Zootaxa 3637 (4): 412–431 www.mapress.com / zootaxa / Copyright © 2013 Magnolia Press ISSN 1175-5326 (print edition) Article ZOOTAXA ISSN 1175-5334 (online edition) http://dx.doi.org/10.11646/zootaxa.3637.4.2 http://zoobank.org/urn:lsid:zoobank.org:pub:54055A0B-A1CD-4C7D-AAA9-F46AFD0B1D5F Re-examination of the eastern Pacific and Atlantic material of Alpheus malleator Dana, 1852, with the description of Alpheus wonkimi sp. nov. (Crustacea, Decapoda, Alpheidae) ARTHUR ANKER1,2,4 & PAULO P.G. PACHELLE3 1 Laboratório de Zoobentos, Instituto de Ciências do Mar (Labomar), Universidade Federal do Ceará, Avenida da Abolição, 3207, Meireles, 60165-081, Fortaleza, CE, Brazil 2 Current address: Tropical Marine Science Institute, National University of Singapore, 18 Kent Ridge Road, Singapore 119227, Republic of Singapore 3 Laboratório de Invertebrados Marinhos do Ceará, Departamento de Biologia, Universidade Federal do Ceará, Fortaleza, Ceará, Brazil 4 Correspoding author. E-mail: arthuranker7@yahoo.com Abstract The bumpy-clawed snapping shrimp, Alpheus malleator Dana, 1852 (Alpheidae), is revised based on the recently collected and older museum material from the eastern Pacific (Panama, Ecuador), Caribbean (Panama, Puerto Rico, Trinidad & Tobago), Brazil (São Paulo), and West Africa (Cape Verde, Senegal, Guinea, Equatorial Guinea, Congo). The eastern Pacific material is assigned to A. wonkimi sp. nov., based on one morphological difference in the colour and thickness of the uropodal spiniform seta, as well as previously published molecular data. The Caribbean, Brazilian and West African material is considered to represent a single, widespread, morphologically variable, amphi-Atlantic taxon, A. malleator. Alpheus pugilator A. Milne-Edwards, 1878 is retained as a junior synonym of A. malleator, whereas A. tuberculosus Osorio, 1892, A. malleator var. edentatus Zimmer, 1913 and A. belli Coutière, 1898, the latter two based on juvenile specimens, are tentatively placed in the synonymy of A. malleator. Illustrations, including colour photographs, are provided for A. wonkimi sp. nov. and A. malleator and their morphological variability is discussed and illustrated. Key words: Alpheidae, Alpheus, snapping shrimp, cryptic species, transisthmian speciation, Pacific Ocean, Atlantic Ocean Introduction The snapping shrimp genus Alpheus Fabricius, 1798 is well known for a remarkable diversity in the shape and sculpture of the major and minor chelipeds (e.g., Coutière 1899; Banner & Banner 1982). The bumpy-clawed snapping shrimp, Alpheus malleator Dana, 1852, is one of the most distinctive members of the genus. This species can be immediately recognised by the heavily sculptured (covered with large, projecting bumps and small tubercles) and setose major and minor claws. Other diagnostic features of A. malleator are the dorsally flattened and posteriorly broadened rostrum, abruptly delimited from the adjacent rostro-orbital furrows; the frontal margin of the carapace with acute orbital teeth and two smaller, acute or blunt adrostral teeth situated between the rostrum and orbital teeth; the antennal basicerite with a very stout distolateral tooth; the antennal scaphocerite with a proximal, large, laterally projecting tooth; the third maxilliped with a very short, broad penultimate article; and the uropodal exopod bearing a stout, dark-coloured, spiniform seta. The original description of A. malleator by Dana (1852a, 1852b) was a very short diagnosis in Latin, but the accompanying figures published three years later in an atlas (Dana 1855: pl. 35, fig. 9) illustrate most of the important diagnostic features of the species. The type locality of A. malleator was given with some doubts as “in portu Rio de Janeiro?”, i.e. the harbour of Rio de Janeiro, southern Brazil. Although many of Dana’s crustacean 412 Accepted by J. Goy: 25 Feb. 2013; published: 12 Apr. 2013 type specimens were lost during the Great Chicago Fire in 1871, the type of A. malleator remains preserved in the dry specimen collection of the Natural History Museum in London (NHM) (Evans 1967). Milne-Edwards (1878) described Alpheus pugilator A. Milne-Edwards, 1878 from Cape Verde, without illustrations. No comparison was made between A. pugilator and A. malleator, although based on the short descriptive text of A. pugilator these two nominal species appeared to share many important characters. Most of the extant A. Milne-Edwards’ material remains deposited in the collections of the Muséum national d’Histoire naturelle in Paris (MNHN). Osorio (1892) described Alpheus tuberculosus Osorio, 1892 from “Iogo-Iogo”, now Baia de Iogoiogo, São Tomé. Like Milne-Edwards, Osorio did not provide illustrations of his material and did not compare A. tuberculosus with A. malleator or A. pugilator, but instead with the non-related A. dentipes Guérin, 1832. Based on Osorio’s description, the only significant difference between A. tuberculosus and both A. malleator and A. pugilator is the presence of a small distoventral tooth on the merus of the third and fourth pereiopods; this tooth was not mentioned in the descriptions of A. malleator and A. pugilator. The type specimens of A. tuberculosus were presumably deposited in the Museu Bocage in Lisbon (see below). Coutière (1898) established Alpheus belli Coutière, 1898 with a single, very small specimen (length of the major chela 4 mm) of “undeterminable sex”, from Fernando de Noronha, Brazil. This author provided two somewhat diagrammatic figures of this species, showing the anterior region of the body in dorsal view and the major cheliped in the taxonomically non-informative, ventromesial view. Importantly, Coutière (1898) listed A. pugilator as a synonym of A. malleator, but did not mention Osorio’s A. tuberculosus, of which he was apparently unaware. Osorio’s study was also not mentioned in Coutière’s famous monograph of the Alpheidae (Coutière 1899). Coutière (1898) compared A. belli with A. malleator (= A. pugilator), which he considered to be very close, and listed several important differences between these two species, e.g., in the configuration of the frontal margin of the carapace, armature of the major chela and shape of the dactylus of the third to fifth pereiopods. However, at the end of the description of A. belli, Coutière stated that despite these differences, A. belli could merely be a juvenile of A. malleator. Although Coutière was based in Paris (MNHN), judging from the title of Coutière’s (1898) study (“Note sur quelques Alphéidés nouveaux de la collection du British Museum”), the type specimen of A. belli was presumably deposited in London (NHM). However, one of us was unable to trace it neither in the NHM nor in the MNHN collections in August 2012 (A. Anker, pers. obs.). Coutière (1899) provided additional drawings of A. malleator, including a very detailed illustration of the major chela, and confirmed the synonymy of A. pugilator. In addition, he assigned A. malleator to the heterogeneous “Macrochirus group”, which, however, is now considered to be non-monophyletic (see below). Nobili (1901) reported A. malleator from Bahía de Santa Elena, Ecuador, listing some differences with Dana’s (1855) and Coutière’s (1899) figures. Rathbun (1902, 1910) reported A. malleator (as Crangon malleator) from Galapagos and Bahía de Santa Elena, without illustrations nor taxonomic remarks. Nobili’s and Rathbun’s records of A. malleator are the first for the eastern Pacific (but see below). Zimmer (1913) established a new variety, Alpheus malleator var. edentatus Zimmer, 1913, based on a single, small (total length 10 mm) specimen from Barbados. Zimmer separated his variety from the typical A. malleator, using a much larger male (total length 35 mm) from Cuba (Gundlach’s collection, Museum für Naturkunde = Zoological Museum Berlin, ZMB), by the posteriorly narrower rostrum, stronger teeth on the frontal margin of the carapace, and differences in the proportions of some antennal parts. Like Coutière (1898), Zimmer (1913) stated that these differences are perhaps due to the young age of the Barbados specimen. Several drawings accompanied the description of A. malleator var. edentatus, but the deposition location of the type specimen was not precised. At the authors’ request, Charles Oliver Coleman (ZMB) confirmed the presence of Gundlach’s specimen of A. malleator in the Berlin collections (ZMB 4114), but not of Zimmer’s type of A. malleator var. edentatus. The latter was also not located in the collections of the Senckenberg-Museum in Frankfurt, during the first author’s short visit in August 2012 (A. Anker, pers. obs.). Balss (1914) provided a description and several diagrammatic drawings of a species he identified as A. tuberculosus, from Annobon, Equatorial Guinea. Balss’ figures, albeit quite schematic and therefore imprecise, show very clearly a species closely related to or identical with A. malleator. Subsequently, Balss (1916) re-assigned these specimens to A. malleator edentatus, to which he also assigned several specimens from Togo. Schmitt (1924a), while reporting A. malleator from Curaçao (as Crangon malleator), placed A. malleator var. edentatus in the synonymy of A. malleator, suggesting that some of the adult features of A. malleator may be MATERIAL OF ALPHEUS MALLEATOR AND A NEW ALPHEUS SPECIES Zootaxa 3637 (4) © 2013 Magnolia Press · 413 absent in juveniles. In another study, however, Schmitt (1924b) treated A. malleator var. edentatus as a possible junior synonym of A. nigrospinatus Rankin, 1898 (now a junior synonym of A. websteri Kingsley, 1880). However, the flattened, abruptly delimited rostrum, the presence of a long distolateral tooth on the basicerite and proximolateral tooth on the scaphocerite, and the third pereiopod with a strongly biunguiculate dactylus in A. malleator var. edentatus all show clearly that Schmitt’s (1924b) synonymy was erroneous (see Anker et al. 2008b for comparison with A. websteri). Holthuis (1951) concurred with Coutière (1899) on the status of A. pugilator and also placed A. tuberculosus sensu Balss (1914) and A. malleator edentatus sensu Balss (1916) in the synonymy of A. malleator. In the same time, Holthuis (1951) treated A. tuberculosus sensu Osorio (1892) as a species distinct from A. malleator, based on the presence of a small meral tooth on the third and fourth pereiopods observed by Osorio. Crosnier & Forest (1966) examined material of A. malleator from several localities in West Africa (Cape Verde, São Tomé, Príncipe, Annobon, Congo) and tentatively placed A. tuberculosus Osorio, 1892 in the synonymy of A. malleator. These authors argued that because (1) in most characters, in particular the uniquely armoured major chela, A. tuberculosus matches A. malleator, and (2) all specimens examined by them, including 18 specimens of both sexes from the type locality of A. tuberculosus (Annobon), were A. malleator, Osorio (1892) may have committed an observation or interpretation error. Crosnier & Forest (1966) stated that they tried to locate Osorio's types of A. tuberculosus in Lisbon's Museu Bocage to verify their hypothesis, but were unsuccessful. Therefore, it has to be assumed that Osorio’s types of A. tuberculosus were probably lost. Kim & Abele (1988) provided a detailed description and very good illustrations of the eastern Pacific A. malleator based on specimens from Costa Rica and Mexico. However, these authors did not compare the eastern Pacific and western Atlantic materials, as they did for some of the other species. Williams et al. (2001) provided a preliminary phylogenetic hypothesis for the genus Alpheus based on segments from three genes (COI, EF-1a, GPI). The trees based on COI and EF-1a revealed that A. malleator—C (Caribbean) and A. malleator—P (eastern Pacific) were genetically clearly divergent, suggesting that they represented two allopatric (transisthmian) geminate species. Material of A. malleator from other parts of the Atlantic Ocean (Brazil, West Africa) was not sequenced by Williams et al. (2001). The molecular divergence in COI and GPH sequences between A. cf. malleator (eastern Pacific) and A. malleator (Caribbean) was also clearly shown in Hurt et al. (2008), who estimated their divergence time at about 4.9 mya. In the present study, we re-examine the morphological variation of Alpheus malleator across its geographic range in the Atlantic and eastern Pacific Oceans. The eastern Pacific material includes several specimens collected by the first author in Panama in 2006–2007 and an older museum specimen from Ecuador. The western Atlantic material includes two specimens from the Caribbean coast of Panama collected by Nancy Knowlton and Javier Jara in 1995, a specimen from São Paulo collected by Valter Cobo in 2006, as well as several museum specimens from Puerto Rico and southern Brazil. The eastern Atlantic A. malleator is represented by two specimens collected by N. Knowlton and her team in Cape Verde in 1998, in addition to the museum material from Senegal, Guinea, Equatorial Guinea and Congo (some of these specimens also examined by Crosnier & Forest 1966). In spite of a remarkable morphological variation observed within the eastern Pacific, Caribbean, Brazilian and West African materials of A. malleator, the eastern Pacific specimens consistently differ from the Atlantic specimens in the thickness and colour of the stout spiniform seta on the uropodal exopod. This morphological difference, albeit a slight one, corroborates the previously reported divergence in the DNA sequences between the eastern Pacific and Caribbean specimens of A. malleator (Williams et al. 2001; Hurt et al. 2008). Thus, the eastern Pacific population clearly represents a morphologically and genetically distinct species, which is herein described as new. On the other side, the Caribbean, Brazilian and West African populations appear to form a single, widespread, amphi-Atlantic species, A. malleator, at least based on morphological evidence (see below). Therefore, A. pugilator, is retained in the synonymy of A. malleator, whereas A. tuberculosus, A. malleator var. edentatus and A. belli, the latter two clearly juvenile specimens, are tentatively placed in the synonymy of A. malleator. All the material used or mentioned in this study is or remains deposited in the collections of the Muséum national d’Histoire naturelle, Paris, France (MNHN); Natural History Museum, London (NHM); Museu de Zoologia, Universidade de São Paulo, São Paulo, Brazil (MZUSP); Nationaal Natuurhistorisch Museum, Naturalis, Leiden, the Netherlands (RMNH); Oxford University Museum of Natural History, Oxford, the United Kingdom (OUMNH); and National Museum of Natural History, Smithsonian Institution, Washington DC (USNM). Carapace length (cl, in mm) was measured from the tip of the rostrum to the posterior margin of the carapace. 414 · Zootaxa 3637 (4) © 2013 Magnolia Press ANKER & PACHELLE Taxonomy Family Alpheidae Rafinesque, 1815 Genus Alpheus Fabricius, 1798 Alpheus malleator Dana, 1852 (Figs. 1–6, 11A–H) Alpheus malleator Dana 1852a: 23; Dana 1852b: 557; Dana 1855: 23, pl. 31, fig. 9a-h; Kingsley 1883: 112; Sharp 1893: 113; Coutière 1899 (see Chace & Forest 1970 for all page numbers); Moreira 1901: 9; Luederwaldt 1919: 430; Oliveira 1940: 142; Holthuis 1951: 73; Sourie 1954: 112, 253; Crosnier & Forest 1965: 605; Crosnier & Forest 1966: 240, fig. 10; Evans 1967: 408; Chace 1972: 68; Coelho & Ramos 1972: 148; Abele 1976: 273; Christoffersen 1980a: 74; Christoffersen 1980b: 39; Abele & Kim 1986: 197, 206-207, figs. i, j; Martínez-Iglesias et al. 1997: 425; Christoffersen 1998: 359; McClure 2005: 145 (partim, not fig. 16); Hurt et al. 2008: 516 et seq.; De Grave & Fransen 2011: 386. Crangon malleator — Schmitt 1924a: 64, fig. 8; Schmitt 1936: 367; Schmitt 1939: 24. Alpheus pugilator A. Milne-Edwards 1878: 229. (?) Alpheus tuberculosus Osorio 1892: 201; Osorio 1898: 194; Rathbun 1900: 313; Balss 1914: 98, figs. 1-5; Balss 1916: 21; Vilela 1949: 56; Holthuis 1951: 73; Crosnier & Forest 1965: 605; Crosnier & Forest 1966: 242. (?) Alpheus Belli Coutière, 1898: 149, fig. 1. (?) Alpheus belli — Coutière 1899 (see Chace & Forest 1970 for all page numbers); Chace 1972: 63; Pequegnat & Ray 1974: 246; Christoffersen 1998: 357. (?) Alpheus malleator var. edentatus Zimmer 1913: 387, figs. G-M; Balss 1916: 22; Schmitt 1924b: 71. Not Alpheus malleator — Nobili 1901: 2; Rathbun 1902: 288; Sivertsen 1933: 4; Abele 1975: 72; Rodríguez de la Cruz 1977: 28; Brusca 1980: 252; Wicksten 1983: 43; Banner & Banner 1984: 42; Hendrickx & Wicksten 1987: 16; Kim & Abele 1988: 31, fig. 12; Wicksten & Hendrickx 1992: 5; Lemaitre & Alvarez-Leon 1992: 42; Wicksten 1993: 152; Hickman & Zimmermann 2000: 40; Wicksten & Hendrickx 2003: 64; Hendrickx & Hermoso-Salazar 2005: 433, fig. 1D (map); Lazarus-Agudelo & Cantera-Kintz 2007: 228 (= Alpheus wonkimi sp. nov., see below). Not Crangon malleator — Rathbun 1910: 607 (= Alpheus wonkimi sp. nov., see below). Material examined. Brazil: 1 male (cl 16.1 mm), MZUSP 00138, São Paulo, Ilha da Vitória (northeast of Ilhabela), leg. Gunther, 1906 [det. M.J. Rathbun, 1915]; 2 males (cl 9.5, 11.3 mm), MZUSP 22296, São Paulo, Ubatuba, Praia do Lázaro, leg. E.J. de Paula, 13 July 1976; 2 males (cl 10.6, 15.4 mm), MZUSP 13094, São Paulo, Ubatuba, north of Praia Grande, leg. M.A. Pinheiro, 18 October 1997; 1 male (cl 9.9 mm), MZUSP 28270, São Paulo, Ilha da Vitória (northeast of Ilhabela), leg. V. Cobo, December 2006. Panama: 1 male (cl 11.1 mm), OUMNH.ZC. 2012-07-0146, Caribbean coast, Isla Grande, intertidal basaltic rocks exposed to high-energy waves, leg. N. Knowlton et al., 6 June 1995 [fcn B-424]; 1 male (cl 8.3 mm), USNM, same collection data as for the previous specimen [fcn B-423]. Puerto Rico: 1 male (cl 10.8 mm), 2 ov. females (cl 10.5, 10.8 mm), MNHN-IU2010-4199, locality not specified, leg. P. Serre, 1907; 1 male (cl 12.4 mm), 1 ov. female (cl 14.0 mm), MNHN-IU2010-4194, locality not specified, bases of hard corals, leg. P. Serre, 1907. Trinidad & Tobago: 1 ov. female (cl 7.9 mm), OUMNH.ZC. 2010-01-0071, Tobago, locality not specified, collector and year unknown. Cape Verde: 1 male (cl 7.5 mm), 1 female (cl 6.0 mm), USNM, Santiago, Cidade Velha, leg. N. Knowlton et al., 3 April 1998. Senegal: 1 male (cl 7.2 mm), MNHN-Na 3109, north of Dakar, C24, leg. Postel, 1949. Guinea: 1 male (cl 13.0 mm), 2 ov. females (cl 11.0 mm, 12.1 mm), MNHN-IU-2010-4192, Tamara, Îles de Los (Loos Islands), west coast, low tide, leg. M. Serrand, 6 July 1913; 1 male (cl 11.8 mm), MNHN-IU-2010-4193, same collection data as for the previous specimens. Equatorial Guinea: 4 males, 4 females (cl 3.7–12.3 mm, male largest), MNHN-IU-2010-4196, Annobon, Punta del Palmar, intertidal, 29 July 1965, leg. A. Crosnier; 5 males, 1 ov. female, 4 females (cl 5.0–12.6 mm, ov. female largest), MNHN-IU-2010-4198, Annobon, near San Pedro, intertidal, 30 July 1965, leg. A. Crosnier. Congo: 1 male (cl 12.8 mm), MNHN-IU-2010-4197, Djeno Rocks near Pointe Noire, intertidal, small tide pools among rocky outcrops, 4 April 1965, leg. A. Crosnier. Description. Body stout, moderately large (maximum carapace length 16.1 mm, maximum total length about 50 mm). Rostrum short to moderately long, typically not reaching mid-length of first article of antennular peduncle, dorsally flattened, more or less widening posteriorly; lateral margins abruptly delimited by deep adrostral furrows; tip subacute or blunt, often descendent (Figs. 1A, B, 3A, E). Orbital hoods swollen, with acute or subacute teeth; anterior margin between each orbital tooth and rostrum with variously developed additional (= MATERIAL OF ALPHEUS MALLEATOR AND A NEW ALPHEUS SPECIES Zootaxa 3637 (4) © 2013 Magnolia Press · 415 FIGURE 1. Alpheus malleator Dana, 1852: male from Isla Grande, Panama (USNM) [A–L]; male from Puerto Rico (MNHNIU-2010-4194) [M], A, frontal region, dorsal view; B, same, lateral view; C, third maxilliped, lateral view; D, second pereiopod, lateral view; E, right third pereiopod, lateral view; F, same, distal propodus and dactylus, lateral view; G, second pleopod, lateral view; H, same, detail of appendix masculina and appendix interna; I, right uropod, dorsal view; J, left uropodal exopod, dorsal view; K, telson, dorsal view; L, left third pereiopod with aberrant number of spiniform setae, lateral view; M, detail of left uropodal exopod. 416 · Zootaxa 3637 (4) © 2013 Magnolia Press ANKER & PACHELLE FIGURE 2. Alpheus malleator Dana, 1852: male from Isla Grande, Panama (USNM), A, major cheliped, lateral view; B, major chela, lateral view; C, same, mesial view; D, dactylus of major chela, lateral view; E, minor cheliped, lateral view; F, minor chela, mesial view; G, same, lateral view. adrostral) tooth (Figs. 1A, B, 3A, E). Post-rostral area without median tubercle; pterygostomial angle rounded; cardiac notch well developed. Abdomen without specific features on sternum. Telson broad, stout, tapering posteriorly, slightly depressed along mid-dorsal line, with two stout dorsal spiniform setae and two pairs of posterolateral spiniform setae, mesial much longer than lateral (Figs. 1K, 3J); anal tubercles well developed. Antennular peduncles moderately stout, with second article longest, about 1.8 times as long as wide; stylocerite short, almost reaching distal margin of first article, with acute tip (Figs. 1A, 3A, E). Antenna with enlarged, stout basicerite bearing strong, sharp distolateral tooth reaching to or slightly beyond mid-length of second antennular article; scaphocerite large, stout; lateral margin shallowly concave, proximally with distinctly projecting, blunt tooth; blade narrow separated from strong distolateral tooth by deep cleft; distolateral tooth reaching well beyond distal end of antennular peduncle, almost reaching distal margin of carpocerite (Figs. 1A, B, 3A, E). Third maxilliped stout; coxa with blunt lateral plate; antepenultimate article not operculate, somewhat compressed and twisted, with numerous slender setae; penultimate and ultimate articles strongly compressed, very setose; penultimate article short, distally widening; ultimate article about three times as long as wide, broad, densely covered with long, very thick setae (Fig, 1C). MATERIAL OF ALPHEUS MALLEATOR AND A NEW ALPHEUS SPECIES Zootaxa 3637 (4) © 2013 Magnolia Press · 417 FIGURE 3. Alpheus malleator Dana, 1852: male from São Paulo (MZUSP 28270) [A–D]; male from Loos Islands, Guinea (MNHN-IU-2010-4193) [E–J], A, frontal region, dorsal view; B, second pereiopod, lateral view; C, third pereiopod, lateral view; D, uropodal exopod, dorsal view; E, frontal region, dorsal view; F, second pereiopod, lateral view; G, third pereiopod, lateral view; H, second pleopod (male), lateral view; I, uropodal exopod, dorsal view; J, telson, dorsal view. Major cheliped very stout; ischium very short; merus subtriangular in cross-section, stout; carpus short, cupshaped; chela broadly ovoid; lateral surface with longitudinal groove running from about mid-length of palm to end of pollex and a low crest delimiting shallow groove running obliquely from linea impressa to distodorsal third of palm; ventrolateral and dorsolateral surfaces with shallow, rounded bumps; distolateral margin of palm with two rounded bumps; mesial surface more or less rugose, with numerous setose tubercles, row of much larger tubercles present close to dorsal margin; distomesial margin of palm with very large, bluntly projecting, dorsal bump and two smaller, blunt or subacute bumps closer to propodo-dactylar articulation; pollex much shorter than dactylus, distally truncate; dactylus hammer-shaped, with short, truncate plunger, latter furnished with stamen-shaped sensillae (Figs. 2A–D, 4B, C, F, G, 5A–D). Minor cheliped stout; ischium short; merus subtriangular in cross-section, stout; carpus short, cup-shaped; chela with palm and fingers subequal in length; lateral and mesial surfaces of palm covered with groups setae, many arising from small bumps, ventral surface somewhat rugose, dorsal margin with larger, rounded bumps; distomesial margin with large, projecting, subacute tooth; fingers moderately curved, stout, setose, with sharp cutting edges, pollex somewhat excavated laterally, finger tips crossing (Figs. 2E–G, 4D, E). 418 · Zootaxa 3637 (4) © 2013 Magnolia Press ANKER & PACHELLE FIGURE 4. Alpheus malleator Dana, 1852: male from São Paulo (MZUSP 13094) [A–E]; male from São Paulo (MZUSP 28270) [F, G], A, frontal region, dorsal view; B, major cheliped, detached, lateral view; C, same, mesial view; D, minor cheliped, detached, lateral view; E, same, mesial view; F, major chela, in situ, lateral view; G, same, mesial view. Scale bars estimated. Second pereiopod with ischium and merus subequal in length; first carpal article shorter than merus, about the same length as sum of second, third and fourth articles; chela comparatively large, simple (Figs. 1D, 3B, F). Third and fourth pereiopods generally similar, stout; ischium unarmed; merus somewhat swollen, length about three times greatest width, distoventral margin unarmed; carpus with small distoventral tooth; propodus with six strong MATERIAL OF ALPHEUS MALLEATOR AND A NEW ALPHEUS SPECIES Zootaxa 3637 (4) © 2013 Magnolia Press · 419 spiniform setae along ventral margin and two (occasionally more) spiniform setae adjacent to dactylus; dactylus stout, strongly curved, biunguiculate (Figs. 1E, F, L, 3C, G). Fifth pereiopod shorter and much more slender than third and fourth (Fig. 6A); propodal brush well-developed. Second male pleopod with appendix masculina densely setose, about twice as long as appendix interna, almost reaching distal margin of endopod (Figs. 1G, H, 3H). Uropod with large subacute lobe on protopod; distolateral spiniform seta black in both sexes, stout, long, reaching to or slightly beyond distal margin of exopod, flanked by large mesial tooth and smaller lateral tooth, latter similar in size and shape to distolateral tooth of exopod; distal margin of exopod and endopod with row of short, stout, spiniform setae (Figs. 1I, J, M, 3D, I). For additional illustrations see Dana (1855), Coutière (1899), Zimmer (1913), Crosnier & Forest (1966). Variation. Alpheus malleator is remarkably variable in the configuration of the frontal margin of the carapace, especially in the width and shape of the post-rostral plate and the size (large, small, reduced to barely noticeable bumps), shape (rounded, subacute, with several distal points), direction (anterior, anteromesial) and position (marginal, submarginal) of adrostral teeth between the rostrum and orbital teeth (Figs. 1A, 3A, 4A, E, 11A–H). This variation was first reported in West African specimens by Crosnier & Forest (1966: fig. 10, a, g), but is also marked in the Caribbean and Brazilian populations (Fig. 11A–H). The chelipeds, especially the major cheliped, also show some degree of variation. Adult males have larger chelae, stronger major chelae, with more pronounced distal bumps on the palm (Figs. 4B, C, 5A, B). In females and younger males, the major chelae are smaller in size, shorter, more rounded, with shorter distal bumps on the palm (Figs. 4F, G, 5C, D; see also 6B, D). The ratio of the first carpal article of the second pereiopods is also somewhat variable (Fig. 3B, F; see also Crosnier & Forest 1966: fig. 10, e, i). The thickness of the spiniform seta on the uropodal exopod is also somewhat variable, being generally wider in males (Figs. 1I, J, M), especially in the Brazilian and West African specimens (Fig. 3D, I). However, the colour of this spiniform seta appears to be black in both males and females, contrary to the statement of Abele & Kim (1986) that the “distolateral spine on uropod of exopod [is] dark-colored in male only”. FIGURE 5. Alpheus malleator Dana, 1852: A, B, major cheliped of male from Puerto Rico (MNHN-IU-2010-4199), in situ, mesial (A) and lateral (B) views; C, major cheliped of male from Guinea (MNHN-IU-2010-4193), in situ, lateral view; D, major cheliped of male from Cape Verde (USNM), detached, lateral view. Scale bars estimated. Colour pattern. Body ground colour variable from dark chocolate-brown to reddish-brown; carapace with more or less marked, whitish, transverse band and numerous irregular patches and spots, many interconnected; rostrum dark-brown; orbital hoods, flanks and anterolateral areas of carapace mostly colourless; abdomen usually 420 · Zootaxa 3637 (4) © 2013 Magnolia Press ANKER & PACHELLE with irregular, white, transverse bands on each somite, some bands broken into rhomboid-shaped patches, bands usually continuing on pleura and splitting into two large white areas (usually two per pleuron: anterior and posterior), bands sometimes very narrow or absent (especially in larger individuals); major and minor chelipeds brown to red-brown, with some paler areas mesially, some tubercles dark brown, others marked by pale or whitish spots; dactylus of major chela red-brown centrally, pinkish dorsally and distally; pollex of major chela mostly dark brown or red-brown; adhesive disks contrasting whitish; fingers of minor chela dark or red-brown; second pereiopods and walking legs red or reddish; antennular and antennal flagella pale brown to green-yellowish; tail fan brown to red-brown, without spots, diaeresis darker brown, spiniform seta of uropodal exopod black; pleopods reddish (Fig. 6). Type locality. Rio de Janeiro ? (“in portu Rio Janeiro ?” in Dana 1852a). Distribution. Western and eastern Atlantic: Gulf of Mexico; Florida; Caribbean Sea (Panama, Cuba, Puerto Rico, Trinidad & Tobago, Barbados); Brazil (Fernando de Noronha, Rio de Janeiro, São Paulo); West Africa (Cape Verde, Senegal, Guinea, São Tomé, Annobon, Congo) (e.g., Coutière 1898, 1899; Holthuis 1951; Crosnier & Forest 1966; Chace 1972; Abele & Kim 1986; Christoffersen 1998; McClure 2005; present study). See remarks below. FIGURE 6. Alpheus malleator Dana, 1852: A, male from São Paulo, Brazil (MZUSP 28270); B, young male from the Caribbean coast of Panama (not deposited); C, male from Santiago, Cape Verde (USNM); D, young female from Santiago Cape Verde (USNM). Photographs: A, Valter Cobo; B, Carl C. Hansen / Nancy Knowlton; C, D, Nancy Knowlton. MATERIAL OF ALPHEUS MALLEATOR AND A NEW ALPHEUS SPECIES Zootaxa 3637 (4) © 2013 Magnolia Press · 421 Ecology. Rocky and mixed sand-rock shores and coral reef substrates (e.g., fossilised coral terraces, bases of hard corals), deep inside crevices, possibly rock-boring or at least capable of enlarging natural cavities. Remarks. Although the geographic range of A. malleator spans almost the entire tropical western and eastern Atlantic (Fig. 12), the species appears to be uncommon or at least to have a patchy occurrence. This is certainly due to its special microhabitat requirements, partly endolithic life style and the resulting difficulties of sampling. Alpheus malleator seems to prefer exposed shores and bioeroded hard substrates, such as porous basaltic rocks, fossilised coral terraces, bases of living corals etc., usually covered with algae and with deep crevices lined with various invertebrates (sponges, ascidians). In addition, most records of A. malleator are from islands situated at some distance from the mainland (e.g., Isla Grande, Ilha Vitória, Loos Islands) or offshore islands (Puerto Rico, Tobago, Cape Verde, São Tomé, Annobon). Thus, the beach profile, exposure and availability of suitable substrates certainly influence the distribution of A. malleator at a local scale. However, due to its cryptic habits, relative uncommonness and preference for exposed, often wave-battered rocks, A. malleator may be present in many localities where it has not yet been collected (marked with “?” in Fig. 12). For instance, despite intensive sampling efforts in presumably suitable microhabitat in São Tomé in 2006, no specimens of A. malletor were found, showing that even a targeted collecting may be unsuccessful. The presence of A. malleator in Florida requires confirmation. The only Florida record of A. malleator is based on keys in Abele & Kim (1986) who did not provide a specific locality. However, it is quite possible that A. malleator occurs in southern Florida, e.g., in the Florida Keys and/or Dry Tortugas, both offering the type of microhabitat required by this species. The presence of A. malleator in Fernando de Noronha off northeastern Brazil is based on the tentative synonymy of A. belli. Currently, A. malleator is not known from the mainland coast of northeastern Brazil. Noteworthy, the authors’ search for A. malleator on exposed rocky shores of Ceará, Rio Grande do Norte and Alagoas in 2011–2012 did not yield any specimens. Most Brazilian records of A. malleator are from much further south, in particular from the near-continental islands of Rio de Janeiro and São Paulo. Similarly, A. malleator has not (yet) been found around oceanic islands with abundance of exposed rocky shores, such as Bermuda and Ascension Island, which are marginal to the species’ geographic range (Fig. 12). Alpheus wonkimi sp. nov. (Figs. 7–10, 11I, J) Alpheus malleator — Nobili 1901: 2; Rathbun 1902: 288; Sivertsen 1933: 4; Abele 1975: 72; Rodríguez de la Cruz 1977: 28; Brusca 1980: 252; Wicksten 1983: 43; Banner & Banner 1984: 42; Hendrickx & Wicksten 1987: 16; Kim & Abele 1988: 31, fig. 12; Wicksten & Hendrickx 1992: 5; Lemaitre & Alvarez-Leon 1992: 42; Wicksten 1993: 152; Hickman & Zimmermann 2000: 40; Wicksten & Hendrickx 2003: 64; Hendrickx & Hermoso-Salazar 2005: 433, fig. 1D (map); McClure 2005: 146, fig. 16; Lazarus-Agudelo & Cantera-Kintz 2007: 228 (not A. malleator Dana, 1852). Crangon malleator — Rathbun 1910: 607 (not A. malleator Dana, 1852). Alpheus cf. malleator — Hurt et al. 2008: 516 et seq. Type material: Panama: holotype: male (cl 11.8 mm), MNHN-IU-2010-7925, Pacific coast, Río Mar, lower part of predominantly rocky intertidal, low tide, in rock crevices near water edge, leg. A. Anker, J. Jara, E. Gomez, 3 March 2006 [fcn 06-266]; paratypes: 1 male (cl 8.2 mm), USNM, same collection data as for the holotype [fcn 06265]; 1 male (cl 9.7 mm), 1 female (cl 8.3 mm), RMNH.D.55154, Pacific coast, Río Mar, lower part of predominantly rocky intertidal, low tide, in rock crevices near water edge, leg. A. Anker, J. Jara, 19 April 2007 [fcn 07-120A, 07-120B]; 1 ov. female (cl 9.3 mm), OUMNH.ZC. 2010-01-0072, Pacific coast, Río Mar, El Higo, intertidal, rock conglomerate, leg. M. Torchin, 20.02.2007 [fcn 07-080]; 1 male (cl 11.9 mm), 1 ov. female (cl 10.2 mm), MNHN-IU-2010-7926, Pacific coast, Río Mar, rocky intertidal, leg. N. Knowlton et al., 20 Feb. 1992 [fcn C361, C-362]. Additional material: Panama: 1 male (cl 11.8 mm), OUMNH.ZC. 2007-07-0145, Pacific coast, Las Perlas Islands, Isla Contadora, rocky intertidal, low tide, in rock crevices near water edge, leg. A. Anker, C. Hurt, E. Gomez, E. Tóth, J. Jara, 31 March 2006 [fcn 06-366]; 1 female (cl 9.2 mm), RMNH.D.55155, same collection data as for the previous specimen [fcn 06-371]. Ecuador: 1 male (cl 14.6 mm), MNHN-IU-2010-4195, locality not specified, collector unknown, donated by A. Crosnier in 1979. 422 · Zootaxa 3637 (4) © 2013 Magnolia Press ANKER & PACHELLE FIGURE 7. Alpheus wonkimi sp. nov.: paratype, male from Río Mar, Panama (RMNH.D.55154) [A–L]; paratype, male from Río Mar, Panama (USNM) [M, N, P]; paratype, female from Río Mar, Panama (MNHN-IU-2010-7926) [O, Q], A, frontal region, dorsal view; B, same, lateral view; C, third maxilliped, lateral view; D, major cheliped, lateral view; E, minor cheliped, lateral view; F, dactylus of major chela, lateral view; G, second pereiopod, lateral view; H, third pereiopod, lateral view; I, second pleopod, lateral view; J, same, detail of appendix masculina and appendix interna; K, uropodal exopod, dorsal view; L, telson, dorsal view; M, second pereiopod, lateral view; N, dactylus of major chela, lateral view; O, same, lateral view; P, distal uropodal exopod, dorsal view; Q, same, dorsal view. MATERIAL OF ALPHEUS MALLEATOR AND A NEW ALPHEUS SPECIES Zootaxa 3637 (4) © 2013 Magnolia Press · 423 FIGURE 8. Alpheus wonkimi sp. nov.: non-type male from Ecuador (MNHN-IU-2010-4195), A, frontal region, dorsal view; B, major chela in situ, mesial view; C, same, lateral view; D, minor chela, detached, mesial view; E, same, lateral view. Scale bars estimated. Description. For full description and illustrations of A. wonkimi sp. nov. see Kim & Abele (1988, as A. malleator); additional illustrations are provided in Figs. 7–10, 11I, J. For separation of A. wonkimi sp. nov. from A. malleator see remarks below. Variation. As in the previous species, the frontal margin of the carapace is variable in the width and shape of the post-rostral plate, the configuration (size, shape and direction) of adrostral teeth (Figs. 7A, 8A, 11I, J), and the size and proportions of the chelipeds, especially the major chela (Figs. 7D, 8B, C, 9, 10; see also Kim & Abele 1988: fig.12e, f.). Colour pattern. Body ground colour variable from purple-brown to greenish or reddish-brown; carapace with broad, whitish, transverse band and numerous irregular patches and spots, many interconnected; rostrum dark purple-, greenish- or reddish-brown; orbital hoods, flanks and anterolateral areas of carapace mostly colourless; abdomen with narrow, irregular, white, transverse bands on each somite, some bands broken into rhomboid-shaped patches, bands continuing on pleura and splitting into two large white areas (usually two per pleuron: anterior and 424 · Zootaxa 3637 (4) © 2013 Magnolia Press ANKER & PACHELLE posterior); major and minor chelipeds brownish to brown-orange, with larger pale-yellow areas mesially, some tubercles dark brown, others marked by pale spots; dactylus of major chela grey-brown centrally, pinkish dorsally and distally; pollex of major chela mostly dark brown; adhesive disks contrasting whitish; fingers of minor chela dark brown-green; second pereiopods and walking legs reddish; antennular and antennal flagella pale brownyellowish to brown-greenish; tail fan purple-, greenish- or reddish-brown, without spots, diaeresis darker brown, spiniform seta of uropodal exopod brown, somewhat paler distally; pleopods reddish-brown (Figs. 9. 10); see also colour photograph in Hickman & Zimmermann (2000). FIGURE 9. Alpheus wonkimi sp. nov.: holotype, male from Río Mar, Panama (MNHN-IU-2010-7925), dorsal (A) and lateral (B) views. MATERIAL OF ALPHEUS MALLEATOR AND A NEW ALPHEUS SPECIES Zootaxa 3637 (4) © 2013 Magnolia Press · 425 FIGURE 10. Alpheus wonkimi sp. nov.: non-type male from Las Perlas Islands, Panama (OUMNH.ZC 2012-07-0145), dorsal (A) and lateral (B, C) views. 426 · Zootaxa 3637 (4) © 2013 Magnolia Press ANKER & PACHELLE FIGURE 11. Variation of the rostro-orbital region of the carapace in Alpheus malleator Dana, 1852 [A–H] and Alpheus wonkimi sp. nov. [I, J]: A, male from Isla Grande, Panama (OUMNH.ZC 2012-07-0146); B, C, males from Puerto Rico (MNHN-IU-2010-4194, MNHN-IU-2010-4199); D, E, two males from São Paulo, Brazil (MZUSP 13094); F, male from Djeno, Congo (MNHN-IU-2010-4197); G, H, ov. female (G) and male (H) from Loos Islands, Guinea (MNHN-IU-2010-4192); I, paratype, female from Río Mar, Panama (MNHN-IU-2010-7926); J, paratype, male from Río Mar, Panama (USNM). Etymology. The new species is named after Prof. Won Kim (Seoul National University, Korea) for his important contribution to the taxonomy of the eastern Pacific members of the genus Alpheus (Kim & Abele 1988). Type locality. Río Mar, Pacific coast of Panama. Distribution. Eastern Pacific: Mexico (Baja California, Sinaloa, Nayarit, Jalisco); El Salvador (Acajutla); Costa Rica (Golfo de Nicoya); Panama (Río Mar, Las Perlas Islands); Colombia (Bahía Málaga, Isla Malpelo); Ecuador (Bahía Santa Elena, Galapagos) (Nobili 1901; Rathbun 1902; Wicksten 1983; Kim & Abele 1988; Wicksten & Hendrickx 2003; present study). Ecology. Lower intertidal of exposed rocky shores, near water edge at extreme low tides, deep inside rock crevices, possibly rock-boring or at least capable of enlarging natural cavities; at some localities, e.g., around Las Perlas Islands, sharing rock crevice microhabitat with Alpheus saxidomus Holthuis, 1980 and A. utriensis Ramos & von Prahl, 1989. Remarks. Alpheus wonkimi sp. nov. can be separated morphologically from its Atlantic sister species, A. malleator, apparently by only one character. In A. malleator, the distolateral spiniform seta of the uropodal exopod is stout and wide at its base, especially in males (length/width ratio 2.9–3.7, measured in four individuals), and is MATERIAL OF ALPHEUS MALLEATOR AND A NEW ALPHEUS SPECIES Zootaxa 3637 (4) © 2013 Magnolia Press · 427 black, sometimes with a pale-brown tip (Fig. 1I, J, M). In A. wonkimi sp. nov., this spiniform seta is more slender, relatively narrower at its base (length/width ratio 4.3–5.5, measured in three individuals) and is dark brown or tancoloured (Fig. 7K, P, Q). The brown or black colour of the uropodal spiniform seta usually persists for decades in alcohol-preserved specimens, but may fade or bleach completely in some very old or inadequately preserved specimens (e.g., in one male of A. wonkimi sp. nov. from Ecuador and in one female of A. malleator from Tobago). The two species can also be distinguished by the barcoding fragment of the COI gene sequence (Williams et al. 2001; Hurt et al. 2008). In addition, they are geographically separated from each other by the Isthmus of Panama. The frontal region of the carapace of both A. malleator and A. wonkimi sp. nov. is remarkably variable in the length and shape of the rostrum, and the development, shape and direction of adrostral teeth (Fig. 11; see also Crosnier & Forest 1966). This variability is reminiscent of the situation in another transisthmian snapping shrimp species pair, A. cylindricus Kingsley, 1878 – A. vanderbilti Boone, 1930 (Anker et al. 2008a). Although some variation was found in the proportions of the major and minor chelipeds, ratios of the carpal articles in the second pereiopod (as in A. malleator), shape of the scaphocerite blade and the uropodal diaeresis, and some other characters, none of them was consistent to be used as a differentiating feature between A. wonkimi sp. nov. and A. malleator. Similarly, no differences were found in the colour pattern of the body, chelipeds and other appendages between A. wonkimi sp. nov. and A. malleator (Figs. 6, 9, 10). However, it must be noted that the presently available information on the colour pattern of A. malleator is rather limited as photographs in Fig. 6 show immature or freshly dead specimens. Therefore, fresh material of A. malleator from both the western and eastern Atlantic, with high-quality colour photographs (like those available for A. wonkimi sp. nov.), as well as DNA sequencing and comparison of the African, Brazilian and Caribbean materials, are desirable to confirm the above proposed taxonomic assignments. FIGURE 12. Map showing distribution ranges of Alpheus malleator Dana, 1852 in the Atlantic and Alpheus wonkimi sp. nov. in the eastern Pacific. Acknowledgements The authors are most grateful to their colleagues who collected some of the specimens used in this study: Nancy Knowlton (National Museum of Natural History, Smithsonian Institution, Washington DC, USA), Javier Jara (Smithsonian Tropical Research Institute, Panama City, Panama), and Valter Cobo (Universidade de Taubaté – 428 · Zootaxa 3637 (4) © 2013 Magnolia Press ANKER & PACHELLE UNITAU, Brazil). Laure Corbari, Anouchka Sato, Paula Martin-Lefevre (MNHN), and Marcos Tavares (MZUSP) arranged loans of MNHN and MZUSP material, respectively. Charles Oliver Coleman (ZMB) provided some informations on alpheid specimens deposited in Berlin. The Panamanian material was collected with collection permits from Autoridad Nacional del Ambiente (ANAM) of the Republic of Panama, and processed at the STRI facilities in Naos, Panama City. CAPES (Coordenação de Aperfeiçoamento de Pessoal de Nível Superior) of the Brazilian Government provided financial support for this study in the form of a postdoctoral fellowship to AA. References Abele, L.G. (1975) The macruran decapod Crustacea of Malpelo Island. Smithsonian Contributions to Zoology, 176, 69–85. Abele, L.G. (1976) Comparative species composition and relative abundance of decapod crustaceans in marine habitats of Panama. Marine Biology, 38, 263–275. http://dx.doi.org/10.1007/BF00388939 Abele, L.G. & Kim, W. 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