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Collected by Mr. Macgillivray During the Voyage of HMS Rattlesnake

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Collected by Mr. Macgillivray During the Voyage of HMS Rattlesnake BULLETIN OF MARINE SCIENCE VOLUME 24 1974 NUMBER 4 BIOLOGICAL RESULTS OF THE UNIVERSITY OF MIAMI DEEP-SEA EXPEDITIONS. 106. THE LOBSTERS OF THE SUPERFAMILY NEPHROPIDEA OF THE ATLANTIC OCEAN (CRUSTACEA: DECAPODA)1 L. B. HOLTHUIS Rijksmuseum van Natuurlijke Historie, Leiden, The Netherlands ABSTRACT An enumeration is given of the 18 known Atlantic species of Nephro- pidean lobsters. Special attention is devoted to the family group taxonomy, and to the morphology of the carapace. Keys are provided to all species. One subfamily, two genera and three species are described as new. The geographic distribution of each species is discussed, and all available infor­ mation on the biology of the tropical forms is brought together. The spe­ cies are provided with descriptions and figures or with references to such descriptions and figures published elsewhere. INTRODUCTION The present paper is based in the first place on the Nephropid material from deeper water collected between 1962 and 1971 by the research ships of the University of Miami, R. V. GERDA and R. V. PILLSBURY. These collections were made in West Indian waters (GERDA, 1962-1971; PILLS­ BURY, 1966-1971) and off the west coast of Africa (PILLSBURY, 1964, 1965). Also all the Nephropid material from the tropical Atlantic present in the collections of the Institute of Marine and Atmospheric Science, Uni­ versity of Miami, Florida, U.S.A. (M), the National Museum of Natural History, Smithsonian Institution, Washington, D.C., U.S.A. (W), the Mu­ seum of Comparative Zoology, Harvard University, Cambridge, Mass., U.S.A. (MCZ), the British Museum (Natural History), London, England (BM), the Rijksmuseum van Natuurlijke Historie, Leiden, Netherlands (L), and the Museum National d'Histoire naturelle, Paris, France (MP) has been consulted. 1 Contribution from the Rosenstiel School of Marine and Atmospheric Science, University of Miami. 724 Bulletin of Marine Science [24(4) This study was started several years ago, but due to several circumstances could not be finished until now. Without the very cordial help that I re­ ceived from many persons, I would not have been able to finish it at all. In the first place I want to thank Dr. Gilbert L. Voss and Dr. Frederick M. Bayer of the University of Miami, who placed all the GERDA and PILLS- BURY material at my disposal, made it possible for me to partake in eight of the PILLSBURY and one of the GERDA cruises and assisted me in every possible way in this project. Dr. Fenner A. Chace and Dr. Raymond B. Manning of the Smithsonian Institution allowed me to examine all the material of the National Museum of Natural History and made arrange­ ments enabling me to work for several months in the Museum on these collections; I am especially grateful to Dr. Manning for generously allowing me to publish on the Eunephrops material on which he himself originally intended to devote a study; he provided me also with all the illustrative material on that genus that he had brought together already. Dr. Herbert W. Levi of the Museum of Comparative Zoology, Drs. Isabella Gordon, R. W. Ingle and A. L. Rice of the British Museum, and Drs. M. de Saint Laurent and J. Forest of the Paris Museum gave me every possible help with the study of the material under their care. The drawings illustrating the present paper are by Mrs. Constance Stolen McSweeny, at that time staff artist of the Institute of Marine and Atmospheric Science, Miami, Mrs. Carolyn Gast, staff artist of the Smith­ sonian Institution, Mr. W. C. G. Gertenaar, staff artist of the Rijksmuseum van Natuurlijke Historie, Leiden, and Mrs. Lilly Manning. The photo­ graphs are made by the staff photographers of the Smithsonian Institution and the Leiden Museum (Mr. Chr. Hoorn), with the exception of Figures 7 and 8, which were placed at my disposal by the U.S. Naval Oceanographic Office, through the kindness of Mr. Timothy W. lanaitis. The abbreviation cl. is used in the text to denote the carapace length measured from the tip of the rostrum to the median point of the posterior margin of the carapace. In the list of examined material the institutions holding the material in question are indicated with the letters that in the first paragraph of this in­ troduction are given in parentheses after the full name of each institution. SYSTEMATIC PART The taxonomy of the true lobsters has been differently interpreted by different authors. In the most generally accepted classification of the De- capoda, Waterman & Chace (1960: 25) considered these animals to form the superfamily Nephropsidea of the section Macrura. Glaessner (1969: 400, 455) raised this group to the rank of an infraorder (= section in the classification of Waterman & Chace), which he named Astacidea; he was followed in this by Hobbs (1974: 2). When compared with the super- 1974] Holthuis: Taxonomy and Zoogeography of the Nephropidae 725 families in Caridea, the Astacidea indeed deserve a higher rank than super- family, and should be treated like an infraorder. The synonymy of this infraorder is as follows: Astacidea Latreille, 1803 Astacini p.p. Latreille, 1802-1803, Hist.nat.Crust.Ins., 3: 32. Macroura Divisio VI Leach, 1815, Trans.Linn.Soc.London, 11: 343. Homardiens Desmarest, 1823, Dict.Sci.nat., 28: 306. Fissilaminata Haworth, 1825, Philos.Mag.Journ., 65: 184. Astacini Latreille, 1831, Cours Entomol., 1: 378. Astacina Burmeister, 1837, Handb.Naturgesch., 3: 563. Astacomorpha Huxley, 1880, Crayfish, (ed. 1): 338, 339, 341, 342, 363. Homaridae Boas, 1880, K.Danske Vidensk.Selsk.Skr., (6)1(2): 156, 174. Homariderne Boas, 1880, K.Danske Vidensk.Selsk.Skr., (6)1(2): 68. Homaridea Ortmann, 1891, Zool.Jb.Syst., 6: 1. Nephropsidea Ortmann, 1898, Bronn's Klass. Ordn.Thier-Reichs, 5(2): 1138. Astacura Borradaile, 1907, Ann.Mag.nat.Hist., (7)19: 464, 467. Astacidea Fowler, 1912, Ann.Rep.New Jersey State Mus., 1911: 332. Herpochelida Beurlen & Glaessner, 1930, Zool.Jb.Syst., 60: 49, 53. Nephlopsidea Yokoya, 1933, Journ.Coll.Agric. Tokyo, 12: 48. Astacidea Glaessner, 1969, Moore's Treatise Invert.Paleontol., R (4): 455. The infraorder Astacidea was divided by Hobbs (1974) into three super- families: (1) the Astacoidea, containing the freshwater crayfishes of the northern Hemisphere, (2) the Parastacoidea, formed by the freshwater crayfishes of the Southern Hemisphere, and (3) the Nephropoidea con­ taining the marine lobsters. In the present paper the Nephropoidea of the Atlantic Ocean are dealt with. Most attention is given to the species oc­ curring in the tropical and southern zones of the Atlantic Ocean. The Northern Hemisphere, (2) the Parastacoidea, formed by the freshwater for completeness's sake; their synonymy and distribution are listed, some remarks are given, but no complete account is provided. The other species are more exhaustively dealt with. Nephropoidea Dana, 1852 The Nephropoidea consist of two families, the Nephropidae (usually indi­ cated as Nephropsidae or Homaridae) and the Thaumastochelidae. It is possible that also the Axiidae should be placed in this superfamily rather than in the Thalassinidea: they are connected with the Nephropidae by so many similarities and so few differences, that it is often difficult to decide whether a genus belongs to one or the other of these two families; so Eu- trichocheles and Enoplometopus, which for a long time have been consid­ ered Nephropidae actually have to be placed with the Axiidae. The posi­ tion of the Axiidae is not further considered here and only the Atlantic members of the families Thaumastochelidae and Nephropidae are dealt with. 726 Bulletin of Marine Science [24(A) Recentiy, Glaessner (1969) divided the heterogeneous assemblage of Nephropid genera into three subfamilies: Nephropinae, Homarinae, and Neophoberinae. The characters shown by Homarus are so close to those of Nephropine genera like Nephrops, Metanephrops, and Eunephrops, that I have not been able to distinguish a subfamily Homarinae, and have to refer Homarus to the Nephropinae. On the other hand the genera Nephrop- sis, Nephropides, and two new genera (Thymops and Thymopsis) described below, differ sufficiently from the other Nephropid genera to constitute a separate subfamily Thymopinae. Glaessner's subfamily Neophoberinae is a very distinct and well characterized taxon. KEY TO THE FAMILIES THAUMASTOCHELIDAE AND NEPHROPIDAE 1 Eyes entirely absent, eventual remnants immovably fused with the ophthalmic somite. Telson unarmed. Chelipeds very unequal, the larger with fingers more than four times as long as the palm; cutting edges of the fingers of the larger cheliped with two rows of diverging slender spines. Fifth pereiopod (at least in the fe­ male) with a chela. Abdominal pleura short, quadrangular, lat­ eral margin broad, truncate, not ending in a point. Scaphocerite with several very large teeth on the inner margin. Thaumastochelidae 1' Eyes well developed or reduced, always present as movable ap­ pendages. Telson with lateral and/or postlateral spines. Cheli­ peds equal or unequal, but fingers always considerably less than twice as long as palm; teeth on the cutting edge placed in the same plane. Fifth pereiopod without a true chela. Abdominal pleura large, triangular or ovate, usually ending in a point. Scapho­ cerite, if present, with the inner margin evenly curved, unarmed. Nephropidae 2 Rostrum laterally compressed for the larger part of its length, with dorsal and ventral, but no lateral teeth. Carapace with branchiostegal spine. Body entirely covered by numerous closely placed and sharply pointed spinules. Lateral margin of the tel­ son with 6 to 12 spines. Neophoberinae 2' Rostrum dorsoventrally depressed with lateral (and sometimes ventral), but without dorsal teeth. Carapace without a branchi­ ostegal spine. Body never uniformly covered with spinules, al­ though granules may be present all over, or spinules may be placed on the carapace. The lateral margin of the telson with at most three lateral spines, which if present, are usually small and irregular.
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