The Snapping Shrimp Genus Alpheus from the Eastern Pacific (Decapoda: Caridea: Alpheidae)
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The Snapping Shrimp Genus Alpheus from the Eastern Pacific (Decapoda: Caridea: Alpheidae) WON KIM and LAWRENCE G. ABELE SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY • NUMBER 454 SERIES PUBLICATIONS OF THE SMITHSONIAN INSTITUTION Emphasis upon publication as a means of "diffusing knowledge" was expressed by the first Secretary of the Smithsonian. In his formal plan for the Institution, Joseph Henry outlined a program that included the following statement: "It is proposed to publish a series of reports, giving an account of the new discoveries in science, and of the changes made from year to year in all branches of knowledge." This theme of basic research has been adhered to through the years by thousands of titles issued in series publications under the Smithsonian imprint, commencing with Smithsonian Contributions to Knowledge in 1848 and continuing with the following active series: Smithsonian Contributions to Anthropology Smithsonian Contributions to Astrophysics Smithsonian Contributions to Botany Smithsonian Contributions to the Earth Sciences Smithsonian Contributions to the Marine Sciences Smithsonian Contributions to Paleobiology Smithsonian Contributions to Zoology Smithsonian Folklife Studies Smithsonian Studies in Air and Space Smithsonian Studies in History and Technology In these series, the Institution publishes small papers and full-scale monographs that report the research and collections of its various museums and bureaux or of professional colleagues in the world of science and scholarship. The publications are distributed by mailing lists to libraries, universities, and similar institutions throughout the world. Papers or monographs submitted for series publication are received by the Smithsonian Institution Press, subject to its own review for format and style, only through departments of the various Smithsonian museums or bureaux, where the manuscripts are given substantive review. Press requirements for manuscript and art preparation are outlined on the inside back cover. Robert McC. Adams Secretary Smithsonian Institution SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY • NUMBER 454 The Snapping Shrimp Genus Alpheus from the Eastern Pacific (Decapoda: Caridea: Alpheidae) Won Kim and Lawrence G. Abele SMITHSONIAN INSTITUTION PRESS Washington, D.C. 1988 ABSTRACT Kim, Won, and Lawrence G. Abele. The Snapping Shrimp Genus Alpheus from the Eastern Pacific (Decapoda: Caridea: Alpheidae). Smithsonian Contributions to Zoology, number 454, 119 pages, 45 figures, 2 tables, 1988.—A revision of the genus Alpheus revealed 53 nominal species reported from the eastern Pacific. Of these, 23 species are considered valid and an additional 21 new species are described, resulting in 44 eastern Pacific species. A key to all 44 species is provided. Forty-four species are described and 41 are illustrated. The species occur in a variety of habitats including coral (18 species), rocky intertidal (29 species), mangroves (3 species), and sand-mud (21 species). Most species occur in several habitats while a few (e.g., Alpheus lottini) are found only in a single habitat. Seventy-seven percent of the species are endemic to the eastern Pacific while 11% also occur on both coasts of the Atlantic. Five percent are shared with the Indo-West Pacific and 2% each with the western Atlantic, eastern Atlantic and Indo-West Pacific and all other tropical regions. Within the eastern Pacific, species of Alpheus occur as far north as Horseshoe Cave, Sonora County, California, and as far south as Puerto Montt, Chile. Twenty species occur in the Galapagos Islands of which only one is apparently endemic. The greatest number of species (33) is found in the subregion between about 12°N and 2°N (Costa Rica, Panama, Colombia) with the number decreasing north and south of this subregion. OFFICIAL PUBLICATION DATA is handstamped in a limited number of initial copies and is recorded in the Institution's annual report, Smithsonian Year. SERES COVER DESIGN: The coral Montastrea cavernosa (Linnaeus). Library of Congress Cataloging-in-Publication Data Kim, Won, 1955- Thc Snapping Shrimp Genus Alpheus from the eastern Pacific (Decapoda: Caridea: Alpheidae) (Smithsonian contributions to zoology ; no. 454) Bibliography: p. Supt. of Docs.: SI1.27:454. 1. Alpheus—Pacific Ocean—Classification. 2. Crustacea—Classification. 3. Crustacea—Pacific Ocean— Classification. I. Abele, Lawrence G. II. Title. HI. Series. QL1.S54 no.454 591 s 86-607946 [QL444.M33] [595 3'843] Contents Page Introduction 1 Acknowledgments 1 Materials and Methods 2 Alpheus Fabricius, 1798 3 Seven Species Groups 7 Key to Eastern Pacific Species of Alpheus 9 1. Alpheus bellimanus Lockington, 1877 13 2. Alpheus rectus, new species 16 3. Alpheus hoonsooi, new species 18 4. Alpheus barbara Lockington, 1878 21 5. Alpheus clamator Lockington, 1877 21 6. Alpheus inca Wicksten and M6ndez, 1981 24 7. Alpheus grahami Abele, 1975 26 8. Alpheus websteri Kingsley, 1880 28 9. Alpheus malleator Dana, 1852 31 10. Alpheus lottini Guerin-Meneville, 1829 33 11. Alpheus normanni Kingsley, 1878 35 12. Alpheus panamensis Kingsley, 1878 38 13. Alpheus felgenhaueri, new species 40 14. Alpheus sulcatus Kingsley, 1878 42 15. Alpheus cristulifrons Rathbun, 1900 44 16. Alpheus cylindricus Kingsley, 1878 47 17. Alpheus paracrinitus Miers, 1881 49 18. Alpheus rostratus, new species 51 19. Alpheus floridanus Kingsley, 1878 53 20. Alpheus aequus, new species 55 21. Alpheus fasciatus Lockington, 1878 57 22. Alpheus bouvieri A. Milne Edwards, 1878 58 23. Alpheus chilensis Coutiere, 1902 60 24. Alpheus hebes, new species 62 25. Alpheus longinquus, new species 65 26. Alpheus lacertosus, new species 68 27. Alpheus californiensis Holmes, 1900 70 28. Alpheus canalis, new species 72 29. Alpheus hyeyoungae, new species 75 30. Alpheus scopulus, new species 77 31. Alpheus tenuis, new species 79 32. Alpheus villus, new species 82 33. Alpheus umbo, new species 84 34. Alpheus mazatlanicus Wicksten, 1983 86 35. Alpheus lams, new species 88 36. Alpheus hamus, new species 91 37. Alpheus firmus, new species 93 38. Alpheus distinctus, new species 95 39. Alpheus martini, new species 98 40. Alpheus pacificus Dana, 1852 100 41. Alpheus galapagensis Siwertsen, 1933, new rank 102 in iv SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY 42. Alpheus spinicaudus Lockington, 1878 103 43. Alpheus antepaenultimus, new species 103 44. Alpheus exilis, new species 105 Zoogeographical Considerations 107 Appendix: Station List 109 Literature Cited 114 The Snapping Shrimp Genus Alpheus from the Eastern Pacific (Decapoda: Caridea: Alpheidae) Won Kim and Lawrence G. Abele Introduction poorly until now. About 150 years ago, H. Milne Edwards (1837) described one species in the genus Alpheus from the The snapping shrimp genus Alpheus is a diverse taxon that coast of Chile (Alpheus spinifrons) and this may be the first contains more than 250 species worldwide. Species are found note of any species of Alpheus from the eastern Pacific. After in four major habitats: (1) on living and dead corals; this is the H. Milne Edwards, several authors reported species of Alpheus environment containing the largest number of species; (2) in from scattered localities in the eastern Pacific. The main works tide pools and in small pockets of water in the sand or were: Lockington (1878) from both coasts of North America, sand-mud intertidal; (3) in burrows in sand and muddy Kingsley (1878a) from the west coast of Central America substrate or occasionally under rocks; and (4) as symbionts including Panama, Holmes (1900) from California, Rathbun with some invertebrates, especially anemones, echinoids and (1910) from the Peruvian coast, Hult (1939) and Sivertsen annelids, and fishes (Banner and Banner, 1966; Abele and (1933) from the Galapagos Islands, Chace (1937) from Baja Felgenhauer, 1982). California, Schmitt (1939) from the Clipperton Islands and the Prior systematic work on this genus has resulted in Galapagos Islands, Holthuis (1952) from Chile, Chace (1962) monographic publications on the fauna of the eastern Atlantic from the Clipperton Islands, Abele (1975) from the Malpelo (Crosnier and Forest, 1966), western Atlantic (Chace, 1972), Islands (Colombia), Hendrickx et al. (1983) from the west and Indian Ocean and central Pacific faunas (Banner, 1953; coast of Mexico, and Wicksten (1983) from the Gulf of Banner and Banner, 1966, 1982). In spite of these and other California. Recently, Wicksten and MSndez (1981) described systematic studies on Alpheus, the status of many species is A. inca from Chile, and Wicksten (1983) described A. uncertain and many names have been synonymized by different mazatlanicus from the Laguna Caimanero, Sinaloa, Mexico. authors. This situation may be the result of difficulties in At present, 53 nominal species have been reported in Alpheus identifying members of Alpheus owing to their small size, Fabricius or Crangon Weber, from the eastern Pacific. Of ambiguous morphological characters, and fragility of spec- these, 23 species may be valid, even though several are imens. uncertain (Table 1). Coutiere (1899; indexed by Chace and Forest, 1970) divided ACKNOWLEDGMENTS.—We thank Raymond B. Manning for Alpheus into 5 nominal "groups" and further divided one of allowing the first author to work at the Smithsonian these into three "subgroups" in 1905. Even though these groups Institution's National Museum of Natural History and for the have no true systematic status, several authors have followed loan of the specimens of Alpheus. We wish to thank Fenner this classification as a means of dealing