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Reprinted from: CRUSTACEANA, Vol. 28, Part 1, 1975 CO ^fjb-' LEIDEN E. J. BRILL

Crustaceana 28 (1), 1975, E. J. Brill, Leiden THE IDENTITY OF SESARMA HANSENI RATHBUN, 1897, A SUPPOSEDLY WEST INDIAN SPECIES, WITH 5. DEHAANI H. MILNE EDWARDS, 1853, FROM THE WEST PACIFIC (DECAPODA, GRAPS1DAE) BY LAWRENCE G. ABELE Florida State University, Department of Biological Science, Tallahassee, Florida, 32306, U.S.A. INTRODUCTION The status of several species of Sesarma from the western Atlantic has recently been reviewed. Chace & Hobbs (1969) presented diagnoses and illustrations of the West Indian species and showed that Sesarma americanum De Saussure, 1853 is a senior synonym of S. tampicense Rathbun, 1914, rather than a junior synonym of S. an gust/pes Dana, 1852 as suggested by Rathbun (1918). Abele (1972) reviewed the status of five nominal species of Sesarminae from the western Atlantic. He concluded that S. angustipes Dana, 1852 is a senior synonym of S. miersii iheringi Rathbun, 1918, rather than a senior synonym of S. roberti H. Milne Edwards, 1853 as suggested by Hartnoll (1965). The present report examines the status of Sesarma hanseni Rathbun, 1897. Rathbun (1897) described the new species S. hanseni based on a single male specimen in the Copenhagen Museum. The only data accompanying the specimen indicate it came from "Vestindien". Additional material of the species has never been taken in the West Indies, although several authors dealing with that fauna have listed the name. Through the courtesy of Dr. Torben Wolff, Copenhagen Museum, I was able to examine the unique holotype of S. hanseni. Study of the specimen and comparisons with other species revealed that it is conspecific with a very common West Pacific species, S. dehaani H. Milne Edwards, 1853. The holotype of S. hanseni is, in all probability, mislabeled and should not be considered a part of the West Indian fauna. The holotype of S. hanseni is illustrated and some descriptive notes are presented. Sesarma (Holometopus) dehaani H. Milne Edwards, 1853 (figs. 1, 2) Grapsus (Pachysoma) quadratus - De Haan, 1835: 62, pi. 8 fig. 3 [not Cancer quadratus FabriciusJ. Sesarma dehaani H. Milne Edwards, 1853: 184; Stimpson, 1858: 106; Heller, 1865: 62; Kingsley, 1880: 214; De Man, 1887: 642; Burger, 1893, 615; Ortmann, 1894: 718; Stimpson, 1907: 134; Kemp, 1918: 235; Parisi, 1918: 111. Sesarma (Holometopus) dehaani - Tesch, 1917: 143; Balss, 1922: 154; Urita, 1926: 19; Shen, 1932: 195, text-figs. 121-123, pi. 9 fig. 1; Sakai, 1934: 324; Sakai, 1936: 234, pi. 65 fig. 1; Sakai, 1939: 681-682, pi. 77 fig. 1; Sakai, 1965: 202, pi. 97 fig. 2. Sesarma neglecta De Man, 1887: 643, 661; Tesch, 1917: 178. Sesarma hanseni Rathbun, 1897: 92.

SESARMA HANSEN! RATHBUN 49 Sesarma (Holometopns) hanseni. Rathbun, 1918: 315, text-fig. 152, pi. 87 fig. 1; Chace & Hobbs, 1969: 179. Material examined. Holotype of S. hanseni-, male, cb (carapace breadth) 16.5 mm; "Vestindien"; Copenhagen Museum. 4 males, cb 16.6 to 21 mm; Pacific Ocean, Formosa, Miao, Li Hsien; USNM (National Museum of Natural History) 123498. Description of the holotype of Sesarma hanseni. The carapace breadth is about 1.27 times its length. The frontal region is deflexed; it does not widen distally and is about 0.55 of the carapace breadth. A shallow median sinus is present. The interorbital area is divided into four distinct lobes. The outer orbital tooth is acute. There is a minute indentation posterior to the outer orbital tooth. Fig. 1. Holotype of Sesarma hanseni Rathbun, 1897. A, dorsal view of carapace; B, fifth pereiopod; C, dorsal view of right chela. Scale = 5 mm for A, 10 mm for B, C. The posterior portion of the carapace narrows distinctly slightly anterior to the midline. There are four oblique rows of granules on each side of the posterolateral portions of the carapace. The gastric and cardiac regions are distinct. The carapace is naked and sparsely punctate. The eyes are well developed and the cornea is pigmented. The third maxillipeds gape widely and have an oblique, hairy ridge on the merus. The chelipeds are subequal and robust. The merus has the margins granulate, almost serrate; the lateral surface is covered with short rows of granules; there is no distal inferior tooth. The carpus has two granular ridges present; one on the medial margin and the other on the dorsal surface. Short rows of granules are present lateral to the ridge on the dorsal surface. An elongate lobe is present at the lateral angle. The dorsal surface of the palm has a raised, granulate ridge 4

50 LAWRENCE G. ABELE which is bifurcate in the distal half and extends as a poorly defined ridge onto the medial surface of the palm. A few large granules are present at the distal margin of the palm. The lateral surface of the palm is covered with low, depressed granules. The dorsal surface of the movable finger is covered with marty small granules. The fingers are slightly spooned at the dark colored tips. Each finger is armed with about six unequal teeth. There is only a single complete walking leg with the specimen. It is the last (fifth pereiopod); the merus is longer than the other segments, its length is about 2.3 times its width. The carpus is a little shorter than the propodus and is unarmed. The propodus is armed with about eight black spines on the ventral surface. The dactylus is shorter than the propodus and is armed with about two small, black spines dorsally and about four ventrally. of S. hanseni. Scale = 1 mm. The abdomen is damaged but it has been figured by Rathbun (1918: 316, text-fig. 152a). The gonopod is simple, the apex is extended and curved laterally. There is a small expansion proximal to the distal portion of the dark colored endpiece. Distribution. China: Liaoting Peninsula (east), Kiangsu (Woosung, Shanghai), Fukien (Foochow, Amoy), Kwangtung (Whampoa). Hong Kong. Formosa (Taiwan). Japan: Bosco Province to Kyusyu and Okinawa. Korea. (Shen, 1932; Sakai, 1965). Habitat. The species occurs along the banks of muddy streams growing with grasses some distance from brackish water. It also occurs in rice fields where its burrows cause some damage (Shen, 1932; Sakai, 1939). Color. Color notes are given by Stimpson (1907), and Shen (1932) and a color plate is given by Sakai (1965, pi. 97 fig. 2). Remarks. - The holotype of S. hanseni is slightly smaller than the available specimens of S. dehaani but there is little doubt that the specimens are conspecific. Dr. Lipke B. Holthuis, of the Leiden Museum, kindly compared the illustrations of the gonopods of S. hanseni with those of the lectotype of S. dehaani (a male specimen, cl 37 mm, cb 40 mm, from Japan, leg. P. F. von Siebold, 1823-1830;

SESARMA HANSENI RATHBUN 51 RMNH no. Crust. D 157) in the Rijksmuseum van Natuurlijke Historie, Leiden, and confirmed the synonymy. Shen (1932) discussed variation in S. dehaani, especially regarding the number of teeth on the anterolateral margins. The majority of specimens have a slight emargination or tooth posterior to the outer orbital angle, while other specimens have none, two or three (Shen, 1932: 198, text-fig. 123). The tooth of S. dehaani is almost as well developed as that of S. reticulatum (Say, 1817) (see Williams, 1965: 221, fig. 205), the type species of the genus Sesarma. Two of the subgenera of Sesarma, Sesarma (type S. reticulatum) and Holometopus (type S. haematocheir), are differentiated by the presence (Sesarma) or absence (Holometopus) of a tooth posterior to the outer orbital tooth. The distinction, however, appears to be more a matter of convenience. Sesarma dehaani and S. benedicti Rathbun, 1897, for example, both have a slight emargination or tooth posterior to the outer orbital tooth yet both are included in the subgenus Holometopus. Among the American species of Sesarma, S. cinereum (Bosc, 1801-1802), S. miersii Rathbun, 1897, S. angustipes Dana, 1852, S. biolleyi Rathbun, 1906, S. magdalenense Rathbun, 1918 and S. occidentale Smith, 1870 form a very closely related group of species in the subgenus Holometopus while S. benedicti is just as closely related to S. reticulatum as to any of the above species. Serene & Soh (1970) recently split the genus Sesarma into about 19 genera, several of which are based on the presence or absence of teeth posterior to the outer orbital angle. It would seem, as the above authors noted, that additional characters are needed before this designation can be fully justified. ACKNOWLEDGEMENTS I thank Dr. Torben Wolff, Copenhagen Museum, for allowing me to examine the holotype of S. hanseni and Dr. R. Serene, National Museum of Singapore, for his comments on the manuscript. Dr. Serene kindly supplied photographs of S. dehaani from the Indo-Pacific for comparison and verified that the gonopods of S. hanseni and S. dehaani are identical. I thank also Dr. Lipke B. Holthuis, Leiden Museum, for his comments and for confirming the synonymy of S. dehaani and S. hanseni through examination of the type of the former species in the Leiden Museum. Messrs. C. Alan Child and Henry Roberts were most helpful during my visit to the National Museum of Natural History, Washington, D. C. A portion of this work was supported by the Department of Invertebrate Zoology, Smithsonian Institution. Dr. Robert H. Gore, Smithsonian Institution, kindly supplied several references which were unavailable to me. RESUME Le reexamen de l'holotype de Sesarma hanseni Rathbun, 1897, decrit des Antilles, a revele qu'il appartenait a l'espece Sesarma dehaani H. Milne Edwards, 1853, du Pacifique occidental. Sesarma hanseni est par consequent un synonyme subjectif de Sesarma dehaani. L'holotype de S. hanseni est, selon toute probability mal etiquete et ne devrait pas etre considere comme appartenant a la faune des Antilles.

52 ABELE, SESARMA HANSENI RATHBUN LITERATURE CITED ABELE, L. G., 1972. The status of Sesarma angustipes Dana, 1852, S. trapezium Dana, 1852 and S. miersii Rathbun, 1897 (Crustacea: Decapoda: Grapsidae) in the western Atlantic. Carib. Journ. Sci., 12 (3-4): 165-170, figs. 1-2. BALSS, H., 1922. Ostasiatische Decapoden, 4. Die Brachyrhynchen (Cancridea). Archiv Naturgesch., 88 (A) (11): 94-166, pis. 1, 2. BURGER, O., 1893. Beitrage zur Kenntniss der Gattung Sesarma. Zool. Jahrb. (Syst.), 7: 613-632. CHACE, F. A., Jr., & H. H. HOBBS, Jr., 1969. The freshwater and terrestrial decapod crustaceans of the West Indies with special reference to Dominica. Bull. U. S. nation. Mus., 292.- i-v, 1-258, figs. 1-76, pis. 1-5. HAAN, W. DE, 1833-1850. Crustacea. In: P. F. VON SIEBOLD, Fauna Japonica: i-xvi, i-xxxi, vii-xvii, 1-243, pis. A-Q, 1-55. HARTNOLL, R. G., 1965. Notes on the marine grapsid crabs of Jamaica. Proc. Linn. Soc., London (Zool.), 176: 113-147, figs. 1-16. HELLER, C., 1865. Die Crustaceen. Reise der oesterreichischen Fregatte "Novara" um die Erde, Zool., 2 (3): 1-280, pis. 1-25. KEMP, S., 1918. Crustacea Decapoda and Stomatopoda. Zoological results of a tour in the Far East. Mem. Asiatic Soc. Bengal, Calcutta, 6: 217-297. KINGSLEY, J. S., 1880. Synopsis of the Grapsidae. Carcinological notes, 4. Proc. Acad. nat. Sci., Philadelphia, 1880: 187-224. MAN, J. G. DE, 1887. Uebersicht der Indo-Pacifischen Arten der Gattung Sesarma Say. Zool. Jahrb. (Syst.), 2: 639-722. MILNE EDWARDS, H., 1853. Memoire sur la famille des Ocypodiens. Ann. Sci nat., (Zool.) (3) 20: 163-228, pis. 6-11. ORTMANN, A. 1894. Die Decapoden Krebse des Strassburger Museum, 8. Cancroidea, Cancrinea, Catametopa. Zool. Jahrb. (Syst.), 7: 683-772. PARISI, B., 1918. Decapodi giapponesi del Museo di Milano, 6. Catometopa e Paguridea. Atti Soc. Ital. Sci. nat., 57: 90-115, pi. 8. RATHBUN, M. J., 1897. Synopsis of the American Sesarmae, with description of a new species. Proc. biol. Soc. Washington, 11: 89-92. 1918. The grapsoid crabs of America. Bull. U. S. nation. Mus., 97: i-xxii, 1-461, figs. J-172, pis. 1-161. SAKAI, T., 1934. Brachyura from the coast of Kyusyu, Japan. Sci. Rep. Tokyo Bunn'ka Daigaku, (IB) 1 (25): 281-330. 1936. Crabs of Japan. 66 plates in life colour with descriptions: 1-239, 1-12, 1-27. (Tokyo). 1939. Brachygnatha, Brachyrhyncha. Studies on the crabs of Japan, 4: 365-741, pis. 42-111. (Yokendo, Tokyo). 1965. The crabs of Sagami Bay: i-xvi, 1-206, figs. 1-27, pis. 1-100 [English}. (Maruzen Co., Tokyo). SERENE, R., & C. L. SOH, 1970. New Indo-Pacific genera allied to Sesarma Say, 1817 (Brachyura, Dccapoda, Crustacea). Treubia, 27 (4): 387-416, pis. 1-8. SHEN, C. J., 1932. The Brachyura Crustacea of North China. Zoologica Sinica, (A) 9 (1): 1-320, pis. 1-10. STIMPSON, W., 1858. Prodromus descriptions animalium evertebratorum quae in Expeditione ad Oceanum Pacificum Septentrionalem, a Republica Federata missa, Cadwaladaro Ringgold et Johanne Rodgers ducibus, observavit et descwpsit. Proc. Acad. nat. Sci., Philadelphia, 10 (4): 31-40. 1907. Report on the Crustacea collected by the North Pacific Exploring Expedition 185 3-56. Smithsonian misc. Coll., 49: 1-240. TESCH, J. J., 1917. Synopsis of the genera Sesarma, Metasesarma and Clistocoeloma with a key to the determination of the Indo-Pacific species. Zool. Mededeel. Leiden, 3: 127-260, pis. 15-17. URITA, T. 1926. A check list of Brachyura found in Kagoshima Pref., Japan: 1-40. (The Tsingtao Times, Tsingtao). WILLIAMS, A. B. 1965. Marine decapod crustaceans of the Carolinas. U. S. Dept. Interior, LJ. S. Fish Wildl. Serv., Fish. Bull., 65: 1-298, figs. 1-252. Received for publication 22 January 1973.