ó^S PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON ll8(2):344-366. 2005. Cocculinid and pseudococculinid limpets (Gastropoda: Cocculiniformia) from off the Caribbean coast of Colombia Néstor E. Ardila and M. G. Harasewych (NEA) Museo de Historia Natural Marina de Colombia, Instituto de Investigaciones Marinas, INVEMAR, Santa Marta, A.A. 1016, Colombia, e-mail: [email protected]; (MGH) Department of Invertebrate Zoology, MRC-I63, National Museum of Natural History, Smithsonian Institution, Washington, D.C. 20013-7012 U.S.A., e-mail: [email protected] Abstract.•The present paper reports on the occurrence of six species of Cocculinidae and three species of Pseudococculinidae off the Caribbean coast of Colombia. Cocculina messingi McLean & Harasewych, 1995, Cocculina emsoni McLean & Harasewych, 1995 Notocrater houbricki McLean & Hara- sewych, 1995 and Notocrater youngi McLean & Harasewych, 1995 were not previously known to occur within the of the Caribbean Sea, while Fedikovella beanii (Dall, 1882) had been reported only from the western margins of the Atlantic Ocean, including the lesser Antilles. New data are presented on the external anatomy and radular morphology of Coccocrater portoricensis (Dall & Simpson, 1901) that supports its placement in the genus Coccocrater. Coc- culina fenestrata n. sp. (Cocculinidae) and Copulabyssia Colombia n. sp. (Pseu- dococculinidae) are described from the upper continental slope of Caribbean Colombia. Cocculiniform limpets comprise two paraphyletic, with the Cocculinoidea related groups of bathyal to hadal gastropods with to Neomphalina and the Lepetelloidea in- global distribution that live primarily on cluded within Vetigastropoda (Ponder & biogenic substrates (e.g., wood, algal hold- Lindberg 1996, 1997; McArthur & Hara- fasts, whale bone, cephalopod beaks, crab sewych 2003). carapaces, or shark egg cases) that sink Among the ten families of cocculiniform from shallow water into the aphotic zone limpets, the best known and by far the most (Haszprunar 1988, 1998; Lesicki 1998). diverse are the Cocculinidae (Cocculinoi- Early records were assigned to the newly dea) and the Pseudococculinidae (Lepetel- proposed families Cocculinidae Dall, 1882 loidea). Both are associated with sunken and Lepeteilidae Dall, 1882. Subsequent re- wood and plant material (Haszprunar 1988, searchers discovered these gastropods to be Lesicki 1998), which tend to be encoun- both diverse and distinctive, apportioning tered more often than other biogenic sub- the many newly collected deep sea species strates (Leal & Harasewych 1999). Based between two superfamilies, Cocculinoidea on the primitive organization of the alimen- and Lepetelloidea, containing two and eight tary systems of the Cocculinidae and Pseu- families respectively (see Haszprunar 1998, dococculinidae, Haszprunar (1988:74) sug- Lesicki 1998, Strong et al. 2003 and refer- gested that a diet of wood was likely the ences therein). These limpets had been unit- primitive condition, and that adaptations to ed in the superorder Cocculiniformia (Hasz- other biogenic substrates were "probably prunar 1987, 1998) but recent phylogenetic the main driving force of the cocculiniform studies have shown Cocculiniformia to be radiation." VOLUME 118. NUMBER 2 345 The western Atlantic fauna of cocculini- croscope (SEM). For specimens in which form limpets contains the first cocculino- the preservation was adequate, bodies were idean and lepetelloidean species to be dis- critical-point dried, coated with carbon and covered (Dall 1882, Verrill 1880). While gold, and the -.xternal anatomy examined this fauna is widespread and diverse, pub- with SEM. lished records from within the margins of The specimens on which this study is the Caribbean tectonic plate are rare. Re- based are deposited in the Museo de His- cords of Cocculinidae are limited to reports toria Natural Marina de Colombia- of two species•Cocculina rathbuni Dall, MHNMC (INV MOL), Santa Marta, and 1882 and Coccocrater portoricensis (Dall the National Museum of Natural History, and Simpson, 1901)•from bathyal depths Smithsonian Institution, Washington, D.C. off the Caribbean coast of Colombia (Diaz (USNM). & Puyana 1994). Among the Pseudococ- Abbreviations.•P = distance from an- culinidae, Notocrater pustulosa Woodring, terior shell edge to apex; SH = shell height; 1928 non Thiele, 1925, was described from SL = shell length; SW = shell width (see the Miocene of Jamaica, whereas Amphi- McLean & Harasewych 1995: fig. 1). plica venezuelensis McLean, 1988 is known only from abyssal depths in the Venezuela Systematics Basin. Recent explorations along the outer shelf Superfamily Cocculinoidea Dall, 1882 and upper continental slope of the Carib- Family Cocculinidae Dall, 1882 bean coast of Colombia during cruises of Genus Cocculina Dall, 1882 INVEMAR-MACROFAUNA I and II in Cocculina rathbuni Dall, 1882 1998-2001, resulted in the collection of a Figs. 1, 2 number of considerable range extensions Cocculina rathbuni Dall, 1882:402.•Dall, and new species from the southwestern Ca- 1889:347, pi. 15, figs. 5, 7.•Pilsbry, ribbean Sea. The present paper reports on 1890:132, pi. 25, figs. 5, 6 [copy of the members of the limpet families Coc- Dall].•Dall, 1908:340.•Thiele, 1909:6, culinidae and Pseudococculinidae that were pi. 2, figs. 1, 2.•Abbott, 1974:34, fig. collected as part of this survey. Range ex- 192.•McLean, 1987:325, figs. 1-4.• tensions and supplemental data are provid- Haszprunar, 1987:321.•Diaz & Puyana, ed for previously known species, and one 1994:113, pi. 35, fig. 345 [copy of new species is described in each of the two Dall].•McLean & Harasewych, 1995:8, families treated. figs. 2-11.•Lesicki, 1998:51. Materials and Methods Material examined.•INV MOLÍ 672, 1 specimen (SL = 6.92, SW = 5.37, SH = Samples were collected using a bottom 1.92 mm), Sta. 75, off Rosario Island trawl (9 X 1 m opening, 16 m in length) (10°09'N, 76°00'W), in 296 m. during cruises of INVEMAR-MACRO- Description.•Shell (Fig. 2A) medium to FAUNA I and II, 1998-2001, aboard the R/ large (SL = 7 mm), thin, white, elliptically V Ancon, working off the Caribbean coast conical, low (SH/SL =^ 0.30). Apex slightly of Colombia at depths of 20-500 m (Fig. posterior of center (P/SL == 0.55). Proto- 1). Specimens of Cocculinidae and Pseu- conch (Fig. 2B, C) just below apex, 260 |xm dococculinidae were sorted and stored in in length, with posteriormost portion envel- 70% ethanol, sometimes with fragments of oped by posterior slope of shell. Protoconch wood on which they were collected. Sub- sculpture of raised cords forming irregular sequently, shells and radulae were dissected reticulate pattern that is aligned in rows, and examined using a scanning electron mi- broader laterally. Surface within retícula- 346 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 7C'0-0-W 74'0'0"W ITOO-^N N A •9-' 7i •r i/ 2 P - o / Q - ^_^' <M •jg,36 115 A, deXîcniza RosarioiÈ'^ z P • D 75 j^ , O («W .T,qua b »50-^ San ' ., 753 Bernardo Is .-Tolu 63-^ ; 'E. de fa Radâ Iff/i «-sSr"^ *P Escondioo , "Arboletes 50 S5 a 50 tCO ''\ ] COLOMBIA Kilome ras 2 b b 3 cc » 76°0 0-W 74"00-W 72-0'0'W Fig. I. Location of stations off the Caribbean coast of Colombia at which cocculiniform gastropods were collected during cruises of INVEMAR-MACROFAUNA 1 and II. 1998-2001, aboard the R/V Amon. Stations range in depth from 70 to 516 m. tions finely pitted (Fig. 2D). Teleoconch port this species from the southwestern Ca- sculpture of prominent, raised concentric ribbean Sea, off the coast of Colombia. growth lines and weak, radial striae. Ante- Their records are from the same area off the rior and posterior slopes nearly straight. Colombian coast but from greater depths Aperture not planar, likely reflecting shape (560-900 m). of substrate. Shell edge thin. Periostracum thin. Pigmented eyes absent. Cocculina messingi McLean & Range.•Off Massachusetts, Florida, Ba- Harasewych, 1995 hamas, Martinique, St. Vincent, Barbados, Figs. 1, 3, 4 Colombia (Dall 1889, Diaz & Puyana 1994, Cocculina messingi McLean & Harasew- McLean 1987, McLean & Harasewych ych, 1995:11, figs. 12-24.•Lesicki, 1995) at depths of 124-1127 m. 1998:51. Remarks.•A. detailed synonymy, as well Cocculina sp. 1 Ardila & Harasewych, as illustrations of the shell, protoconch, ex- 2002. ternal anatomy, and radular morphology of this species are provided by McLean & Material examined.•INV MOL 1661, 1 Harasewych (1995:8-11, figs. 2-11). Diaz specimen (SL = 5.80, SW = 4.59, SH = & Puyana (1994:113) were the first to re- 1.87 mm), Sta. 46, off Punta Gloria VOLUME 1 18, NUMBER 2 347 Fig. 2. Cocculhui ivlliinmi Dali. 1882. INV MOL1672. Sla. 15. off Rosario Island (IO°09'N, 76°00'W), in 296 m. Scanning electron micrographs (SEM) of shell. A. dorsal, left lateral, and ventral views of shell; B. dorsal and C. left lateral views of protoconch; D. detail of protoconch .sculpture. (1I°II'N, 74°]7'W), in 282 m. USNM 75, off Rosario Island (10°09'N. 76°00'W), 1008179, 3 specimens (SL = 10.9, SW = in 296 m. INV MOL3437 1 specimen (SL 8.9, SH = 3.3 mm), INV MOL 1677, 1 = 7.13, SW = 5.61, SH = 2.73 mm), Sta. specimen (SL = 10.36, SW = 8.64, SH = 36, off Neguanje. (11°24'N, 74°09'W), in 3.23 mm), and INV MOL 1678, 1 specimen 296 m. (SL = 10.08, SW = 8.56, SH = 3.28 mm), Description.•Shell (Figs. 3, 4A) medi- Sla. 56, off Arboletes (9°02'N, 76°36'W), um to large (SL to 10.9 mm), thin, white, in 290 m. INV MOL 1665, 1 specimen (SL elliptically conical, low to moderate in = 6.20, SW = 4.78, SH = 1.87 mm), Sta. height (SH/SL = 0.37-0.41). Apex slightly 348 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON Fig. 3. Cacculina messinai McLean & Harasewych, 1995. INV MOLI665, Sta. 75, off Rosario Island C10°09'N, 76°00'W), in 296 m.