Part 28 Benthic, Shelled Gastropods Leonora Rodríguez-Sevilla, Rita Vargas, and Jorge Cortés Cyphoma gibbosum on a seafan in the coral reef of Parque Nacional Cahuita, Caribbean of Costa Rica (Photo: Jorge Cortés) I.S. Wehrtmann, J. Cortés (eds.) Marine Biodiversity of Costa Rica, Central America, © Springer Science + Business Media B.V. 2009 333 334 L. Rodríguez-Sevilla et al. Abstract Here we present a compilation of 1,198 benthic, shelled gastropod (prosobranch and pulmonate) species records from Costa Rica published in the scientific literature. Of these, 389 species are from the Caribbean and 818 from the Pacific; 9 species are common to both coasts. In terms of families, the Caribbean has 63, while the Pacific has 75. Including a list of 84 unpublished species (identified by specialists), the total of benthic, shelled gastropods for Costa Rica is 1,282, and the number of species from the Pacific coast increases to 902. Compared to a study of Colombia’s Caribbean coast, our list includes 60% of the species reported there, but only 31% of the 2,682 species reported for the Panamic province. The difference in species number between the Caribbean and Pacific of Costa Rica may be due to several facts, including the length of the coast (the Caribbean coast is five times shorter), number of expeditions (many more on the Pacific), and difference in coastal morphology (the Caribbean less complex). Compared to other areas, Costa Rica has lower species numbers, which may be due to patchy sampling (mostly in shallow waters with few samples from deeper than 100 m), and to the study of only macromolluscs. All of this indicates that more sampling is needed on both sides of Costa Rica and in the other Central American countries. Introduction Molluscs are the second most specious group of invertebrates in the world, and three fourths of them are gastropods (Morton 1960; Ruppert & Barnes 1994). Mollusc collecting has been popular for centuries, and amateurs have contributed significantly to the advancement of mollusc studies. There are inmense collections both in museums and in private hands. Molluscs are also important as food source, for the curio trade and in some traditional practices (use of dyes or the shells themselves, e.g., as musical instruments). The Caribbean and Pacific coasts of Costa Rica measure approximately 212 and 1,254 km in length, respectively, totaling 1,466 km. The Caribbean coast of Costa Rica is relatively straight, with strong surf, sandy beaches on the north section, and only two small mangrove areas, one in the central section and one in the south; tide range is less than 50 cm. The southern section, from Limón to the border with L. Rodríguez- Sevilla Museo de Zoología, Escuela de Biología, Universidad de Costa Rica, 11501-2060 San Pedro, San José, Costa Rica. e-mail: leonorars@racsa.co.cr R. Vargas Apdo. 3144-1000, San José, Costa Rica. e-mail: ritav@cariari.ucr.ac.cr J. Cortés Centro de Investigación en Ciencias del Mar y Limnología (CIMAR), and Escuela de Biología, Universidad de Costa Rica, 11501-2060, San José Costa Rica. e-mail: jorge.cortes@ucr.ac.cr 28 Benthic, Shelled Gastropods 335 Panama, is formed by rocky headlands of coralline origin, alternating with sandy beaches. In that section, coral reefs with lagoons and seagrass beds predominate (Vargas & Cortés 1999a; Coll et al. 2001; Cortés & Jiménez 2003a). The Pacific coast is longer and more irregular. It has different types of environments such as estuaries, mangroves, muddy bottoms, rocky areas, sandy beaches, and coral reefs, and the tides reach up to 4 m (Vargas & Cortés 1999b; Cortés & Jiménez 2003b). Malacological Studies in Costa Rica The first marine invertebrate species described for Costa Rica belongs to the phylum Mollusca; it was the polyplacophora Chaetopleura scabricula Sowerby, 1832. The first species described from Costa Rica came from Puerto Potrero (now known as Bahía Potrero), Guanacaste, and from the Golfo de Nicoya and Golfo Dulce, Puntarenas (Cortés 1998). Other mollusc species were described, thanks to the collections done by Hugh Cuming between 1827 and 1830, along the west coast of the Americas. The majority of the molluscs collected in Costa Rica are the product of expeditions organized by foreigners, especially from the USA, from the late 19th century to the mid-20th century. The main objective of these expeditions was to explore the coasts of the Americas. One of the most visited sites by these expeditions was Isla del Coco, since it is en route to the Galapagos Islands, and the focus of attention of many naturalists (Hertlein 1963; Cortés & Jiménez 2003b). The Caribbean coast was not studied until the 1960s. Bakus (1968a) did a comparative study of the littoral zonation of gastropods collected in 1962 at three sites on the Caribbean: Portete, Puerto Limón, and Isla Uvita; he reported 29 species of gastropods. Houbrick (1968a) studied the marine molluscs in 1966 at two localities (Portete and Barra del Colorado), in shallow waters and in the littoral zone; he collected 250 species of molluscs of which 181 were gastropods. Wellington (1974) studied the fauna of Cahuita in 1972 and reported 70 species of gastropods. Robinson & Montoya (1987) collected materials between 1982 and 1986 from the intertidal to 5 m depth in ten locations distributed along the coast: Barra del Colorado, Río Matina, Moín, Portete, Puerto Limón, mouth of Río Banano, Río Estrella, Cahuita National Park, Puerto Viejo, and Manzanillo. They collected 395 species of which 288 were gastropods. Espinosa & Ortea (2001) collected from Cahuita to Gandoca between 1999 and 2001, and listed 484 molluscs of which 340 were gastropods. Finally, Rodríguez-Sevilla et al. (2003) published a compedium of the marine gastropods of the Caribbean coast of Costa Rica; they reported 384 species, plus 106 additional in two appendixes, for a total of 490 species. Many papers have been published on molluscs of the Pacific of Costa Rica, including Isla del Coco. We can divide the publications into three periods. The first publications are descriptions of species based on specimens collected by ship captains or by people who collected for museums during the early part of the 19th century (e.g., Sowerby 1832a, b). The second period, from the late 19th to the mid20th century, produced a large series of papers based on scientific expeditions to the eastern Pacific, e.g., Dall (1900) with the US Fish Commission Expedition of 336 L. Rodríguez-Sevilla et al. 1888 on board the steamer Albatross. Some of the most productive expeditions were organized by the New York Zoological Society, described by Beebe (1938). An example of one of many papers on molluscs is by Hertlein & Strong (1943). Another productive series of expeditions were the Allan Hancock Expeditions to the eastern Pacific, presented by Fraser (1943). For an example of a paper, among many, on molluscs see Hanna & Hertlein (1938). The third period is from the 1950s to the present date when individuals have done collections, e.g., Jung (1989) studied Strombina and published on them. Others collected as part of expeditions; e.g., Høisæter (1998) participated in the Costa Rican–German Expedition and published on the gastropods of the Golfo Dulce. Many papers have been published on the molluscs of Isla del Coco as a result of expeditions between 1983 and 1987 (e.g., Montoya & Shasky 1987; D’Attilio et al. 1987; Montoya 1988; Montoya & Kaiser 1988; Kaiser 1998, 2001a, 2002; Kaiser & Hertz 2001). During the review of this part, Carol Skoglund sent us a list of 84 species in 59 genera and 20 families of gastropods (Table 28.1) collected during three dredging Table 28.1 List of species collected during three dredging trips (depth: 10–37 m) to Playa del Coco, Guanacaste, by Carol Skoglund and her late husband Paul. These are now, with a few exceptions, in the Skoglund collection, Phoenix, Arizona. This collection will go to the Santa Barbara Museum of Natural History. The species from the late Robert Koch’s collection are now at the Santa Barbara Museum of Natural History. The Philadelphia Academy of Sciences (ANSP) has duplicate specimens of all turrids listed, plus the six species shown as ANSP which are not in the Skoglund collection. The list and information was provided by Carol Skoglund (C. Skoglund, personal communication 2005) Family FISSURELLIDAE Lucapinella eleanorae McLean, 1967 Leurolepas roseola McLean, 1970 Family TROCHOIDAE Calliostoma leanum (C.B. Adams, 1852) Calliostoma rema Strong & Hertlein, 1933 Family TURBINIDAE Arene balboai (Strong & Hertlein, 1939) – SBMNH Family RISSOIDAE Rissoina axeliana Hertlein & Strong, 1951 – SBMNH Family CALYPTRAEIDAE Crucibulum castellum Berry, 1963 Family NATICIDAE Natica othello Dall, 1908 Polinices caprae (Philippi, 1852) Sinum debile (Gould, 1853) Family TRIVIIDAE Hespererato columbella (Menke, 1847) – SBMNH Family MURICIDAE Pterynotus pinniger (Broderip, 1833) Folinia ericana (Hertlein & Strong, 1951) Folinia insignis (de Folin, 1867) Dermomurex indentata (Carpenter, 1857) Favartia perita (Hinds, 1844) Trachypollia lugubris (C.B. Adams, 1852) Coralliophila orcuttiana Dall, 1919 Family CAECIDAE Caecum parvum C.B. Adams, 1852 – SBMNH Caecum lohri (Strong & Hertlein, 1939) – SBMNH Family BUCCINIDAE Caducifer nigricostatus (Reeve, 1846) (continued) 28 Benthic, Shelled Gastropods 337 Table 28.1 (continued) Family TURRITELLIDAE Turritella mariana Dall, 1908 Family VERMETIDAE Serpulorbis oryzata (Mörch, 1862) Family EPITONIIDAE Epitonium minuticostatum (Deboury, 1912) Epitonium togatum Hertlein & Strong, 1951 Epitonium hindsii (Carpenter, 1856) Opalia spongiosa Carpenter, 1864 Family EULIMIDAE Scalenostoma rangii (de Folin, 1867) Sincola dorsata (Sowerby, 1832) – SBMNH Family NASSARIIDAE Nassarius corpulentus (C.B. Adams, 1852) Nassarius pagodus (Reeve, 1844) Nassarius nucleolus (Philippi, 1846) Family OLIVIDAE Oliva kaleontina Duclos, 1835 Monostiolum crebristriatum (Carpenter, 1856) Phos veraguensis Hinds, 1843 Phos dejanira (Dall, 1919) Phos cumingii Reeve, 1846 Family COLUMBELLIDAE Columbella haemastoma Sowerby, 1832 Mitrella harfordi Strong & Hertlein, 1937 Nassarina helenae Keen, 1971 – SBMNH Nassarina perita Keen, 1971 Steironepion tincta (Carpenter, 1864) Strombina carmencita Lowe, 1935 Strombina elegans (Sowerby, 1832) Strombina maculosa (Sowerby, 1832) Strombina recurva (Sowerby, 1832) Sincola gibberula (Sowerby, 1832) Cochlespira cedonulli (Reeve, 1843) Pyrgospira obeliscus (Reeve, 1843) Crassispira cortezi Shasky and Campbell, 1964 Crassispira brujae Hertlein & Strong, 1951 Crassispira epicasta Dall, 1919 Crassispira adana (Bartsch, 1950) – ANSP Crassispira nigerrima (Sowerby, 1834) – ANSP Crassispira tepocana Dall, 1919 Hindsiclava militaris (Reeve, 1843 ex Hinds, MS) – SBMNH and ANSP Olivella gracilis (Broderip & Sowerby, 1829) Family CANCELLARIIDAE Aphera tessellata (Sowerby, 1832) Narona exopleura (Dall, 1908) – SBMNH Tritonoharpa siphonata (Reeve, 1844) Family TEREBRIDAE Terebra puncturosa Berry, 1961 Terebra argosyia Olsson, 1971 Family TURRIDAE Calliclava albolaqueata (Carpenter, 1865) Imaclava unimaculata (Sowerby, 1834) Agladrillia pudica (Hinds, 1843) Drillia acapulcana (Lowe, 1935) Cerodrillia cybele (Pilsbry & Lowe, 1932) – SBMNH Gemmula hindsiana Berry, 1958 Miraclathurella bicanalifera (Sowerby, 1834) Carinodrillia dichroa Pilsbry & Lowe, 1932 Carinodrillia halis (Dall, 1919) – ANSP Strictispira ericana (Hertlein & Strong, 1951) Compsodrillia alcestis (Dall, 1919) Compsodrillia duplicata (Sowerby, 1834) Compsodrillia haliplexa (Dall, 1919) Compsodrillia jaculum (Pilsbry & Lowe, 1932) Compsodrillia olssoni McLean & Poorman, 1971 – ANSP Nannodiella fraternalis (Dall, 1919) Glyphostoma candida (Hinds, 1843) Kurtziella cyrene (Dall, 1919) – ANSP Tenaturris verdensis (Dall, 1919) Daphnella bartschi Dall, 1919 – SBMNH Philbertia doris Dall, 1919 – SBMNH Microdaphne trichodes (Dall, 1919) – SBMNH 338 L. Rodríguez-Sevilla et al. Fig. 28.1 Prunum holandae from the Caribbean coast of Costa Rica (Photo: Paul V. Scott) Fig. 28.2 Jenneria pustulata from Pacific Costa Rica (Photo: Yolanda Camacho) trips to Playa del Coco (depth: 10–37 m) by herself and her late husband Paul, which were not included in her publication (Skoglund 1990a). Herein we report 1,198 benthic, shelled gastropod (prosobranch and pulmonate) species present in both coasts, 389 from the Caribbean (Fig. 28.1) and 818 from the Pacific (Fig. 28.2), with nine species in common. In terms of families, the Caribbean has 63, while the Pacific has 75. Including the species in Table 28.1, the total of benthic, shelled gastropods for Costa Rica is 1,282, and the number of species from the Pacific coast increases to 902. The systematics used in the present part are based on Abbott (1974), Skoglund (1992, 2002), and Beesley et al. (1998). 28 Benthic, Shelled Gastropods 339 Comparative Species Richness A recent report by Díaz Merlano and Puyana Hegedus (1994) on the Caribbean and western Atlantic of Colombia revealed 653 species of benthic, shelled gastropods. The number of species reported herein for the Caribbean constitutes 60% of the above-mentioned number from Colombia. Keen (1971) and Skoglund (2002) reported 2,682 for the Panamic province; our number for the Pacific coast comprised 31% of the Panamic species. The difference in number of species between both Costa Rican coasts and between Costa Rica and the other regions sampled could be explained by several reasons: 1. The Caribbean coast is about five times shorter than the Pacific coast. 2. The collecting efforts have been greater in the Pacific. 3. The morphological differences between both coasts make the Pacific a more heterogeneous region with more diversity in habitats and, hence, more biodiversity. 4. The majority of the sites sampled in the Caribbean coast are located in the central and southern sections of the coast; the northern part has been poorly sampled. However, diversity must be higher in the southern and central areas of the coast, due not only to more intense sampling, but also to the larger variety of habitats in the south. 5. The majority of the collections were made on the shore or by diving in shallow waters; thus, the deep water fauna remains unknown. 6. Most species reported in the literature are macromolluscs. Micromolluscs have received little attention due to the difficulties associated with their identification. In other areas, however, the study of the micromolluscs has received more interest lately, revealing a large number of new species and families. 7. Sampling of the Pacific coast has been patchy and mostly in shallow waters, there are few samples of waters deeper than 100 m. For all these reasons it is concluded that more sampling is needed on both sides of Costa Rica. Therefore, the number of species presented in this part is probably an underestimation of what really exists. Species Lists 28.1 and 28.2, included on the CD-Rom, are based on the literature and museum collections in Costa Rica. In the collection of the Museo de Zoología of the Universidad de Costa Rica, there is a number of specimens that have not been identified, and some might be new species; these have not been included in the lists. Specialists Henry Chaney: Director of Collection and Research, Santa Barbara Museum of Natural History, Santa Barbara, California, USA – Eastern Pacific. hchaney@ sbnature2.org Kirstie L. Kaiser: Museum Associate, Los Angeles County Museum of Natural History, Los Angeles, California, 90007, USA – Eastern Pacific. 76511.3463@ compuserve.com 340 L. Rodríguez-Sevilla et al. José Espinosa: Instituto de Oceanología de Cuba, La Habana, Cuba – Caribbean. espinosa@oceano.inf.cu Michel Montoya: San José, Costa Rica – Eastern Pacific. michelmontoya@correo.co.cr Carol Skoglund: Santa Barbara Museum of Natural History, Santa Barbara, California, USA – Eastern Pacific. carolskoglund@msn.com Collections ● ● ● ● ● ● ● ● ● ● Museo de Zoología, Escuela de Biología, Universidad de Costa Rica: http:// museo.biologia.ucr.ac.cr/ Academy of Natural Sciences, Philadelphia: http://www.acnatsci.org/ California Academy of Sciences, San Francisco: http://www.calacademy.org/ Instituto de Oceanología, Cuba: http://www.cuba.cu/ciencia/citma/AMA/p_a.html Los Angeles County Museum of Natural History, California: http://www.nhm. org/ National Museum of Natural History, Smithsonian Institution, Washington, DC: http://www.mnh.si.edu/ Natural History Museum, London: http://www.nhm.ac.uk/ Naturhistorisches Museum, Basel, Switzerland: http://www.nmb.bs.ch/ San Diego Natural History Museum, California: http://www.sdnhm.org/ Santa Barbara Museum of Natural History, California: http://www.sbnature. org/ Acknowledgments We are grateful to José H. Leal for his help during the preparation of this part. José H. Leal and David G. Robinson identified and revised specimens. J. Michel Montoya (Costa Rica), Kay Hale and Helen D. Albertson (previously at the RSMAS Library, University of Miami), Marien Faber (Museum of Zoology in Amsterdam), and Emilio Rolán (Sociedad Española de Malacología) provided many papers that contained information on Costa Rican gastropods. Useful comments by anonymous reviewers and by Carol Skoglund are greatly appreciated. References 1. Abbott RT (1974) American Seashells. Van Nostrand Reinhold, New York, 663 p 2. Abbott RT, Dance SP (1982) Compendium of Seashells. E.P. Dutton, New York, 411 p 3. Adams CB (1852) Catalogue of shells collected at Panama, with notes on synonymy, station, and geographical distribution. Ann Lyc Nat Hist New York 5:229–548 4. Allmon WD (1988) Ecology of recent turritelline gastropods (Prosobranchia, Turritellidae): current knowledge and paleontological implications. 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