture adjacent to apical depression 17. Escape aperture on side opposite apical depression. 18. .Alliunien layers 19. Stalk and adhesive. Positions at which photographs were taken are indicated on figure 13. a = adhesive, ad = apical depression, bm = basal mucoid material, da = dense albumen, LI = orthochromatic first lamina, L2 = second lamina, L2m = metachromatic component, L2o = orthochromatic component, mp = mucoid plug, os = outer surface of LL ta = thin albumen. C. N. D'Asaro 1988 Page 141

Table 2. Comparative niicromorplinlogy of buccinacean egg capsules stained in most cases with toluidine blue (mcta = meta- c'hronialic, nip = mucoid plug, nr = not reported, ortho = orthoclironiatic, pre = present, tt = two types of albumen cells in the oviduct)

Layers Width closing Layers of Species of wall LI L2 L3 L4 aperture albumen Author

Cantharus ntul- 23 iim ortho, ortho and vestige meta, LI, L2, mp. langulus 1-2 /im meta, 1-2 /tm 21 ^m Cantharus cancel- 22 fim ortho, ortho and vestige meta,

larius 2 ^m meta, 3—i ^m 17 urn Btisycon carica nr pre pre pre pre and B. canalicu- latum Nassarius reticiila- nr three layers of uncertain relationship mentioned tus Buccinium undo- nr pre two fibrous mucoid nr turn layers layer

Ilyanassa obsoleta 11-22 meta pre, meta, 1 ^m pre, 60 nm (LO pre) Mm 2 nm 9-10 ^m Page 142 THE NAUTILUS, Vol. 102, No. 4

Figures 20-23. Sections of the Cantharus cancellarius egg capsule. 20. Wall showing laminae 21. Escape aperture opposite apical depression. 22. Escape aperture adjacent to apical depression. 23. Basal plate and adhesive. Positions at which photographs were taken are indicated on figure 14. a = adhesive, ad = apical depression, bm = basal mucoid material, da = dense albumen, LI = orthochromatic first lamina, L2 = second lamina, L2m = metachromatic component, L2o = orthochromatic component, mp = mucoid plug, OS = outer surface of LI, sL2 = outer portion of L2 in the stalk, ta = thin albumen.

laminae, homologous to three of the four muricacean albumen with fine fibers surrounds more coarsely fibered, laminae, are present. These are: LI, a very thin, ortho- less dense albumen in which embryos are positioned (fig- chromatic outer lamina; L2, an orthochromatic and ure 20, da, e, ta). metachromatic central lamina with tubular vacuoles; and The escape aperture is closed by four layers arranged L4, a metachromatic albumen retaining layer (figure 20; in a pattern essentially identical to that of C. multan- table 1). LI is often denser on the apical plate. Distinct gulus. Apically, LI forms the outermost layer (figures tubular vacuoles are present in the central portion of L2, 21, 22). The expanded, orthochromatic part of 1,2 is the especially where ridges e.xist in the wall and apical plate. second component. It has \ertical folds or fractures (fig-

Longitudinally directed vacuoles are positioned nearer ures 21; 22, L2o). The broadest component is a meta- to the outer surface, while more circularly directed vac- chromatic mucoid plug lying between the orthochro- uoles are nearer to the ituier surface. Apically, where the matic part of L2 and L4 (figures 21; 22, mp). This plug wall is folded, fibers in the lamina are visible (figure 21). tapers basally between L2 and L4, but does not extend

As shown for C multangulus, L3 is not present as a to the base as a continuous lamina equivalent to L3. distinct and continuous lamina throughout the whole There is a layer of similar mucoid material positioned capsule. The metachromatic albumen retaining layer is between L2 and L4 at the base of the lumen (figures 14, extremely thin and can be identified only where it is BM; 23, bm). pulled away from the albimien or the capsule wall and Basally, the plate is formed from LI and most of L2 folded to one side (figure 20, L4). Dense metachromatic (figures 14; 23, sL2). Only the orthochromatic, proximal C. N. D'Asaro 1988 Page 143

part of L2 extends across the base, where it, some mucoid material, and L4 form the floor of the lumen. Unlike most specimens of C. multangulus. the lumen is separated from the substratum b\ only a narrow layer of metachromatic adhesive (figure 23, a).

Contis jioridanus flohdensis Sowerby, 1870 (figures 24, 26-29; table 3; USNM 847141)

Median capsular sections typically have the longitudinal axis angling away from the axis of the stalk and show a slightly irregular outer surface (figure 24). Macromor- phology was described by Perry and Schwengel (1955: fig. 360, as C. spurius atlanticits Clench, 1942) and

D'Asaro (1986a:88, fig. 4). The capsule wall has three distinct components: LI, a fibrous metachromatic outer lamina; L2, a finely fibered, orthochromatic central lamina; and L4, a thin, metachromatic albumen retaining layer (figure 26; table 3).

LI is very transparent and has fibers circular in cross section. Occasional corrugations mark this lamina, especially near the escape aperture and external ridges (figure 27, LI). Fibers in L2 tend to form a cross-hatched pattern in the upper three quarters of the capsule, but some align with the long axis in the stalk and the base (figures 26-29). Globular structures buried in the fibrous Figure 24-. Schematic drawing of a longitudinally sectioned layer occur frequently (figure 27). No homologue of L3 egg capsule of Conus floridanus floridensis. Figure is present as a distinct layer, nor is there evidence that 25. Drawing, like figure 24, of a Conus jaspideus stearnsi the mucoid plug in the escape aperture is homologous egg capsule. Numbers with arrowheads indicate positions at to L3. The albumen retaining layer (L4) is not obviously which photographs for figures 26-32 were taken. \ = adhesive, fibrous. Although this layer is the third in sequence, it is .\L = albumen, E = embryo, LI = metachromatic first lamina designated as L4, the albumen retaining layer, because — solid line, L2 = second lamina— light stipple, L4 = meta- its position and probable function appear the same as L4 chromatic albumen retaining layer—broken line, MP = mucoid in Vluricacea and Buccinacea. plug. in this species, only one layer of fibrous, metachromatic albumen is visible (figure 26, al). Granules are In this plane, the basal plate is the widest part (figure attached to the widely separated fibers. Embryos are 25). randomly positioned in the albumen. Three laminae were identified: LI, a finely fibered, The sealed escape aperture is less complex than similar metachromatic outer lamina; L2, a complex orthochro- structures formed by muricaceans or buccinaceans. It is matic and metachromatic lamina with fibrous and mu- lined on its outer surface by LI and on its inner surface coid components; and L4, a metachromatic albumen re- by L4. L2 is replaced by metachromatic, layered mucoid taining layer (figure 30; table 3). LI, with circular fibers, plug with which it interdigitates extensively (figure 27, is tightly fused to the fibrous portion of L2. Except near mp). The degree of interdigitation suggests that L2 and the escape aperture and on the stalk, this layer is uni- the mucoid plug were formed at the same time. formly corrugated (figures 30; 31, os). L2 has an ortho- The stalk is composed of LI and almost all of L2 from chromatic outer component with coarse, circular fibers both sides, while the basal plate on each side is composed (figure 30, L2). The inner component of L2 consists of of LI and L2 from the same side (figures 28, 29, sL2). at least three metachromatic layers of mucoid material A fibro-mucoid, metachromatic adhesive, which may ex- containing scattered fibers arranged parallel with the tend into the stalk, attaches the basal plate to the sub- long axis (figures 31, 32, mL2). Of the species studied, stratum (figure 29, a). Fibers from L2 radiate widely into only C. /. floridanus has a similar arrangement of fibers the adhesive, especially where it extends into the stalk. in mucoid material, and that occurs only in the stalk. No

distinct lamina equivalent to L3 is present. L4 may have Conus jaspideus stearnsi Conrad, 1869 fragments of albumen fused to its inner surface (figure 32, is). (figures 25, 30-32; table 3; USNM 847148) Albumen in this species is metachromatic and includes When sectioned, the lamellate capsules, described by scattered transparent spherules and irregular granular D'Asaro (1986a:88, fig. 4), appear pointed at the apex material that may form layers (figures 30-32). Embryos with a broad stalk and an escape aperture on one side. are distributed throughout it. Page 144 THE NAUTILUS, Vol. 102, No. 4

Figures 26-29. Sections ot the Conu:> jioridanm floridensis egg capsule. 26. Wall showing laminae. 27. Escape aperture. 28.

Stalk 29. Basal plate and adhesive. Positions at which photographs were taken are indicated on figure 24 \ = adhesive, al = albumen, is = inner surface of L4, LI = metachromatic first lamina, L2 = orthochromatic second lamina, L4 = metachromatic albumen retaining layer, mp = mucoid plug, as = outer surface of LI, sL2 = outer portion of L2 in the stalk.

Table .'{. Comparative micromorphology of conid and marginellid egg capsules stained in most cases with toluidine blue (meta irutai hromatic. mp = mucoid plug, ortho = orlhochromatic. pre = present).

Lasers Width closing Layers of Species of wall LI L2 L.3 L4 aperture albumen Author

Conus floridanus 49 ^m meta, ortho, not pre meta, LI, mp, L4, one meta this report

floridensis .5 nm 40 M'li 3-4 nn\ 330 ^m

Conus jaspidius 25 niu meta, ortho and not pre meta, I.I, mp. L4. one meta this report

slearnsi 2-3 Mm meta, 1-2 ^m I N ^m 22 nm Granulina ovuli- \r> fim one lamina present suture in wall two meta this report; formia D Asaro, 1986 Marginella aureo- 12 ^m one lamina present suture? two meta this report; cincta D'.Asaro,

1 9.Sfi C. N. D'Asaro 1988 Page 145

LI forms the outer covering of the escape aperture,

w hile the inner lining is L4. A metachromatic, mucoid plug replaces L2 (figure 31, mp). The edges of the mucoid plug are separated from L2 b> an interdigitating metachromatic boundary zone, similar but less complex than

that of C. /. floridensis. None of the scattered fibers in the metachromatic mucoid layers of L2 enter the mucoid plug. The two outer laminae form the stalk and basal plate, w hile the base of the capsule is composed of the albumen retaining layer and the inner, mucoid part of L2 (figure 32, mL2). The stalk varies in length; therefore, in some specimens the lumen of the capsule may be nearly level with the substratum. A dense mucoid adhesive is present basalK (figure 32, a).

Granulina ovuUjormis (Orbigny, 1841) (figure 33; table 3; USNM 836973)

Macro- and micromorphology of the egg capsules, based in part on eosinophilic features, were described by D'Asaro (1986b: 196, figs. 3-5). Because of structural differences, marginellid capsule micromorphology is not describable with reference to Tamarin and Carriker's (1967) system of enumerating capsule laminae. The simple, pustulate capsules are constructed from an inner component, enclosing two layers of albumen and an embryo, and basement and outer components that sandwich and fuse the previously mentioned structure between them (D'Asaro, 1986b). Because these components were uniformly unstained and did not delaminate during sectioning, the capsule wall is defined as having a single structural lamina of uniform composition, possibK formed by the same portion of the oviduct that produced the layers of L2 in muricaceans and buccinaceans (figure 33, cw). The outer surface of the capsule

is coated with metachromatic mucus. No escape aperture with a mucoid plug exists. The point at which the capsule wall fractures at hatching is marked by a distinct meta-

chromatic suture (figure 33, s). Dense granular albumen

lies just below the capsule wall, while a less dense component immediately surrounds the single embryo. Both albuminous layers stain metachromatically. The adhesive

on the basal layer is not stained (figure 33, a, bl).

Marginella aureocincta Stearns, 1872 (figure 34; table 3; USNM 836974)

Macro- and micromorphology of the egg capsule were described by D'Asaro (1986b: 195, figs. 3-5). 'With application of toluidine blue, all parts of the capsule are metachromatic, except the unstained basement adhesive. Figures 30-32. Sections of the Conus jaspideus stearnsi egg The capsule wall does not delaminate and stains intensely capsule. 30. Wall showing laminae 3 1 . Escape aperture toward to the point of obscuring its layered structure suggesting apex. 32. Stalk, basal plate, and adhesive. Positions at which ovuliformis, it should be defined that, as in Granulina photographs were taken are indicated on figure 25. a = ad- (figure cw, osw). Dis- as a single structural lamina 34, hesive, al = albumen, is = inner surface of L4, LI = meta-

tinctive granulations, described by D'Asaro (1986b: 195, chromatic first lamina, L2 = orthochromatic and metachro- fig. 3), are obvious on the outer surface. No specific matic second lamina, L4 = metachromatic albumen retaining fibrous layer surrounds the albumen. The metachro- ia\er, mL2 = mucoid component of L2, mp = mucoid plug, matic, finely granular albumen occurs in two compo- OS = outer surface of LL Page 146 THE NAUTILUS, Vol. 102, No. 4

Figure 33. Section of the GranuHna ovitlifurmis egg capsule attached to a Thalassia leaf. Figure 34. Section of the Marginella aureocincta egg capsule, a = adhesive, bl = basement layer, cw = capsule wall, da = dense albumen, osw = outer surface of capsule wall, s = suture, ta = thin albumen.

nents, a narrow band of dense material and a more fluid, function of the outer structural lamina, LI. In most central zone in which the embryo lies (figure 34, da, ta). species, LI is thin (5 ^m or less), usually dense and finely fibered, and reflects the final shape (corrugations and DISCUSSION ribbing) imparted by the ventral pedal gland (Gruber, 1982; Sullivan and Maugel, 1984). LI seals the surface Separation of structural laminae by fracturing and to- of the w hole capsule including the escape aperture, but luidine blue staining allows differentiation and identifi- not the basal plate in contact with the adhesive. Since cation of homologous components of neogastropod egg the contents of some prosobranch egg capsules are axenic capsules. Laminae can be identified in this manner be- but not bacteriostatic (Lord, 1986), the fine structure of cause the fibers of which they are composed appear to this lamina could serve as a ph\sical barrier to invasion have been secreted and chemically bound together dur- by microorganisms, especially in species with a vacuo- ing separate phases of assembly. Separate or combined lated and fluid-filled L2. (The basal adhesive may also functions related to protection, structural support, release serve as a barrier.) In Urosalpinx cinerea, LI is thick of larvae or juveniles, and albumen retention are inferred with separated fibers, but the interstices are filled with for the structural laminae. dense mucus (Tamarin and Carriker, 1967). LI could ,\mong the iieogastropods studied, three microstruc- also serve as a reactive substrate during the molding and tural patterns of laminae were identified, each charac- hardening process in the ventral pedal gland which fixes terizing specific higher taxa. Complex, four-layered cap- the capsule in its final shape (see Gruber, 1982 and Sul- sules with an escape aperture sealed by four layers, livan and Maugel, 1984). including two that are mucoid, were found in nearly all L2 comprises the internal skeleton of the capsule wall muricaceans and buccinaceans examined. Three-layered in all neogastropod taxa studied except Marginellidae, capsules having an escape aperture sealed by three layers, which has a single structural lamina (tables 1-.3). In some including one that is mucoid, were found in the Conidae. taxa, extensive cross-linkages between the protein fibers Uncomplicated, single-layered capsules that may only form a homogeneous L2. In others, there are dense ho- have a suture in the wall to facilitate hatching appear mogeneous zones on inner and outer surfaces gradually to be characteristic of the Marginellidae. .separating into directionally oriented fibers forming most Muricacea and Buccinacea are species-rich taxa hav- of the lamina. In ribs or other sculpture, these fibers are ing family or species-specific egg capsule morphologies loo.sely packed. The homogeneous zones, in addition to with final a shape that results from a molding process in skeletal support, could pro\ide another ph\sical barrier the ventral pedal gland (.Ankel, 1929; Gruber, 1982; Sul- to invasion by microorganisms. livan and Maugcl, 1984). Micromorphological similari- In Muricacea and Buccinacea, L2 may have fibrous ties suggest that oviducal mechanisms common to both components with different axial orientation often de- superfamilies are u.sed to construct structural laminae scribed as separate layers (Fretter, 1941; Gruber, 1982). that serve the same respective functions in these taxa Resistance to delamination and cohesiveness suggest that (tables 1, 2). Some species may add additional nonstruc- these layers were constructed from the same capsular tural layers in the ventral pedal gland (table 2, LO; Sul- protein during a continuous process; thus, the\ should livan and Maugel, 1984). be described as parts of a single capsular lamina. If all Protection (sealing the fibrous wall) appears to be a similar fibrovis components anil contiguous homogeneous C. N. D'Asaro 1988 Page 147

zones from previously described egg capsules of these Muricacea and Buccinacea and contain smaller structural superfamilies are viewed in this manner (as organized fibers, usually embedded in a mucoid component. The

in tables 1, 2), then more meaningful comparisons be- capsule wall has two laminae (LI and L2) and an al- tween taxa can be made. bumen retaining layer (L4). Rather than having ribbon-

L3 is distinct in most muricaceans and buccinaceans like fibers in the laminae (see Flower et ai, 1969, for a and appears to be a mucoid extension of the apical plug, discussion of fiber ultrastructure), there are very fine,

a part of the hatching mechanism (tables 1, 2). In both short, and folded or twisted fibers that overlap into a

species of Cantharus, L3 is not present in most of the cross-hatched pattern. A singular mucoid plug lying be- capsule wall; however, the mucoid plug in the escape tween the outer laminae, LI, and the albumen retaining aperture tapers gradually into the wall in a homologous layer, L4, and interdigitating extensively with the fibrous

position. There is also mucoid material with chromo- middle lamina seals the escape aperture. Extensive in- tropic characteristics similar to the apical plug in a ho- terdigitation would require simultaneous formation of

mologous basal position suggesting that a thin layer of L2 and the mucoid plug. There is no evidence in the L3 mucopolysaccharide may be present throughout the conids studied of a lamina equivalent to L3 associated wall but could not be identified with the histological with the mucoid plug. These differences in micromor- technique applied. In the egg capsule of Eupleura cau- phology suggest that important differences in oviducal data etterae, which was described by Gruber (1982: fig. structure and function exist between conids and muri- 14), the singular mucoid plug extending as a layer into caceans and buccinaceans. the capsule wall and surrounding the lumen can be de- Pustulate marginellid egg capsules are microstruc- scribed as L3. Lying between the outer structural layers turally the least complex of the taxa studied, and differ and L3 in E. c. etterae, is another layer (Gruber s sixth markedly from capsules of other neogastropods. Each is layer). The position of this layer, in contact with the constructed of three homogeneous components fused al- equivalent of L2 and surrounding the outer edge of the most indistinguishably into the others to form a capsule mucoid plug formed by L3, and its composition suggest in which the parts do not delaminate during sectioning.

that it is homologous with the first mucoid plug in Chi- Resistance to delamination and a uniform response to coreus florifer dilectus (table 1). toluidine blue suggest that only one protein was used to

L4. the albumen retaining layer, is the primary lamina form the capsule wall. There is no complex escape ap-

in muricaceans and buccinaceans completely surround- erture, but in many marginellids it is possible to identify ing albuminous fluids and embryos (tables 1, 2). Often a preformed suture at which the capsule will break dur- it is not reported in the literature, perhaps because it is ing hatching. The suture is visible at one end and on the 4 nm or less in thickness and usually bound tightly to sides, which corresponds exactly to the position where other components. A function of the layer could be to the capsule wall breaks during hatching. These features prevent nonrefractory albumen from mixing with re- also suggest that the structure and function of the mar- fractory capsular proteins when the outer structural lam- ginellid oviduct is different from that of the major su- inae are assembled by the ciliary mechanism Fretter perfamilies studied. (1941) described. Although this study does show that there are micro- Albumen in egg capsules of muricaceans and bucci- morphological features common to the egg capsules of naceans is stratified to some degree at oviposition (tables some prosobranch taxa, it does not provide clear evidence 1, 2). This was rather obvious in capsules of Chicoreus that can be used to explain exactly how capsules are florifer dilectus, which had a core of dense albumen, formed in the oviducal glands. It can be inferred that free of embryos, that was structurally different from the L4, the albumen retaining layer, is deposited around the surrounding, less dense material. More typically in other albumen and embryos to hold them in a central position species, the core albumen contained the embryos. Dif- \\ hile the more refractory parts of the capsule are formed. ferent layers of albumen in newly deposited capsules It can also be inferred that for most species the remaining suggest that two or more kinds of albumen producing parts of the process involve sequential deposition of struc- cells exist in the oviducts of the species studied, as Fretter tural laminae, as Fretter (1941) described for Nassarius (1941) has demonstrated for several neogastropods. reticulatits, with L3 and formation of the innermost mu- Muricaceans and buccinaceans have structurally sim- coid plug being the second part of the process. The third ilar barriers closing the escape apertures, which are five part of assembly would involve formation of the main to nine times as thick as the capsule wall (tables 1, 2). structural lamina, L2, and the outer mucoid plug. LI The increased thickness of the less refractory mucoid would be the last structural lamina added. Sculpturing, plugs possibly serves to prevent premature hatching of hardening, addition of nonstructural la\ ers, and attach- a capsule. In muricaceans, no fibers from the more re- ment with an adhesi\e are functions of the ventral pedal fractory parts of L2 extend into the mucoid plugs. During gland (Sullivan and Maugel, 1984). sectioning, mucoid plugs in the escape aperture fractured across the width of the capsule wall, while the more ACKNOWLEDGEMENTS refractors parts of the capsule delaminated lengthwise.

Boundary laminae in the wall (LI and L4) appear to I wish to acknowledge the revisions to the manuscript hold the mucoid plugs in position. suggested by a colleague. Dr. Paul V. Hamilton. Some Conid egg capsules are less complex than those of the of the specimens sectioned were provided by Mr. Larry Page 148 THE NAUTILUS, \'ol. 102, No. 4

Dilmore. This project was financed in part with funds Fretter, V. 1941. The genital ducts of some British stenog- from the U.S. Environmental Protection Agency (CR- lossan prosobranchs. Journal of the Marine Biological As- 811649) and the University of West Florida. sociation of the I iiited Kingdom 25173-211

Goldsmith, L. A., H, Hanigan, J M, Thorpe, and K. A. Lind- berg. 1978. Nidamental gland precursor of the egg cap- LITERATURE CITED sule protein of the gastropod mollusc Busycon carica. Comparati\e Biochemistr\ and Physiology 59B:I33-138. Abe, .Naoya. 1983. Breeding of Thais clavigera (Kiister) and Gruber, G. L. 1982. The role of the ventral pedal gland in predation of its eggs by Cronia margariticola (Broderip). formation of an egg capsule by the muricid gastropod In: Morton, B. and D. Dudgeon (eds.). Proceedings of the Euplcura caudata etterae B. B Baker. 1951: an integrated Second International Workshop on the Malacofauna of belia\ ioral. morphological, and histochemical study Mas- Hong Kong and Southern China Hong Kong University ter s thesis. University of Delaware, 142 p. Press, Hong Kong, p. 381-392. Gurr, E. 1962. Staining animal tissue, practical and theoret- Ankel, W. E. 1929. Lber die Bildung der Eikapsel bei ical. Leonard Hill, London. 631 p. Nassa-Arten. V'erhandlungen der Deutschen Zoologischen Hancock, D. A. 1956. The structure of the capsule and the Gesellschaft, Zoologischer Anzeiger, supplement 4:219- hatching process in Urosalpinx cinerea (Say). Proceedings 230. of the Zoological Society of London 127565-571. Ankel, W. E. 1937. Der feinere Bau des Kokons der Purpur- HarasewNch, M. G. 1978. Biochemical studies of the hatching schnecke Nucella lapiUus (L.) und seine Bedeutung fiir process in Busycon. Master's thesis. University of Dela- das Laichleban. Verliaiidlungen der Deutschen Zoolo- ware, Newark, 52 p gischen Gesellschaft, Zoologischer Anzeiger, supplement Hunt, S. 1971. Compari.son of three extracellular structural 10:77-86. proteins in the gastropod mollusc Buccinum undatum L., Bandel, K. 1975. Embryonalgehause Karibischen Meso- und the periostracum, egg capsule, and operculum. Compar- Neogastropoden (Mollusca). .\kademie der Wissenschaf- ative Biochemistry and Physiology 40B:37-46. ten und der Literalur. Mainz. Abhandiuiigen der Math- Lord, .\ 1986. .\Te the contents of egg capsules of the marine ematisch-Xaturwissenschaftlichen Klasse. 1975(1): 1-133. gastropod Nucella lapillus (L.) axenic':* American Mala- Bandel, K. 1976a. Morphologic der Gelege und okologische cological Bulletin 4(2):201-204. Beobachtungen an Muriciden (Gastropoda) aus der siid- Moore, E. A. and F Sander 1978 Spawning and early life lichen Karibischen See. Verhandlungen der Naturfor- historv of Murex pomum Gmelin, 1791. N'eliger 20:251- Gesellschaft in Basel 85(1/2): 1-32. schenden 259. Bandel, K 1976b Spawning, development and ecology of Pechenik, J. .\. 1979. Role of encapsulation in in\ertebrate some higher Neogastropoda from the Caribbean Sea of life histories. American Naturalist 104(6):859-870. C;olombia (South America), X'eliger 19:176-193. Pechenik, J. A. 1983. Egg capsules of Sucella lapillus (L.) Bandel, K. 1976c Morphologic der Gelege und okologie Beo- protect against low-salinit\ stress. Journal of Experimental Buccinacean (Gastropoda) aus der siidlich- bachtungen an Marine Biology and Ecologv 71:165-179. en Karibischen See. Bonner Zoologische Beitraege 27:98- Pechenik, J .A,. 1986. The encapsulation of eggs and embryos 133, by molluscs: an overview .American Malacological Bul- Bandel, K. 1982. Morphologic und Bildung der friihontoge- letin 4(2): 165-172. netischen Gehause bei conchiferen Mollusken Facies (Er- Perry, L. M. and J. S. Schwengel. 1955. Marine shells of the langen) 7:1-153. western coast of Florida. Paleontological Research Insti- Bayne, C. 1968 Histochemical studies on the egg capsules J. tution, Ithaca, New York, 318 p. of eight gastropod molluscs. Proceedings of the Malacol- Radwin, G. E. and J. L. Chamberlin. 1973. Patterns of larval ogical Society of London 38:199-212. development in stenoglossan gastropods. Transactions of D'Asaro, C.N. 1970. Egg capsules of prosobranch mollusks the San Diego Society of Natural History 17(9): 107-1 18. from south Florida and the Bahamas and notes on spawn- Sullivan, C. H. and D. B. Bonar 1984. Biochemical charac- laboratory Bulletin of Marine Science 20:414- ing in the terization of the hatching process of llyanassa ohsoleta. 440. Journal of Experimental Zoology 229(2 ):223-234. D'Asaro, C. N. 1986a. Egg capsules of eleven marine proso- Sullivan, C. H. and T. K Maugel 1984. Formation, organi- branchs from northwest Florida Bulletin of Marine Sci- zation, and composition of the egg capsule of the marine 39(1):76-91. ence gastropod llyanassa obsoleta. Biological Bulletin 167:378- 1986b. Laboratory spawning, egg membranes, D'Asaro, C.N. 389. and egg capsules of 14 small marine prosobranchs from Tamarin, A. and M H C^arriker. 1967. The egg capsule of Florida Bimini, American Malacological and Bahamas. the muricid gastropod I'rnsalpinx cinerea an integrated Bulletin 4(2): 18.5- 199. stud) of the wall b\ ordinarv light, polarized light, and N. E. 1973. The storage and structure of proteins Flower, electron microscop\ Journal of Ultrastructure Research of by the mollusc used in the production egg capsules 21:26-40. CominvUa maculosa. Journal of l'ltra.*tructure Research Webber, H. H. 1977. Gastropoda, Prosobranchia. /n- Giese. 44:1.34-145. in- J. S. and J. S. Pearse (eds). Reproduction of marine Flower, N. E., A. (;eddes, and K M Kudall 1969 I'ltra- J. vertebrates. Vol. IV, molluscs: gastropods and cephalopods. structurc of the fibrous prolcin from the egg capsules of Academic Press. New York, 114 p. the whelk Buccinium undatum Journal of Ultrastructure Research 26:262-273. Franc, A. 1940, Recherchessur ledeveloppement d'Ocinefcra ackulala Lamarck (Mollus()uc gasteropode). Bulletin Biol- ogicjue de la France ct dc la Belgique 74327-345. THE NAUTILUS 102(4):149-153, 1988 Page 149

Two New Species of Metula (Gastropoda: Buccinidae) with a Description of the Radula of the Genus

Philippe Bouchet Museum National d'Histoire Naturelle 55, rue Buffoii 75005 Paris, France

ABSTRACT relatively small or vestigial in the remainder. It is possible that the whole Metula-Ratifusus series discussed by Cer- Two species of Metula H. and A. Adams, 1853, lack a radula, nohorsky (1971) belongs in the Colubrariidae as they all and it is very small in a third species. The morphology of the have similar shell features" (Ponder, 1973:328). teeth is significantly different from that in Pisaiiia, and the two family Colubrariidae is treated as a synonym of genera are probably not closeK related Metula crosnicri new The Buccininae by Ponder and Waren (1988), while Beu and species, from 400-450 m off SW Madagascar, is a large, broad species, with \ery convex whorls and a deep suture. Metula Maxwell (1987) recognize a subfamily Pisaniinae, where africana new species, from the deep continental shelf off West they include Metula, Colubraria, and a number of other

Africa, is considered the descendant of the Mediterranean Plio- genera. cene M. milraeformis (Brocchi, 1814). This lineage cannot be The purpose of the present paper is to provide a name connections taken as evidence for Mediterranean-Indo-Pacific for the West African species that has been known in the in the lower Pliocene as claimed by Grecchi (1978). recent literature as Metula clathrata Adams and Reeve, and to describe another new Metula from the upper continental slope in the Mozambique channel. Several INTRODUCTION additional Indo-Pacific species of Metula, now under study, can be distinguished only on the basis of their The chequered taxonomical history of the generic name protoconch, which has already been emphasized by Al- Metula H. and A. Adams, 1853, has recently been sta- tena (1949) as a taxonomical character. bilized by Emerson (1986), who clarified the identity of its type species, Buccinum clathratitm Adams and Reeve, 1850. Additional information was provided by Beu and SYSTEMATICS Maxwell (1987). The familial position of the genus has been the subject The radula of a species of Metula is figured here for the of a controversy between Ponder (1968, 1973) and Cer- first time (figure 1). It is very small for a buccinid (ribbon nohorsky (1971). Ponder (1968) described the anatomy 25 ;um wide; central tooth 6.5 ^m wide, lateral teeth 12 of Ratifitsus Iredaie, 1919, and Iredalula Finlay, 1927; ^m wide) and very similar to the radulae of Ratifusus he concluded that their peculiar glandular mid-esoph- and Iredalula figured by Ponder (1968): the central tooth agus as well as other features of the anterior alimentary has a narrow arched basal plate with 3 long, slender, and canal justified their inclusion in the family Colubrariidae, equal cusps; the lateral teeth also have a narrow basal which he considered to be anatomically distinct from plate and 3 long slender cusps, the outermost one being the Buccinidae. Cernohorsky (1971) regarded the pres- longest. ence of a vestigial radula in Ratifusus and Iredalula to A radula has been looked for, but not found in Metula indicate placement in the Buccinidae, since the species amosi Vanatta, 1913, and M. cumingi (Adams, 1853);

of Colubrariidae have no radula at all, and he suggested several specimens were examined in each case by A. a placement in the buccinid subfamily Pisaniinae. This Waren (personal communication). I do not consider pres- view has been accepted by most subsequent authors deal- ence or absence of this very reduced radula to be of ing with Metula (Olsson and Bayer, 1972; Kilburn, 1975; generic importance. Houbrick, 1984; Emerson, 1986), who apparently over- The radula (figure 2) of Pisania striata (Gmelin, 1791),

looked Ponder s (1973) refutation of Cernohorsky s point tvpe species of Pisania, is 150 ^m wide; it differs in of view. Ponder (1973) confirmed that Ratifusus, Ire- having a central tooth with a large square basal plate dalula. and Colubraria shared the same anatomical char- and 5 short and broad cusps, the outermost 2 being small- acters that separate them from the Buccinidae, and com- er; the lateral teeth are more strongly built, with 3 un- mented on the radular differences: "It thus appears that equal cusps.

some Colubrariidae have lost the radula and that it is In view of the small variation of radular types in buc- Page 150 THE NAUTILUS, Vol. 102, No. 4

cinids, this difference is remarkable and probabK indicates that the two genera are not closeK- related. What- ever rank (subfamily or tribe) the Pisania group is given in Huccinidae, additional research is needed before the

Colubraria group is considered a mere s>nonym of it.

DESCRIPTIONS

[For a diagnosis of the genus see Altena (1949) as An- temetula]

Metula crosnieri new species (figures 3-5, 9)

Description: Shell solid, fusiform, consisting of 2.5 protoconch and 6.2 teleoconch whorls. Spire high, body whorl comprising 64% of total shell height. Protoconch (figure 9) with large nucleus of two smooth convex whorls abruptK demarcated from teleoconch. Teleoconch whorls convex, without sutural ramp, with deeply impressed suture. Sculpture of raised spiral cords and curved opisthocline axial ribs; cords and ribs producing beaded intersections and pitted intervals. Eight spiral cords on spire whorls; 4 minor, intermediate cords on penultimate whorl; about 18 cords above periphery of body whorl, principal and secondary cords alternating rather regularly, and 33 cords below periphery, of which about 15 are set close together in siphonal region. Axial ribs about equal in strength to spiral cords on spire whorls; weaker on penultimate whorl, with main sculpture being spiral on body whorl. In addition to axial ribs, several incremental scars are obvious, especially on body and penultimate whorls. Aperture ovate; inner lip thin, smooth, adherent to body whorl, thicker in columellar region; outer lip bearing 12 very weak teeth that do not correspond with position of external spiral cords; two most apical teeth slightly stronger. Peristome thickened, forming broad varix, also covered by spiral cords. Siphonal canal long, broad, widely open, and recurved. Fasciole indistinct. Colour light tan, with a very indistinct darker spiral band at periphery of body whorl; incremental scars light- Figures 1, 2. Scanning electron niicrographi ol raciulac I. er; aperture white. Metula africana new species, scale bar lO/iiii. 2. Fisania striata Dimensions of the holotype: Height 51 3 mm, width ((Jnielin. 1791), scale bar .50 nm. 19.4 mm; height of the aperture 24.5 mm, width 8 mm; height of the body whorl 33 mm. Remarks: Metula crosnieri has remarkably convex The largest paratype is 55.4 mm high. whorls and a deep suture when compared with its con- Type lf>cality: Mozambique channel, SW Madagascar, geners. The protoconch (figure 9) indicates non-planc- off Baie de Faiiemotra, 22°15'S, 43°05'E, 470-475 m. totrophic larval development. The combination of these

two characters distinguishes it from all known Indo-Pa- Type material: Holotype and paratype 1 (MNHN) from cific Metula. the type locality, collected by A. Crosnier, Dec. 2, 1973 In the Atlantic, Bartschia significans Rehder, 1943, aboard R.V. "N'auban"; paratype 2 (MNHN), Mozam- type species of Bartschia Rehder, 1943, has even more bique channel, SW Madagascar, 22°17'S, 43°04'E, 400- convex whorls, and a multispiral protoconch. I cannot 450 m, collected by R. v. Cosel, Nov. 30, 1986 aboard find characters other than the convexit\- of the whorls trawler "Mascareignes III". that sharply distinguish Bartschia from Metula, and con-

Distribution: Known only from llic type material, off clude that Bartschia should be considered at most a sub- SW Madagascar. genus of Metula. p. Bouchet 1988 Page 151

I am naming this species after my colleague Alain

Crosnier, who first collected it during a survey of deep water shrimp populations oft Madagascar.

Metula ajricana new species

(figures 1, 6-8, 10)

Metula clathrata Knudsen, 1956;39, plate 1, figure 1; non M. clathrata (Adams and Reeve, 1850). iKin Metula knudseni Kilburn, 1975:592 (replacement name for Buccinum clathratum Adams and Reeve, 1850: see Emerson, 1986).

Description: Shell solid, fusiform, consisting of 7 teleoconch whorls. (Protoconch of holotype partly broken, on a paratype consisting of large nucleus and 1.5 smooth convex whorls, figure 10.) Teleoconch whorls convex, with faint but distinct sutural ramp on early whorls; ramp indistinct on penultimate whorl, body whorl evenly convex. Teleoconch sculpture of raised spiral cords and slightly curved opisthocline axial ribs; intersections distinctly beaded in sutural ramp area, only slightly nod- ulous below ramp. Spiral and axial sculpture of similar strength on early whorls; later spiral sculpture gradually dominates. Nine primary spiral cords per whorl on spire whorls, first (adapical) and third cords stronger, limiting sutural ramp. Fine spiral threads present between cords on penultimate and bod\ whorl, several eventually developing into secondary spiral cords, with one present in sutural ramp area of body whorl. About 22 primary cords below periphery of body whorl, plus another 10 in siphonal area. Incremental scars distinct; growth lines very distinct between axial ribs. Aperture ovate, narrow. Inner lip thin, smooth, adherent to body whorl, thicker in columellar region. Outer lip regularly convex, bearing small teeth that correspond at least partly with position of spiral cords on peristome; a group of 5 teeth forms a small callus in apical portion of outer lip, delimiting small anal canal. Peristome forming a thickened varix, over which spiral sculpture extends. Siphonal canal short, narrow, open, only slightly recurved. Ground colour of shell beige cream, with 3 brown spiral bands; 3 adapical cords brown, interval between them beige cream. Below, uniformly beige band extends over next 2 spiral cords, occupies central position on spire whorls. Darkest band occupies next 4 spiral cords and intervals between them; this band occupying suprasu- tural position on spire whorls, and a central position on body whorl. Below brown band a second beige band extends over 4 spiral cords; next 2 cords brown, with Figures 3-8. Back, side, and front views. 3-5. Metula cros- brown colour fading towards base of shell. Aperture cream nieri new species, holotype, 51.3 mm. 6-8. Metula ajricana colored. new species, holotype, 54.5 mm.

Dimensions of the shell: Height 54.5 mm, width 17.5 mm; height of the aperture 27 mm, width 7 mm; height of the body whorl 35 mm. Other material examined: SENEGAL: off Saint Louis, "deep water", 4 shells (MNHN, leg. M. Pin), 10 shells Type locality: Off Saint Louis, Senegal, in 300-600 m. (coll. M. Pin, Dakar); IVORY COAST: off Abidjan, 1

Type material: Holotype (MNHN), paratype 1 (MNHN) shell, coll. Marche-Marchad (MNHN), and 1 shell, P. Le and paratype 2 (AMNH 198755), all from the type lo- Loeuff coll. (MNHN); EQUATORIAL GUINEA: Atlan- cality, collected by M. Pin on the trawler "Louis Sauger". tide Sta. 120, 02°09'N, 09°27'E, 250-850 m, 1 specimen Page 152 THE NAUTILUS, Vol. 102, No. 4

calit\, and distribution of M. clathrata, Grecchi (1978)

speculated that the presence of its presumed ancestor M. mitraeformis in the Mediterranean Pliocene was an indication of Mediterranean-Indo-Pacihc connections in the lower Pliocene after the Messinian salinity crisis. With M. clathrata now known to be a West American species distinct from M. africana, the history of the M. mitraeformis-africana lineage can not be taken as an indication of such connections. This lineage probably has an Eastern .Atlantic history dating to the Miocene; although the Neogene West African fossil record is lacking,

it is far more probable that M. mitraeformis reinvaded the Pliocene Mediterranean from West Africa rather than from the Indo-Pacifie through unproven maritime connections. Figures 9. 10. Protoconchs. 9. Metula crosnieri. 10. Metula ajricana. Scale line 1 mm. ACKNOWLEDGEMENTS

I thank Dr. A. Waren who prepared the radula of Metula (Kiiudsen, 1956) (ZMC); CONGO: West of Pointe-Noire, africana and commented on the manuscript. A. Crosnier, 100 m, 1 shell, A. Crosnier coil. 1962 (ANSP 333810). M. Pin, and R. von Cosel collected most of the material cited in this paper. Dr. W. Emerson generousK refrained Distribution: Deep continental shelf and upper slope from describing M. africana when he learned 1 was of West Africa, from Senegal to Congo. working on it. Photography is b\ P. Lozouet. Remarks: Metula africana has been figured three times in the literature: by Knudsen (1956: plate 1, figure 1) LITERATURE CITED under the name Metula clathrata; by Emerson (1986: figures 4, 5) as Metula sp.; finally by Kaicher (1987: card Alteiia, C O van R 1949. The genus Antemetula Rehder

4851) as Metula sp. I refer to Emerson (1986) who re- in the Indo-West Pacific area, with the description of two viewed the nomenclature of this West African Metula new fossil species. Bijdragen tot de Dierkdunde 28:385- 393. and concluded that it represents a new species. Beu, A. G. and P. A. Maxwell. 1987. A revision of the fossil The protoconch (figure 9) indicates non-planctotro- and living gastropods related to Plesiotriton Fischer, 1884 phic larval development. (Family Cancellariidae, Subfamily Plesiotritoninae n. There may be clinal variation in adult size along the subfam.) with an appendix: genera of Buccinidae Pisani- West African coast: The 18 shells examined from Senegal inae related to Colubraria Schumacher. 1817. New Zea- 41.7 the 2 shells from Ivory have a mean height of mm; land Geological Survey Paleontological Bulletin 54:1-140, that Coast, although fully adult, measure 32 and 36 mm; Cernohorsky, W. 1971. Indo-Pacific Pisaniinac and related from Equatorial Guinea is 30.6 mm high, and that from buccinid genera. Records of the .\uckland Institute and Congo 25.5 mm. Museum 8:137-167. The tiny rachiglossate radula (figure 1) was prepared Emerson, W. K. 1986. On the type species of Metula H. & clathratum A. .\dams and from the specimen taken during the .-Ktlantide expedition \. Adams, 1853: Buccinum (ZMC). Reeve, 1850. The Nautilus 100(1 ):27-:30 Grecchi, G. 1978. Problems connected with the recorded The close connection between Mediterranean Pliocene occurrence of some mollusks of Indo-Pacific affinit\ in the and Recent West African marine faunas has been dem- Pliocene of the Mediterranean area Rivisla Italiana di onstrated in a number of paleontological papers (for re- Paleonlologia 84:797-812. see Ruggieri, 1967; Marasti and Raffi, 1979; cent reviews Houbrick, R. 1984. A new "Metula" species from the Indo- species of Me- Sahelli and Taviani, 1984). With a single West Pacific. Proceedings of the Biological Society of tula present in the Pliocene of Italy and a single Recent Washington 97(2):420-424.

species in West Africa, it is reasonable to assume that Kaicher, S. D. 1987. Card catalogue of world-wide shells. Metula mitraeformis (Brocchi, 1814) (for figures and Pack 48: Buccinidae, part 3 Published by the author, St. references see Pelosio, 1966) is the direct ancestor to M. Petersburg. Killnirn, R N 1975. Taxonomic notes on South African ma- africana. I have examined material of the fossil species rine Mollusca (5): including descriptions of new ta\a of and found it to differ from the Recent one by its much Rissoidae, Cerithiidae, Tonnidae, Cla.ssididae, Buccinidae, weaker axial sculpture, which on the body and penul- p-asciolariidae, Turbinellidae, Turridae, Architectonici- timate whorls is limited to growth lines. In M mitrae- dae, Epitoniidae, Limidae, Thraciidae. Annals of the Natal sutural ramp that extends formis, there is a very broad Museum 22(2):577-622. over the adapical third or half of early teleoconch whorls, prosobranchs of tropical West Af- Knudsen, J. 1956. Marine and becomes obsolete on the penultimate and body whorls; rica (Stenoglossa). Atlantide Report 4:7-110.

the siphonal canal is also broader. Marasti, R and S. Raffi 1979 Observations on the paleocli- Because he was mistaken about the identity, type lo- matic and biogeographic meaning of the Mediterranean p. Bouchet 1988 Page 153

Pliocene molluscs. State of the problem. 7th Intenuilinnal Ponder, W. and A Waren 1988 Classification of the Cae- Congress on Mediterranean Neogene, Athens, 8 unnum- nogastropoda and I leteroslropha—a li.sl of the family-group bered pages. names and higher taxa. Malacological Review, supplement Olsson, A. and F. Bayer. 1972. American Metulas (Gastro- 4:286-32-1. poda: Buccinidae). Bulletin of Marine Science 22(4):900- Ruggieri, G. 1967. The Miocene and later evolution of the 925. Mediterranean Sea. In: Adams, C. G. and D. V. Ager (eds.). Pelosio, G. 1966. La Malacofauna dello stratotipo del Tabi- Aspects of Tethyan biogeography. The Systematics Asso- aniano (Pliocene inferiore) di Tabiano Bagni (Parma). Bol- ciation, London. P. 283-290. lettino della Societa Paleontologica Italiana 5(2):l()l-183. Sabelli, B. and M. Taviani. 1984. The paleobiogeographic Ponder, W. 1968. Anatomical notes on two species of the distribution of the Mediterranean benthic mollusks and Colubrariidae. Transactions of the Roval Society of New- the Messinian salinity crisis or where did the mollusks go? Zealand, Zoology 10(24):217-223. Annales de Geologic des Pays Helleniques 32:263-269. Ponder, W. 1973. The origin and evolution ot the Neogas- tropoda. Malacologia 12(2):295-338. THE NAUTILUS 102(4):154-158, 1988 Page 154

A New Species of Intertidal Terebra from Brazil

Kurl Auffenberg Harry G. Lee Malacolog> Division 709 Uomax Street Florida Museum of Natural History Jacksonville, FL 32204, USA University of Florida Gainesville, FL 32611. USA

ABSTRACT micrometer and converted to millimeters. Terminology follows that of Miller (1970, 1971). Two preserved but Terebra imitatrix new species is described from northern Brazil completeK retracted adult specimens of the new species and compared to morphologically similar species of Hastnla. (ANSP 299957), Hastula maryleeae Burch (UF 113539) The three known terebrid feeding types are briefl\ reviewed. Hastula salleana (Deshayes) (UF 48197, 113540), Dissection of the foregut of this new species revealed characters and lot. dried-in specimens of that overlap two of the major feeding types. were dissected from each Two Hastula maryleeae Burch (T. Bratcher collection) were rehydrated in a weak solution of potassium h\dro.\ide, Key words: Gastropoda; Terebridae; Terebra; anatom\ : Bra- transferred into water, and dissected. The radular sac was extracted and dissolved in a weak solution of potassium hydroxide. Radular teeth were individual!} mount- INTRODUCTION ed on scanning electron microscope specimen stubs covered with double-sided tape. Micrographs were made While compiling distributional records of the West At- with a Hitachi 5-415.^ scanning electron microscope. lantic Hastula cinerea species group, the junior author Repositories of examined specimens are indicated by located an unidentified lot of 23 specimens from Brazil the following abbreviations: in the Academy of Natural Sciences of Philadelphia AMNH American Museum of Natural Histor\' (ANSP 299957). These specimens were compared to the ANSP Academy of Natural Sciences of Philadelphia known terebrids from the Atlantic and were found to UF Florida Museum of Natural History belong to a distinct undescribed species. An additional lot of the same species was subsequently found in the American Museum of Natural History (AMNH 129280). SYSTEMATICS Matthews et al. (1975:99, fig. 31) in their treatment of Hastula ciiterea describe and illustrate a protoconch con- Family Terebridae Morch, 1852 Bruguiere, sistent with this new species. The Brazilian specimens of Genus Terebra 1789 figured Rios (1970: Hastula salleana (Deshayes, 1859) by Terebra imitatrix new species pi. 47, 1975:127, pi. 38, fig. 560, 1985:131, pi. 45) 123, (figures 1-6, 8, 9, table 1) are probably referable to this new species, but the figured specimen was unavailable for examination. This paper Description: Shell (figures 1-3, 6) medium in size, describes this new species and compares it with mor- broadening anteriorK ; color variable, ranging from phologically similar species of Hastula. A description of banded, or cream, to purplish-brown; teleoconch whorls the foregut anatomy reveals that it does not conform to 10-12; sides flat to slightly convex. Protoconch whorls any of the three known feeding types, which are re- 1.5-2.0; glassy, transparent (figure 6). .\xial sculpture of viewed herein. close-set, recurved ribs of \ariable strength, generally distinct near the suture, becoming obsolete anteriorly; = penultimate MATERIALS AND METHODS major axial ribs 34-57 (x 41.6) on the whorl. Spiral sculpture of very faint microscopic incised

Only shells possessing 10 or more teleoconch whorls and lines most distinct in the intercostal spaces on the upper with intact protoconchs and apertures were measured whorls, obsolete on later whorls, rarely cro.ssing the axial with Vernier calipers. All dissections were made under ribs; spiral rows of pits absent. Last whorl with obsolete a Wild M-5 dissecting micro.scope and line drawings axial ribs and spiral incised lines; color pattern variable, made with the aid of a camera lucida. Description and typicalK of five diffuse color bands: one white pre-sutural ob- discussion of the anatomy is limited to the foregut due band usiialK w ith distinct brown spots that become to poor preservation in the upper whorls. Anatomical solete toward apertural lip, one broad bluish-black zone and protoconch measurements were made with an ocular at shoulder, one pinkish band at periphery, one purplish- K. Auffeiiberg and H. G. Lee 1988 Page 155

Figures 1-3. Terebra imilatrix new species. 1. Holotype, ANSP 299957 (27.5 mm shell length) 2, 3. Paratypes, ANSP 369293, all from Rio Grande do Norte, Brazil, -sand island at mouth of the inlet at Areia Branca.

brown band below periphery, and one white basal zone. bulbs anteriorly (figure 5). Radular organs attached to Columella brown, centrally concave and slightly re- right side of anterior portion of buccal cavity by very curved; rounded rib present on anterior edge. Parietal short duct (0.1 mm). Two rows of radular teeth situated callus thin, transparent to light brown. Fasciole white to obliquely in radular sac and caecum (figure 5). Radular bluish-gray with distinct white rib. Anterior siphonal teeth about 30; 0.1-0.2 mm in length, slightly curved, notch moderately broad, straight. Aperture light to dark not barbed (figures 8, 9). Poison gland long (6.2 mm), brown with white band. extremely convoluted, entering right side of buccal cavity slightly behind radular sac (figure 5). Poison bulb External anatomy: Animal cream-colored with no ap- small (0.9 mm), weak, seemingly vestigial, lying at ven- parent pattern in alcohol-preserved specimens. Oper- tro-posterior end of cephalic hemocoel. culum corneous, small, thin. Eyes on short, broad eye- stalks. Labial tube large, spoon-shaped (figure 4). Anterior Etymology: From the Latin feminine noun meaning end of labial tube terminating in thick muscular lips one who imitates, in reference to the deceptively close bounding mouth slit; well-developed sphincter lacking. resemblance of the shell to that of Hastula cinerea (Born,

1778), with which it occurred in the type lot. Foregut anatomy: Labial cavity large, dominated by massive (3.7 mm), extendable, muscular organ (accessory Table I. Measurements (mm) of shell characters of Terebra feeding apparatus; Miller, 1970, 1971) that tapers an- imitatrix new species and Hasttila rnaryleeae Burch teriorly; muscular organ with two rows of papillae on ventral side (figure 4); attached posteriorly to the left Species side of the cephalic hemocoel by connective tissue; retractor muscle large, originating in foot below and slightly posterior to anterior siphon, passing through cephalic hemocoel. Buccal tube muscular, short (0.9 mm), tapering anteriorly. Buccal cavity small (0.6 mm), rounded. Pre-ganglionic esophagus enters buccal mass posteriorly

(figure 4). Salivary glands not located. Radular sac blade- shaped, small (0.8 mm). Radular caecum (0.4 mm) with well-developed groove running posteriorly, two distinct Page 156 THE NAUTILUS, Vol. 102, No. 4

•s be

Figures 4-6. Terebra irnitatrix new species. 4. Diagrammatic dorsal view nf the organs of the foregut 5. Right side of the buccal tulie showing radular organs and insertion of the poison gland. 6. Protoconch. 7. Hastula manjleeae Burch, protoconch (.-WINH

191815) Scale bars = 1 mm. af, accessory feeding apparatus; be, buccal cavity; bt, buccal tube; It, labial tube; pb, poison bulb; pe, pre-ganglionic esophagus; pg, poison gland; re, radular caecum; rm, retractor muscle; rs, radular sac.

Type locality: Brazil, Rio Grande do Norte, sand island paratypes. Although these two species may occur micro- at mouth of the inlet at Areia Branca, 04°57'S, 37°08"W, s\ mpatrically, we assume there is no trophic competition

G. & M. Kline et al., 14 December 1963. due to the strikingly divergent feeding organs (see discussion). Holotype: ANSP 299957, shell length 27.5 mm, width 6.7 mm. Comparative remarks: Terelrra irnitatrix is similar in Paratypes: Paratypes 1-20, ANSP 369293, from the shell morpholog\- to the West Atlantic Hastula cincrea tvpc locality (15 dry, 5 in alcohol). Paratypes 21-22, UF group. It can be easily separated from Hastula cinerea 115180, from type locality. Paratypes 23-24, AMNH and Hastula salleana by the lack of spiral rows of pits 129280, Brazil, Ceara, Acaraii, 02°53'S, 40°07'W. and the generally more numerous, but less prominent, axial ribs. .Although similar in size, T. irnitatrix is usually Distribution: PresentK know n onl\- from northern Bra- broader anteriorly . Hastula cinerea and H. salleana have zil. 3.5-4.0 protoconch whorls, while T. irnitatrix has 1.5-

Ecology: Based on the locality label of the type lot, this 2.0. The typical color pattern of T. irnitatrix is more species is found in sand near or at inlets much like some distinctK banded with larger and more distinct brown populations of Hastula. One specimen of Hastula ci- spots al the suture and a broader white subsutural band. nerea (Born, 1778) was found in the type lot and another Terelna irnitatrix and Hastula manjleeae are more dif- (AMNH 129269) from the same locality as the AMNH- ficult to separate, particularly southern Caribbean pop- K. Auffenberg and H. G. Lee 1988 Page 157

Illations of H. niarijleeae described as Terehra tobagoensis Nowell-l'slicke, 1969, now placed in the s\non\ my of H. manjleeae by Bratcher and Cernohorsky (1987). Examination of the lectotype (AMNH 195453 designated bv Bratcher and Cernohorsky, 1987:194, pi.

60, fig. 235c)' and paralectotypes (AMNH 191819) of T. tobagoensis. as well as several other lots from Tobago, typical specimens of H. manjleeae from Te.xas, and specimens of a weakly ribbed form of H. maryleeae from the Dominican Republic (see Bratcher and Cernohorsky, 1987, for discussion) revealed consistent shell characters b\ which the two species may be separated. Anatomical features are discussed below. Typical H. maryleeae is from T. imitatrix by the distinctive shell easily separated Figures 8, 9. Radular teeth of paratype (ANSP 369293) of shape caused by the enlarged nodes of the axial ribs at x Terehra imitatrix. new species. 8. Whole tooth, 60 . 9. Tip the suture and by the slight crenulations at the sutures of lootli, 200 X. of the upper whorls. Hastula manjleeae has a much smaller shell than T. imitatrix at the same whorl count (table 1). The shell of T. imitatrix broadens more an- from the West Atlantic Hastula. We have dissected H. teriorK- and has a proportionately slightly larger aper- salleana and H. manjleeae for comparison, and both 1.5-2.0 ture. The protoconchs of both species have whorls, possess a Type IIA polyembolic proboscis (Miller, 1970, however, the protoconch of T. imitatrix is more bulbous 1971). Terebra imitatrix has a large spoon-shaped labial (figures 6, 7; table 1). Typical T. imitatrix also resembles tube, presumably ineversible, a labial cavity dominated the West African Hastula aciculina (Lamarck, 1822), by the accessory feeding apparatus, an extremely short particularK- in color pattern. However, T. imitatrix lacks buccal tube incapable of extending outside the mouth the supra-sutural groove and callosity found in H. acic- and the buccal cavity, and the associated radular organs ulina (Bouchet, 1982; Bratcher and Cernohorsky, 1987). are minute in comparison to those of Hastula. The vestigial poison gland and bulb are barely recognizable as DISCUSSION such and are considered homologous to the massive poison gland and bulb of Hastula only by the similarity of Miller (1970) proposed a division of the Terebridae into position and the site of its entrance into the buccal cavity. three major groups based on the anatomy of the foregut The radular teeth are similar in size and shape to those (feeding type) and later published a series of papers of Hastula maryleeae (0.15 mm). The teeth of H. sal- (Miller, 1971, 1975, 1979) on this subject supported by leana and H. cinerea are larger (0.5 mm) and barbed. in-depth life history studies. These data are reviewed The presence of an accessory feeding apparatus and the and the species assigned to each group are listed in small size of other foregut organs place T. imitatrix near

Bratcher and Cernohorsky (1987). Type I species have the group possessing a Type III polyembolic proboscis. a long labial tube and a short buccal tube. They do not However, the presence of radular and poison organs, such possess a radular apparatus or poison organs. This group as occur in T. imitatrix, has not been reported for this is further divided into two subgroups, lA and IB, based proboscis type. Also, the shell of T. imitatrix is very primarily on habitat and pre%' (see Bratcher and Cer- different from the species in this group, all of which have nohorsky, 1987). Type II encompasses species exhibiting shells with deeply impressed sutures, a sub-sutural groove typical toxoglossan feeding characters. The labial tube and strong to moderate axial sculpture. is long and eversible and the buccal cavit\ is relatively The feeding behavior of Hastula cinerea and of H. large. The buccal tube is long and retractile. They have inconstans (Hinds, 1844) have been well documented a poison gland and bulb as well as a radular sac containing by Marcus and Marcus (1960) and Miller (1979), re- two rows of harpoon-like radular teeth. This feeding type spectively. The long eversible labial tube forages for prey is further divided into two subgroups, types IIA and IIB, items. A single radular tooth is passed through the labial based primarily on habitat and behavior. Type IIA in- tube, held in the tip, and inserted into the prey to fa- cludes several Indo-Pacific species, as well as the West cilitate penetration of the venom. The prey is then in- Atlantic Hastula discussed in this paper (see Bratcher gested via the labial tube. Miller (1970) suggests that and Cernohorsky, 1987). Type III species possess an ac- species with a Type III proboscis forage by utilizing the cessory feeding organ which grasps prey and pulls it into accessory feeding apparatus, and that food items are the labial cavity. The\' lack a radular apparatus and some passed into the opening of the interior buccal tube. We have lost the buccal tube and salivary glands. This group do not know if the radular and poison organs are func-

presentK contains no .\tlantic species, but is represented tional in Terebra imitatrix. If so, the radular tooth is b\' several Indo-Pacific taxa (see Bratcher and Cerno- either transferred from the buccal tube to the accessory horsky, 1987). feeding apparatus and inserted into prey outside the

Terebra imitatrix has very different foregut anatomy body, or once it reaches the buccal tube. Page 158 THE NAUTILUS, Vol. 102, No. 4

The presence in Terebra imitatrix of the accessory LITERATURE CITED feeding apparatus recorded in a few Indo-Pacific Tere- Bouchet. P. 1982. Les Terebridae (Mollusca, Gastropoda) de bra and the poison anil radiilar organs of Hastula make r.\tlantique Oriental. Bollentino Malacologico, Milano Terelna imitatrix may be closely this species unique. 18(9-12):18.5-2I6. aciculina, related to the West African species, Hastula Bratcher, T. and \V. O. Cernohorsky 1987 Living terebras however, this species must be studied anatomically be- of the world .\ monograph of the Recent Terebridae of fore this assignment can be verified Generic limits within the world .Xmerican Malacologists, Inc.. Melbourne, FL, the Terebridae are presently p<}ori\ understood with much 240 p. overlap in shell and anatomical characters. Additional Marcus, E. and E. Marcus 1960. On Hastula cinerea. Bo- Univer- anatomical studies are needed to clarify the taxonomic leiitim Faculdade de Filosofia Ciencias e Letras. sidade de Sao Paulo, no. 260, Zool. no. 23:25-54, pis. 1-5. and e\oiutionar\ significance of the foregut in the Tere- Matthews. H. R., \. C. Dos Santos Coelho, P. De Sa Cardoso, bridae. and M. Kempf. 1975. Notas sobre la familia Terebridae no Brasil (Mollusca, Gastropoda). .-Vrquivos do Museu Na- cional. Rio de Janiero 5585-104. ACKNOWLEDGEMENTS Miller. B .A 1970 Studies on the biologN of Indo-Pacific Terebridae. Ph D. dissertation. I'niversity of New Hamp-

sliirc. Durham. 213 p. We wish to thank several people who have assisted us Miller, B. .\. 1971. Feeding mechanisms in the family Tere- during this study. Special thanks go to Robert Robertson, bridae. .Annual Report of the American Malacological George M. Davis, and Mar) \. (iarback of the .\cademy I'nion for 1970. 1970:72-74. Miller. B. .A. 1975. The biolog\ of Tereiwa gouWi Deshayes, of Natural Sciences of Philadelphia for allowing us to 1859, and a discussion of the life history similarities among borrow and dissect the specimens on w hich this descrip- other terebrids of similar proboscis tvpe Pacific Science tion is based. William K. Emerson, Harold Feinberg, and 29:227-241. Walter Sage at the .American Museum of Natural Histor\ Miller, B. .\. 1979. The biology of Hastula inconstans (Hinds, several allowed us to visit their department and borrow 1844) and a discussion of life histor\ similarities among lots of \ery pertinent material. Twila Bratcher of Hol- other Hastula of similar proboscis type. Pacific Science lywood, California kindly provided comparative mate- 33(3):289-306. rial, as did Eliezar C. Rios (Funda^ao Universidade do Nowell-l'sticke, G. W. 1969 .A supplementary listing of new Rio Grande do Sul, Brazil), the late Joseph Rosewater of shells (illustrated). Privately published. St. Croix, Virgin the U.S. National Museum, and Thomas Pulley and Con- Islands, 31 p., 4 pis. Rios, E. C. 1970, Coastal Brazilian shells Funda(,ao Cidade stance Boone of the Houston Museum of Natural Science. do Rio Grande, Museu Oceanografico de Rio Grande, Bra- Marcos Guianaldo and Armando Garcia, Florida Mu- zil, 255 p.. 60 pis. seum of Natural History, prepared figures 4-7. The mi- Rios, E. C. 1975. Brazilian marine mollusks iconograph\. with the scan- crographs were made by the senior author Fundagao Universidade do Rio Grande, Centro de Cien- in of ning electron microscope housed the Department cias do Mar, Museu Oceanografico, Brazil, 331 p., 91 pis. Zoology, University of Florida. In addition, we thank Rios, E. C. 1985. Seashells of Brazil Funda^ao Cidade do Richard S. Houbrick, William K. Emerson, and Fred G. Rio Grande, Fundagao Universidade do Rio Grande, Mu- pis. Thompson for reviewing earlier drafts of the manuscript. seu Oceanographico, Brazil, 329 p , 1421 THE NAUTILUS 102(4):159-163, 1988 Page 159

Notes on the Biology and Morphology of Margaritifera hembeli (Conrad, 1838) (Unionacea: Margaritiferidae)

Douglas G. Smith Museum of Zoology Universit\ of Massachusetts Amherst, MA 01003-0027, USA

ABSTRACT range in the Red River drainage as indicated by museum records, particularly a specimen in the American Mu- The freshwater pearl mussel Margaritifera hembeli is found seum of Natural History (AMNH 193786) from the Red onK in the Red River basin and a few nearby drainages in River in Arkansas. During the present century, however, Louisiana. Though of concern to conservationists because of its deterio- declining numbers, M. hembeli remains virtually unknown with the range has contracted considerably due to is respect to its anatomy and biology. The species contains all the rating environmental conditions. The present range anatomical characters that typify margaritiferid species. The limited to the Bayou Teche drainage (Vidrine, 1985) and sexes are separate and the gonads show a definite seasonality a single stream in the Red River drainage. The drastic in activity. Gametogenesis is pronounced in specimens collected reduction in range has elicited concern from the federal in the fall, followed by degeneration of reproductive tissues in government, which has provided protection for the re- winter through to late spring. It is concluded, on the the late maining populations (Stewart, 1988). basis of observed gonadal activity, that oviposition and spawn- Despite increased concern for M. hembeli, very little ing take place between late November and late January Char- is known about this species other than the characteristics acters are evident in the morphology of the visceral nervous of its shell. Ortmann (1912) provided a brief description svstem and the stomach of M. hembeli that clearly distinguish ' M. hembeli from M. marrianae and other eastern North Amer- of the anatomy of M. "hembeli, but the specimens upon ican margaritiferid species. A distinct relationship between M. which he based his description came from Alabama and,

hembeli and M. marrianae, however, is suggested by the mu- therefore, are appropriately referred to M. marrianae. tual occurrence of lateral hinge teeth and a corrugated surface Hence the anatomy of M. hembeli remains undescribed,

of the posterior portion of the shell. Due to the lack of knowl- and nothing is known about its biology. The present study edge of the anatomy and biology of other margaritiferid species, provides some information on gonadal activity and de- especially those living in Asia, it is premature to suggest rela- tails of the anatomy of the stomach and nervous system. tionships lietween M. hembeli and other described margari- Comparisons are made with other North American mar- tiferid species, particularly those with lateral hinge teeth. garitiferid species, including M. marrianae, as studied Key uords: Margaritifera: anatomv, reproduction. North by Smith (1979a, 1980, 1986, and unpublished). .America MATERIALS AND METHODS INTRODUCTION A total of 43 partially or completely relaxed, preserved, The North American freshwater mussel Margaritifera specimens were studied. All were collected on various hembeli (Conrad, 1838) was once believed to comprise dates from 1973 to 1986 from Brown Creek, Gardner, two geographically discontinuous populations ta.xonom- Rapides Parish, Louisiana. Specimens had been fixed in ically linked b\ vague similarities of the shell. Johnson 10% formalin, washed in water, and stored in 50% iso- (1983) separated the two populations taxonomically by propyl alcohol. Five specimens lacked information on describing the Alabama group as M. marrianae, thus date of collection and were utilized for dissection pur- restricting the M. hembeli group to Louisiana. His de- poses only. The remaining lots were used for histological scription included characters of the shell only, principally investigations of gonadal activity and sexual character- the degree of sculpturing on the shell surface and the istics, as well as for anatomical studies. The collection shape of the ventral shell margin. Additional concho- dates and numbers of specimens used in the study of

logical differences between the Louisiana and Alabama gonadal activity were as follows: October 1, 1973 (1 populations were noted by Smith (1983) who pointed out specimen); October 5, 1974 (4 specimens); February 22, dissimilarities in the mantle-shell attachment scars on the 1975 (8 specimens); March 28, 1975 (4 specimens); April inner nacreous surface. 25, 1975 (3 specimens); June 21, 1975 (2 specimens); Margaritifera hembeli probably had a more extensive March 30, 1986 (5 of 16 specimens). Page 160 THE NAUTILUS, Vol. 102, No. 4

^.^:

^d.r't-'^r'

Figure 1. Photomicrograph of a transverse section through the gill of Margaritifera hcmheli showing the interlamellar junction (ILJ), 30 x

Portions of the viscera of each specimen were infil- Figure 2. Photomicrograph of a transverse section through trated with paraffin, sectioned at seven micrometer thick- the posterior portion of the mantle of Margaritifera hembeli ness, and stained with Ehrlich's hematoxylin and eosin. showing the partial!) contracted diaphragmatic septum (DS), sections were stained in a picro- ponceau connective Some which separates the exhalent (EX) and inhalent (IN) chambers, tissue stain following the method described in Humason SO X. (1979:147). At least five slides were prepared for each specimen. The barren gills of two specimens and a portion of the posterior mantle lobe of one specimen were lamellae of each gill held together b> solid, separate also sectioned in a similar manner and stained with picro- interlamellar junctions (figure 1, ILJ), lined with squa- ponceau connective tissue stain. mosal epithelium and composed of loose connective tis- Dissections were undertaken on the stomach and vis- sue and fine fibers that are more muscular appearing ceral nervous system of eight specimens. Three speci- than collagenous (in M. margaritifera. see Smith, 1979a). were investigated for gro.ss morphology of the gills, mens Interlamellar junctions for the most part arranged in nervous system, and excretory system. The method of oblique rows in typical margaritiferine fashion (Ort- dissection and exposition of specific internal organs, and mann, 1912; Smith and Wall. 1983: fig. lb), similar to the terminologv used to describe various organ compo- gills of A/. niarr!ana<'(Ortmann, 1912:235). Gill junctions nents, follows Smith (1980, 1986). are somewhat patternless along lower margin and an- .\11 material relevant to this study has been cataloged terior and posterior extremities of each gill plate. in the invertebrate collections of the Museum of Zoology Mantle lobes free all around, no indication of division (Nos. MO. 1643-1645), University of Massachusetts, Am- of exhalent region of mantle margins into .separate anal Massachusetts. herst, and supra-anal apertures. Inhalent and exhalent chambers separated posteriorly by diaphragmatic septa (figure RESULTS 2, DS), and though not observed in the living animal, are presumed to function similar to M. margaritifera Gross Anatomy (Smith. 1980) Inhalent margin of mantle with densely Anterior and posterior adductor muscles subequal, foot pigmented papillae, exhalent region pigmented, with musculature and associated pedal and retractor muscles crenulate margin. well developed Cerebral and pleural ganglia fused, kidney with both glandular and non-glandular chambers, Gonadal Activity and Sexuality renal pore and gonopore closely set but clearly separate. Labial palps falcate and large, gills or demibranchs la- .All animals examined histologicalK were sexually main shell mellar, inner gill larger than outer gill, both inner and ture, including several small specimens ranging fj9 to be from 6 to 9 outer gills free from mantle posterior to pallial line. Both length from 49 to mm (believed .

D. G. Smith 1988 Page 161

'f ^

^i V >*.. s' Figures 3-6. Photomicrographs of histological sections through male and female gonads of Margaritifera hembeli. 3. Gonad of a male specimen of M. hembeli collected in Februar\, following spawning of gametes, 65 x. 4. Gonad of a female specimen collected in February following spawning, 65 x. 5. Gonad of a male specimen of M. hembeli collected in October and showing x collected in male gametes which have filled the entire gonadal stroma, 100 . 6. Gonad of a female specimen of M. hembeli October and containing fully developed ova, 100 x years old on the basis of shell annuli). This would suggest metes takes place sometime between late November and that M. hembeli matures at an earlier age than North late December with release of larvae occurring in late American M. margaritifera (Smith, 1979b). No evidence December or January. Based on examined specimens, of hermaphroditism was observed. there is no evidence that production of glochidial larvae Although no gravid females were among the available takes place at any other part of the year. specimens, a specific reproductive cycle was indicated by the gonads of sectioned specimens. Animals collected Visceral Nerve Anatomy in Februarv', March, and April showed characteristic post- spawning features (figures 3, 4) including partial or com- In M. hembeli, the first pallial bifurcation (figure 7, BI) plete occlusion of gonadal acini by granules, presumed of the posterior nerve is well anterior of the mantle nerve pycnotic cells, and unspawned gametes in various stages separation, usually arising from the visceral ganglion it- of development or cytolysis. Animals collected in June self at a point near the origin of the posterior nerve cord showed little change from April specimens indicating (figure 7, PNC). Some variation exists in the position of that complete resorption of reproductive tissues, corre- this bifurcation relative to the visceral ganglion. The sponding to an undifferentiated stage of non-reproduc- overall pattern differs from that observed in other eastern tive activity, apparently does not occur. By early Octo- North .American margaritiferids including A/, marrian- ber, gonadal activity is resumed and sex cells, including ae. in which the first bifurcation is normally posterior of the latter stages of spermatogenesis and oogenesis, are the visceral ganglion, but similar to that observed in abundant within all observed acini (figures 5, 6). It is Cumberlandia monodonta (Smith, 1980, unpublished therefore hypothesized that the oviposition of eggs into data). Moreover, in both M. hembeli and M. marrianae, marsupial demibranchs and the spawning of male ga- the accessory nerve, which is typically present in M. Page 162 THE NAUTILUS, Vol. 102, No. 4

VG stomach roof of M. hembeli contains a well developed anterior sorting area and a continuation of the posterior sorting area extending along the right side of the stomach roof and separated from the anterior sorting area by a

ridge (figure 8).

The floor of the stomach interior (figure 9) is char-

acterized by an anterior sorting area that is somewhat

similar to M. marrianae (Smith, 1986; fig. 4b); however,

the right side sorting area of M. hembeli is completely

PNC unlike ,V/. marrianae in that a distinct platform is absent in A/, hembeli and the sorting ridges run primariK lat-

eralK , rather than in the anterior-posterior pattern seen in M. marrianae. A groove sets off the right side sorting area from the anterior sorting area in A/, hembeli and the morphologN of the stomach Door sorting areas of OES M. hembeli can be considered more similar to M. margaritifera in overall appearance than to other eastern North .\merican species.

DISCUSSION

As demonstrated by anatomical investigations of the visceral nervous system and the stomach (this studv), and conchological differences discussed elsewhere (Johnson, 1983; Smith, 1983), A/, hembeli and A/, marrianae clear-

1\ represent distinct lineages within the genus Marga- ritifera. Although both species have lateral teeth, these teeth are also present in M. auricularia (southern Europe) and A/, laosensis (southeast .\sia). Lateral teeth may therefore represent structures that have arisen separately in different "stocks of margaritiferid species, or ma> be symplesiomorphies indicative of an ancestral relationship. The presence of a corrugated surface of the posterior slope of the shell (weakK expressed in A/, hembeli) and the close geographical proximit\ of these two species are the strongest lines of evidence indicating a relationship between them. Ne\ ertheless, the pattern of \isceral nerve bifurcation and stomach morphologv in A/, hembeli and M. marrianae show interspecific variation as great as that observed between either of these two species and M. margaritifera or C. monodonta. Biochemical data of Figures 7-9. .\natoniy of the visceral nervous system and the sort de\eloped for M. margaritifera. C. monodonta, stomacli of Margarilifcra hcmbeli. 7. Visceral ganglion (VCJ) and A^. hembeli b> Davis and Fuller (1981) is not avail- of M. hemheli. 20 x. 8. Morphology of the stomach roof of able for M. marrianae, precluding a comparison of ge-

x . M. hembeli, 5 9. Morphology of the stomach floor of M. netic distances among the four species. However, the hemheli. 5 x . APR, anterior protractor muscle; ASA, anterior available biochemical data i Davis and Fuller, 1981), sorting area; .ASR, anterior sorting area of roof; BI, first pallial combined u ith anatomical information presented in this bifurcation; BN, branchial nerve; C, commissure to cerebro- stud) and elsewhere (Smith, 1980, 1986), clearK supports pleural ganglion; MN, mantle nerve; PNC, posterior nerve cord; the concept that the North .American species of the Mar- OES, esophagus; PSf\, posterior sorting area; HD, right duct; RSA, right side sorting area; T, major typhlosole. garitiferidae have been isolated from one another for a considerable period of time (Smith, 1976). margaritifera and C. monodonta (Smith, 1980), is almost aKva> s absent, being observed only once in M. marrianae ACKNOWLEDGEMENTS (Smith, unpublished data).

I thank Malcolm F. Vidrine, Mark E. Gordon, and James Stomach Anatomy E. Williams for gracioiisK- supplying me with the pre-

In addition to the posterior sorting area, which is typical served specimens used in this studv. Mark E. Gordon of other eastern North American margaritiferids, the kiiulK read an early draft of the paper. D. G. Smith 1988 Page 163

LITERATURE CITED margaritifera (Linnaeus) populations in central New En- gland. Veliger 21:381-383. Conrad, T. A. 1838. Monography of the family Unionidae, Smith, 1) (; 1980 Anatomical studies on Margari/i/era mar- or naiades of Lamarck (fresh water bivalve shells) of North garitifcra and Cumherlandia monodonia (Mollusca: Pe- America, Numbers 10, 11. Philadelphia, PA, 81-102 p. lec> poda: Margaritileridae). Zoological Journal of the Lin- Davis, S. G. M. and L. H. Fuller. 1981. Genetic relationships nean Society (i9:257-270. among Recent Unionacea (Bivalvia) of North ,\merica. Smith, D. G. 1983. On tlie so-called mantle muscle scars on Malacologia 20:217-2.53. shells of the Margaritiferidae (Mollusca: Pelecypoda). with

Humason, G. L. 1979. .Animal tissue techniques, 4th ed . W observations on mantle-shell attachment in the Unionoida H. Co., Freeman & San Francisco, 661 p. and Trigonioida. Zoologica Scripta 12:67-71. Johnson, R. I. 1983. Margaritifera marrianae, a new species Smith, D. G. 1986. The stomach anatomy of some eastern of Unionacea (Bivalvia: Margaritiferidae) the from Mo- North .American Margaritiferidae (Unionoida: L'niona- bile-.Alabama-Coosa and Escambia River systems, Ala- cea). .American Malacological Bulletin 4:13-19. bama. Occasional Papers on Mollusks (Harvard Universitv) Smith, D. G. and W. P. Wall. 1983. The Margaritiferidae 4:299-304. reinstated: a reply to Davis and Fuller (1981), "Genetic Ortmann, .\. E. 1912. Notes upon the families and genera of relationships among Recent Unionacea (Bivalvia) of North the najades. .Annals of the Carnegie Museum 8:222-.365. America." Occasional Papers on Mollusks (Harvard Uni- Smith, D. G. 1976. The distribution of the Margaritiferidae: versity) 4:321-330. a re\iew and a new synthesis Bulletin of the .American Stewart, J. H. 1988. Endangered and threatened wildlife and Malacological Union 1976:42 (abstract). plants; final endangered status for the Louisiana pearl shell Smith, D. G. 1979a. Marsupial of the anatomy demibranch "Margaritifera hembeli." Federal Register 53:3567-3570.

of Margaritifera margaritifera (Lin. ) in northeastern North \'idrine. M. F. 1985. Fresh-water mussels (Unionacea) of America (Pelecypoda: Unionacea). Journal of Molluscan Louisiana; a zoogeographical checklist of post-1890 rec- Studies 4.5:39-44. ords. The Louisiana Environmental Professional 2:50-59. Smith, D. G. 1979b Se.xual characteristics of Margaritifera THE NAUTILUS 102(4): 164-166, 1988 Page 164

The Two Printings of J. F. Gmelin's Systema Naturae, 13th Edition (1788-96)

Alan R. Rabat Richard E. Petit Deparlmenl of Mollusks Research Associate Museum of Comparative Zoology Department of Invertebrate Zoology Harvard University National Museum of Natural History Cambridge, MA 02138, USA Smithsonian Institution Washington, DC 20560, USA

One of the more important 18th century reference works printings, and noted for the Lugduni printing: type

s reset, seen . Frequently , librarx copies will in systematics is J. F. Gmelin l.'3th edition of Linnaeus' no copy have Systema Naturae. This work represented a considerable the same date stamped on the binding of each part or updating of and expansion upon the 12th edition of the hand-written on the first page of each part; this is in- Stjstema Saturae (Linnaeus, 1767). Gmelin not only pro- correct. The important facts are that the Lugduni print-

\ ided additional bibliographic references for the species ing was published after the Lipsiae printing, and was described by Linnaeus, but he also described numerous printed from reset type w hich resulted in a number of new species, using the same format as had Linnaeus. differences due to printer s error. Therefore, the date of Gmelin's magnum opus was published in three volumes, publication of each part of the Lugduni printing must comprising 10 parts altogether: Regntim Animalium (7 post-date that of the relevant part of the Lipsiae printing. parts); Regnum VegetaMe (2 parts); and Regnum La- L'nfortunately , we have not been able to determine pideuni (1 part). For comments on the structure and the date of publication for the succeeding parts of the content of Gmelin's work, see Dodge (1958:157-158). Lugduni printing (only for the first part in the multi- Few systematists, however, have realized that Gme- part volumes). However, we present this comparison here lin's work actually underwent two separate printings. in order to alert s\ stematists to the existence of these two The initial German printing was by Georg. Emanuel. printings. Should the reader know of further bibliograph- Beer of Lipsiae [= Leipzig]; the later French printing ical sources bearing on this matter, we would greatly was by J. B. DelamoUiere (subsequently as Bernuset, appreciate hearing from them.

Delamolliere, Falque et Soc. ) of Lugduni [= Lyon]. Both, Herein we present as complete a collation as possible of course, were written in Latin. of the two printings of Gmelin's Systema Saturae. based Hopkinson (1907) provided a valuable collation of the on Hopkinson (1907) and on our examination of complete Lipsiae printing, based on an analysis of various German sets of this work in the libraries of the Museum of Com- literature abstracts and review publications. He noted parative Zoology , the British Museum (Natural History), that only the first part in the two multi-part volumes had and the Museum National d Histoire Naturelle, Paris. a date on the title page and that this date "did not apply to the whole of the parts in that \olume " (Hopkinson, Gmelin, jo. Prid. [Joliann P^iedrich]. 1788-1793. Ca- 1907:1036), since the later parts were published in fol- roli a Linne, Systema Naturae per Regna Tria Naturae, lowing years. Hopkinson did not refer to the Lugduni Secundum Classes, Ordines, Genera, Species, cum cha- printing. Agassiz (1852:69) listed this work as "Leipz. ractcribus, diftcrentiis, synonymis, locis. Editio Decima

1788-1793,. . . Lugduni, 1789-1796." Engelmann (1846: Tertia, Aucta, Reformata. Georg. Emanuel. Beer, Lip-

103) provided the same dates as had Agassiz, but erro- siae; [Second printing, 1789-1796], J. B. Delamolliere, neously gave the place of publication as "Lugd. Batav." Lugduni. which is actually Leiden [= Lugduni Batavorum]. Sher- born (1902:.\.\xv) gave the same dates, made the same Tomus 1, REGNUM ANIMALIUM geographical transposition as had Engelmann, and spec- ified that the second printing was a reprint of the first. 1(1) Mammalia to Aves Picae, pp. xii -I- 1-500 Hulth (1907:13) and Soulsby (1933:16, 50) provided a Lipsiae, 1788; Lugduni, 1789. partial collation ol these two printings; however, their 1 (2) Aves Anseres—A\es Passeres. pp. 501-1032 dates are based solely on those stated on the title page Lipsiae, 1789; Lugduni (post 1789?). for the first part of each of the two multi-part volumes. I (3) Amphibia— Pisces, pp. 1033-1516 Stafleu and Cowan (1976:956) briefly mentioned the two Lipsiae, 1789; Lugduni (post 1789?). ,

A. R. Kabat and R. E. Petit 1988 Page 165

I (4) Insecta General, Coieoptera—Hemiptera, pp. 1517- treated as a reprint (or reissue) in which errors occur, 2224 and not as a "second edition "; (3) systematists who are Lipsiae, 1790; Lugduni (post 1790?). studying species described by Gmeiin should compare

I (5) Insecta Lepidoptera — Aptera, pp. 2225-3020 the two printings in order to uncover any discrepancies Lipsiae, 1790; Lugduni (post 1790?). with respect to species descriptions or references. 1(6) Vermes, pp. 3021-3910 Lipsiae, 1791; Lugduni (post 1791?). ACKNOWLEDGEMENTS 1(7) Inde.x 1-3, pp. 3911-4120 Lipsiae, 1792; Lugduni (post 1792?), Mr. Andre Verhecken, Mortsel, Belgium, first brought the discrepancy in the Gmeiin citations to our attention, and then followed up with information on the existence Tomus II, REGNUM VEGETABILE of the two printings. His interest and assistance is greatly appreciated. We thank the librarians of the Museum of II (1) Monandria to Polyandria, pp. .\1 + 1-884 (Natural His- Lipsiae, 1791; Lugduni, 1796. Comparative Zoology, the British Museum tory), and the Museum National d'Histoire Natureile, II (,2) Didynamia to Cryptogamia, pp. 885-1662 us to examine the sets of Gmelin's work in Lipsiae, 1792; Lugduni (1796 or later?). who allowed their collections. A number of other hbraries were checked for this work; most had only partial sets of one or the Tomus 111, REGNUM LAPIDEUM other printing which did not yield additional useful information. Prof. Kenneth Boss provided a helpful re- 111, pp. 1-476, plates 1-3 (of crystals) J. Lipsiae, 1793; Lugduni. 1796. view of this manuscript.

It has been our experience that molluscan taxonomists who have used Gmelin's work have usually referred sole- LITERATURE CITED ly to the Lipsiae printing. Among others, these include Agassiz, L. 1852. Bibliographia Zoologiae et Geologiae, Vol. (1958:157-158); and Kohn (1966) who provided Dodge 3 (of 4). Ray Society, London, 657 p. biographical information on Gmeiin and a useful critique Dodge, H. 1958. .\ historical review of the mollusks of Lin- of his work. .\n example of the taxonomic problems that naeus. Part 6. The genus Trochus of the class Gastropoda. can arise with respect to the two printings of Gmelin's Bulletin of the American Museum of Natural History

\\ ork is that of certain species of Cancellariidae (Gastrop- 116(2):153-224. oda) reviewed by Petit (1984). For Buccinum piscato- Engelmann, W. 1846. Bibliotheca Historico-Naturalis. Ver-

zeichniss der Biicher iiber Naturgeschichte . . . Erster Band. rium Gmeiin. 1791 (p. 3496), the Lipsiae printing gave 89"' Verlag von Wiihelm Engelmann, Leipzig, x -I- 786 p. two references: "List. Conch, t. 1024 f. and "Martin. of the separate parts Hopkinson, J. 1907. Dates of publication Conch. 4. t. 124 f. 1151, 1152". Based on this, Petit chose of Gmelin's edition (13th) of the 'Systema Naturae' of the figures from the latter reference (i.e., Chemnitz, J. Linnaeus. Proceedings of the Zoological Society of London H., 1780, Neues Systetnatisches ConchyHen-Cabinet 1907(49):1035-1037. \"ol. 4, Nurnberg) to be representative of this species. materiaux Hulth, J. M. 1907. Bibliographia Linnaeana: pour Subsequently, A. Verhecken [in litt.) inquired as to how servir a une Bibliographie Linneenne. Kungliga Veten- the Chemnitz illustration could be used in this manner skaps Societeten I Upsala, 170 p. -H 11 pis. 1954. since it was not cited by Gmeiin. In later correspondence, International Commission on Zoological Nomenclature, N'erhecken advised that he had found that he was re- Opinion 296. Suppression, under the Plenary Powers, for nomenclatorial purposes, of volume 3 (Regnum Lapi- ferring to a cop)- of the Lugduni printing. In that printing deum) of the Twelfth Edition of the Systema Naturae of only one reference, the aforementioned "List. Conch, t. Linnaeus published in 1768 and of the corresponding vol- 1024 f. 89" (i.e.. Lister, M.. 1692/1770, Historiae sive ume in the Houttuyn (1785), Gmeiin (1793) and Turton Synopsis Methodicae Conchyliorum, London), was giv- U806) editions of the above work. Opinions and Decla- in question; the Chemnitz reference en for the species rations rendered by the International Commission on Zoo- being omitted. ObviousK', the omission of the reference logical Nomenclature 8(13):167-178. to Chemnitz was a printer's error, as a comparison of the identity of the de- Kohn, A. J. 1966. Type specimens and two printings makes it evident that the type was reset scribed species of Conus III. The species described by and errors were made. Gmeiin and Blumenbach in 1791. The Journal of the Lin- Incidentally, the third volume of this work (the Reg- nean Societv of London, Zoology 46(308):73-102, pis. 1-3. num Lapideum) was placed on the Official Index of Linnaeus, C. 1766-67. Systema Naturae, per Regna Tria Na- Rejected and Invalid Works in Zoological Nomenclature Sal- turae, . . . Editio Duodecima Reformata. Direct. Laur. (along with the related sections of Linnaeus and of Tur- vii, Holmiae. Regnum Animale, 1328 -t- 36 p. ton), since the "generic names of fossil animals used in Petit, R. E. 1984. Some early names in Cancellariidae. Amer- these works actually corresponded to the classes of Recent ican Malacological Bulletin 2:57-61. animals (ICZN, 1954, Opinion 296). Sherborn, C. D. 1902. Index Animalium sive index nominum We conclude that (1) since the Lipsiae printing was quae ab .\.D. MDCCL\'III generibus et speciebus ani-

published first, it should serve as the basis for discussion malium imposita sunt . . Sectio Prima. L niversity Press, of Gmelin's species; (.2) the Lugduni printing should be Cambridge, Ix + 1195 p. Page 166 THE NAUTILUS, Vol. 102, No. 4

Soulsby, B. H. 1933. A catalogue of the works of Linnaeus Stafleu, F. A. and R. S. Cowan. 1976 [in 1976-86]. Taxonomic (and publications more immediately relating thereto) pre- literature: a selective guide to botanical publications and served in the libraries of the British Museum (Bloomsbury) collections with dates, commentaries, and types. Second and the British Museum (Natural History) (South Ken- Edition. Regnum X'egetabile; Bohn. Scheltema & Holke-

sington). British Museum (Natural History), London, 2nd ma, Utrecht, \'ol. 1 (of 6):xl + 1136 p.

ed., p. xii -I- 1-312; 7 pis. INSTRUCTIONS TO AUTHORS

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