Zootaxa 3905 (1): 091–106 www.mapress.com /zootaxa / ISSN 1175-5326 (print edition) Article Copyright © 2015 Magnolia Press ZOOTAXA ISSN 1175-5334 (online edition) http://dx.doi.org/10.11646/zootaxa.3905.1.5 http://zoobank.org/urn:lsid:zoobank.org:pub:F66C212C-1B72-44E1-B2F8-66215DBC6928 A new genus and two new species of hymenosomatid crabs (Crustacea: Brachyura: Hymenosomatidae) from the southwestern Atlantic and eastern Australia MARCOS TAVARES1,3 & WILLIAM SANTANA2 1 Museum of Zoology, University of São Paulo, Ave. Nazareth 481, Ipiranga 04263-000, São Paulo, SP, Brazil. E-mail: mdst@usp.br Laboratório de Sistemática Zoológica, Pró-Reitoria de Pesquisa e Pós-Graduação, Universidade Sagrado Coração – USC, Rua Irmã Arminda, 10-50, Jd. Brasil, 17011-160, Bauru, SP, Brazil. E-mail: william_santana@yahoo.com.br 3 Corresponding author 2 Abstract A new genus, Teramnonotus n. gen., is erected for Elamena gordonae Monod, 1956. The new genus includes two new species, T. johnlucasi n. gen., n. sp. and T. monodi n. gen., n. sp., described herein from eastern Australia (Queensland) and the southwestern Atlantic (Brazil), respectively. Teramnonotus n. gen. superficially resembles Elamena H. Milne Edwards, 1837, and Trigonoplax H. Milne Edwards, 1853, of which it can be easily distinguished by a combination of characters: eyes and ocular peduncle, rostrum, carapace, thoracic sternite 8 and thoracic pleurite 8, and the inhalant water openings. The validity of the obscure species Elamena mexicana H. Milne Edwards, 1853, is not supported and it is synonymised with Halicarcinus planatus (Fabricius, 1775). Key words: Australia, Brazil, biodiversity, false spider crab, Elamena, Decapoda, Eubrachyura Introduction The family Hymenosomatidae consists of approximately 118 species in 19 genera, mostly Indo-West Pacific (Guinot 2011a). Many hymenosomatid species are conspicuously small and cryptic, and many are only known from their type material. Such is the case of Elamena gordonae Monod, 1956, to which Monod referred to as "... une très petite forme, qui risque de passer inaperçue" (Monod 1956: 472). It was originally described from two small females (one ovigerous) obtained in 1948 in the Atlantic coast of Africa and never again collected. Two ovigerous females collected in 1952 from eastern Australia (Queensland) were assigned to E. gordonae by Lucas (1980), but additional specimens have not been recorded from Australia ever since. Three ovigerous females were dredged in 1994 from Rio de Janeiro State (southeastern Brazil) by the first author. Additional specimens attributed to E. gordonae were obtained in 2002 and following years from northeastern Brazil (Almeida et al. 2007; Almeida & Coelho 2008; Coelho Filho & Coelho 2002; Coelho et al. 2008). The Atlantic and Australian specimens of Elamena gordonae are said to form a single species, but such status has not been conclusively established because the above-mentioned authors did not compare their specimens directly with West African ones. A direct comparison of the southwestern Atlantic and the Australian specimens with the type specimens of E. gordonae revealed that they actually belong to two undescribed species. These two new species are described herein. A comparison between Elamena gordonae s. str. and representatives of the genus Elamena (including its type species) showed that E. gordonae cannot be retained in Elamena H. Milne Edwards, 1837. A new genus, Teramnonotus n. gen., is therefore proposed herein to receive E. gordonae Monod, 1956, along with the new species from the southwestern Atlantic and eastern Australia, namely Teramnonotus monodi n. gen., n. sp. and Teramnonotus johnlucasi n. gen., n. sp. The descriptive terminology follows that used by Lucas (1980) with additions from Ng & Chuang (1996) and Guinot (2011a, b). Abbreviations: Mxp3, third maxilliped; P1, cheliped; P2–P5, second to fifth pereiopods; Pl2–Pl5, second to fifth pleopods; PlT, pleotelson; St 8, thoracic sternite 8. Measurements are expressed in Accepted by P. Castro: 24 Nov. 2014; published: 9 Jan. 2014 91 millimeters (mm): carapace length (cl); carapace width (cw). The material studied has been deposited in or belongs to the collections of the following museums: AM (Australian Museum, Sydney), MNHN (Muséum national d'Histoire naturelle, Paris), MZUSP (Museu de Zoologia da Universidade de São Paulo), UESC (Universidade Estadual de Santa Cruz), RMNH (Nationaal Natuurhistorisch Museum, Leiden), WAM (Westerm Australian Museum, Perth), and ZRC (Zoological Reference Collection, Lee Kong Chian Natural History Museum, National University of Singapore). Taxonomy Family Hymenosomatidae MacLeay, 1838 Teramnonotus n. gen. Elamena (pro parte)—Monod 1956: 469; Lucas 1980: 170–171; Ng & Chuang 1996: 6; Ng et al. 2008: 108; Guinot et al. 2013: 191 [not Elamena H. Milne Edwards, 1837. Type species Hymenosoma mathoei Desmarest, 1823]. Type species. Elamena gordonae Monod, 1956, by original designation. Gender masculine. Included species. Teramnonotus gordonae (Monod, 1956) n. gen., n. comb., eastern Atlantic (between Conakry, Guinea and Monrovia, Liberia), T. johnlucasi n. gen., n. sp., western Pacific (Australia: Queensland), and T. monodi n. gen., n. sp., southwestern Atlantic (Brazil: Ceará, Rio Grande do Norte, Sergipe, Bahia and Rio de Janeiro). Comparative material. Elamena longidactylis Yang & Sun, 1998: 1 male cl 2.8 mm, cw 2.3 mm, 1 juvenile female cl 2.5 mm, cw 2.2 mm, Xiamen, Gu-Lang-Yu, Fujian Province, China, mud flat, under stones, 13.xii.1984 (ZRC 2004.0736). Hymenosoma mathoei Desmarest, 1823: Gesira, Somalia, coll. x.1981, reef, two females, one juvenile female (ZRC 1994.4235). Gesira, Somalia, coll. x.1981, reef, young male (ZRC 1994.4234). Trigonoplax spathulifera Lucas, 1980: Western Australia, Woodside Dampier Expedition 2, Dampier Archipelago, Brigadier Island, 4.8 km E of East Point, stn DA2/99/85, between 20º26.38'S, 116º39.76'E and 20º26.09'S, 116º40.10'E, S. Slack-Smith & M. Hewitt coll., 26.vii.1999, 29 m, 1 ovigerous female (WAM C27116); Woodside Dampier Expedition 3, Dampier Archipelago, Nelson Rocks, stn DA3/99/68, 20º27.99'S, 116º39.71'E, M. Hewitt coll., 7.ix.1999, 6 m, 1 juvenile female (WAM C27685). Woodside Dampier Expedition 1, Dampier Archipelago, Angel Island, stn DA1/98/09, 20º28.69'S, 116º47.95'E, M. Hewitt coll., 20.x.1998, 1 juvenile female (WAM C25639), 1 male (WAM C25640); Searipple Passage, Burrup Peninsula, 20º31'S, 116º51'E, D. S. Jones & B. Morton coll., 7.x.1999, 1 juvenile female (WAM C33088). Trigonoplax unguiformis H. Milne Edwards, 1853: Indonesia, Moluccas, Ambon, Hitu, Ambon Bay, outer bay, in front of Ruhmatiga, up to river Lela; Rumphius Biohistorial Expedition, stn 41, L.B. Holthuis coll., 10.xii.1990, dredge, 4–28 m, female (RMNH). Diagnosis. Carapace pear-shaped, moderately convex, dorsal carapace surface with faint longitudinal ridge, edges slightly upturned; rostrum triangular, upturned, continuous with carapace surface; ventral rostral keel absent. Epistome long, almost as long as broad; pterygostomial region with 2 prominent blunt teeth. No orbits, eyes-stalk immovable, short, stout, fused with carapace; most of eyes hidden beneath rostrum in dorsal view, only cornea visible dorsally; small post-ocular tooth present; cornea pigmented. Antennules fold beneath rostrum; interantenullar septum short, reaching to less than half-length of second antennular article. Antenna conspicuously slender, short, concealed by rostrum in dorsal view. Third maxillipeds broad, almost completely filling mouth square, ischium subequal to merus along lateral edge; merus heart-shaped; epipod lamella about same size as basal portion (operculum); palp articulating near merus anterolateral angle; exopod with distinct flagellum. Chelipeds slender, subequal, stouter than pereiopods; dactyli usually with 2 subterminal teeth. Thoracic sternite 4 extended laterally, conspicuously convex anteriorly distinctly separating Milne Edwards openings from chelipeds. Thoracic sternite 8 fully covered by carapace in dorsal and posterior views. Female abdomen very large, broader than cephalothorax width, soft; somites 1, 2 free, somites 3–5, fused; somite 6 separated from previous somites, fused with telson; Pl2–Pl5 articulated close to lateral edge of somites enlarging brood cavity. Vulvae located on medially fused part of thoracic sternites, oblique, slit-like, peripheral border swollen. Remarks. Elamena H. Milne Edwards, 1837 (type species Hymenosoma mathoei Desmarest, 1823) is the second largest hymenosomatid genus in terms of species, 25 species (Ng et al. 2008). Elamena has been reviewed 92 · Zootaxa 3905 (1) © 2015 Magnolia Press TAVARES & SANTANA and partially revised in a number of occasions (e.g., Alcock 1900, Kemp 1917, Tesch 1918, Gordon 1940, Melrose 1975, Lucas 1980, Ng & Chuang 1996), yet its species composition is still in need of a reappraisal. Guinot (2011b), for instance, commented on the puzzling position of Elamena truncata (Stimpson, 1858) in Elamena, and Ng & Chuang (1996) stated that E. magna Ng & Chuang, 1996, might actually belong in a new genus. Elamena longirostris Filhol, 1885, also leaves doubt as to its generic position; Filhol (1885) stated that E. longirostris belong in the same genus as Elamena whitei Miers, 1876, a species subsequently reallocated in the genus Halicarcinus White, 1846 (Ng et al. 2008). A comparison between Elamena gordonae Monod, 1956 s. str. and representatives of Elamena (including its type species) revealed that E. gordonae can no longer be attributed to Elamena, nor to the closely related genus Trigonoplax H. Milne Edwards, 1853. Elamena gordonae Monod, 1956 s. str. is therefore herein transferred to Teramnonotus n. gen. as its type species along with the two new species being described, one from the southwestern Atlantic and a second from eastern Australia. The genus Teramnonotus n. gen. can be distinguished from Elamena H. Milne Edwards, 1837 s. str. in having: (i) an immovable ocular peduncle, completely fused to the carapace (Figs. 1B, D, 5B, C) (freely movable in Elamena, Fig. 5D); (ii) rostral keel absent (Figs. 5A–C, 6A, B) (a distinct, vertical keel on the ventral surface of the rostrum in Elamena; Figs. 5D, 6C); (iii) subhepatic region with two strong lobes (Figs. 1B, D, 4A–C, 5B, C) (smooth in Elamena; Figs. 2B, 5D); (iv) thoracic sternite 8 concealed by the carapace in dorsal and posterior views (Fig. 6D) (thoracic sterni.te 8 partially visible in dorsal and posterior views between the carapace, the abdominal somites 1 and 2, and the P5 coxa in Elamena; Fig. 6F); (v) narrow thoracic sternite 8 so that the P5 coxae are close to each other (Fig. 1A, B) (wide thoracic sternite, so that the coxae of P5 are far from each other in Elamena; Fig. 2A); (vi) first female abdominal somite distinctly narrower than the second (Fig. 6D) (somites 1 and 2 of about the same size in Elamena); (vii) female abdominal somites 1 and 2 free, somites 3 to 5 fused, lateral clefts still discernible even if minute, and a pleotelson (Fig. 7 A–D) (somites 1–5 free and a pleotelson in Elamena). Trigonoplax H. Milne Edwards, 1853 [type species Ocypode (Trigonoplax) unguiformis De Haan, 1839, by monotypy] share a number of characters with Elamena and, in the past, was even considered a subgenus of Elamena by many authors (e.g., Alcock 1900; Kemp 1917; Gordon 1940). Trigonoplax can be distinguished from both Elamena and Teramnonotus n. gen. by (i) the inhalant water openings being fused laterally for more than half of its length (see Lucas, 1980: 154) (not fused in Elamena and Teramnonotus n. gen.; Fig. 5 B, C), and (ii) a large portion of the female thoracic sternite 8 is not dorsolaterally covered by the carapace (Fig. 6E) (thoracic sternite 8 fully covered by the carapace in Teramnonotus n. gen. and a small portion of thoracic sternite 8 dorsally exposed in Elamena in dorsal and posterior views respectively; Fig. 6D, F). Trigonoplax also differs from Teramnonotus n. gen. in the presence of well-developed keel on the ventral surface of the rostrum (rostral keel absent in Teramnonotus n. gen.; Figs. 5A–C, 6A, B). Etymology. The generic name Teramnonotus is a combination of the Greek words teramnos (soft) and notus (back), in allusion to the soft and smooth dorsal surface of the carapace. Gender masculine. Teramnonotus gordonae (Monod, 1956) n. gen., n. comb. (Figs. 4A, 5A, 8A–C) Elamena (Trigonoplax) gordonae Monod, 1956: 469, figs. 629–637. Elamena gordonae—Lucas 1980: 171 [pro parte]; Ng & Chuang 1996: 6 [pro parte]; Yang & Sun 1998: 2; Almeida et al. 2007: 29 [pro parte]; Almeida & Coelho 2008: 197 [pro parte]; Coelho et al. 2008: 18 [pro parte]; Ng et al. 2008: 108. Material examined. Elamena (Trigonoplax) gordonae Monod, 1956: ovigerous female holotype cl. 3.1 mm, cw. 2.6 mm, West Africa, between Conakry and Monrovia, Debyser coll., 1–8.iii.1948, 30–40 m (MNHN-B25803); Sierra Leone, Trawler Maid Honour, J. Cadenat coll., 1–8.iii.1948, between 8º38'N and 8º42'N, 8–12 m, female paratype (abdomen absent) cl. 2.4 mm, cw. 2.2 mm (MNHN-B25802, currently MNHN-IU-2014-4029). Diagnosis. Rostrum well detached from the carapace outline, long, outreaching by far the second antennular article, apex sharp. Pterygostomial lobes large, first ending in acute tip, second much smaller but well discernible, followed by strong branchiostegal carina extending posteriorly to level of P3/P4. Subhepatic lobes clearly of distinct sizes, first much larger, lobes visible dorsally. Antennules not visible dorsally when folded. Tuft of setae on metagastric region of carapace small. Cheliped fingers moderately curved, laterally compressed; fingers edentate along cutting margins, not interlocking distally; fixed finger distal end inflated, minutely serrate, denticles rounded. Male unknown. NEW HYMENOSOMATID CRABS Zootaxa 3905 (1) © 2015 Magnolia Press · 93 Remarks. Teramnonotus gordonae (Monod, 1956) n. gen., n. comb., described by Monod (1956) as Elamena gordonae from two females (one ovigerous) collected between Conakry and Monrovia, and from Sierra Leone, has never been reported from West Africa since its collection in 1948. The male of the species is currently unknown. In T. gordonae, the P2–P5 dactyli end in an acute unguis preceded by two ventral, subdistal spines. Monod (1956: 470, fig. 630) erroneously figured only one ventral, subdistal spine in P5, but his figure 636 of the P5 and his description of the pereiopods are correct: "Péreiopodes grêles, avec au bord supéro-distal du mérus une forte saillie tuberculiforme; dactyles 3-dentés (1 apicale + 2 subapicales internes)" (Monod 1956: 472, fig. 636). Teramnonotus gordonae (Monod) has been mistakenly recorded as Elamena gordonae from Australia (by Lucas 1980) and from Brazil (by Coelho Filho & Coelho 2002; Almeida et al. 2007; Almeida & Coelho 2008; Coelho et al. 2008). The Australian and the Brazilian specimens actually represent two separate new species herein described. Yang & Sun (1998) commented on the supposed resemblance between Elamena longidactylis Yang & Sun 1998, and Teramnonotus gordonae (Monod, 1956) n. gen., n. comb. (as Elamena gordonae). Their resemblance, however, is only superficial because in E. longidactylis (as typical for Elamena), the ocular peduncle is not fused to the carapace and is freely movable, and the female thoracic sternite 8 is not concealed by the carapace in dorsal and posterior views (in contrast to Teramnonotus n. gen., see above). As stated by Yang & Sun (1998), the examined male and female of E. longidactylis also have thinner dactyli with three subterminal teeth on their ventral surface (instead of two in T. gordonae). Both sexes nevertheless have one anterior prominent pterygostomial lobe followed by a much smaller one (as in T. gordonae). Distribution. So far known only from West Africa (between Conakry, Guinea and Monrovia, Liberia and from Sierra Leone), between 8 and 40 meters deep. Teramnonotus johnlucasi n. gen., n. sp. (Figs. 1A–B, 4B, 5B, 6A, 7A–B, 8D–F) Elamena gordonae—Lucas 1980: 174, figs. 2B, 6E, Ng & Chuang 1996: 6 [pro parte], Davie 2002: 243, Almeida et al. 2007: 29 [pro parte], Almeida & Coelho 2008: 197 [pro parte], Coelho et al. 2008: 18 [pro parte]. [Not Elamena (Trigonoplax) gordonae Monod, 1956] Type material. Ovigerous female holotype cl. 2.8 mm, cw. 2.7 mm, P1–P5 detached from the body, Australia, Queensland, Tyroom Roads, Gt. Sandy Strait, J. S. Hynd coll., iii.1952, from a spiny alcyonarian from 18 m (10 fms) (AM P.12208); ovigerous female paratype cl. 2.5 mm, cw. 2.4 mm, P1–P5 detached from the body, same data (MZUSP 32909). Type locality. Australia, Queensland, Tyroom Roads, Gt. Sandy Strait, 18 m. Diagnosis. Rostrum integrated into carapace outline, short, not outreaching second antennular article when fully extended in dorsal view, apex rounded. Tuft of setae on metagastric region of carapace absent. Inconspicuous postocular teeth. Antennules visible dorsally when folded. Antennular basal article with distinct but short anteroexternal tooth. Subhepatic region with 2 pronounced lobes of about same size. Mxp3 exopod nearly reaching distal margin of mxp3 merus. Mxp3 dactylus longer than propodus. Teeth of cutting edges of cheliped small, equal. Two pterygostomian lobes. Description. Carapace pear-shaped, longer than wide, slightly convex; regions undefined, except for poorly marked gastrocardiac groove; dorsal surface with faint longitudinal ridge, without setae. Carapace anterolateral, posterolateral angles almost imperceptible, posterior margin rounded. Narrow sharp edged rim around carapace, rostrum. Rostrum straight, directed forward, broadly triangular, short, without subrostral keel; with minute setae on lateral margin, ventral surface. Cornea visible dorsally. Postocular tooth short. Carapace without orbits. Eyestalk immovable, with protuberant anteromesial lobe. Antennules short, stout, visible dorsally when folded, 3 peduncular articles subequal in length; basal article subquadrangular, with acute anteroexternal tooth; second somewhat longer, slender than first; distal article thinner. Interantennular septum small. Antenna very slender, short, concealed by rostrum in dorsal view. Epistome as long as wide. Subhepatic region with 2 marked lobes of approximately same size. Subtriangular pterygostomian region large, with 2 lobes; first prominent, acute, second smaller. Anterolateral angle of buccal frame projected anteroventrally. Setae absent. 94 · Zootaxa 3905 (1) © 2015 Magnolia Press TAVARES & SANTANA FIGURE 1. A–D, habitus, dorsal and ventral views. A, B, Teramnonotus johnlucasi n. gen., n. sp., ovigerous female holotype cl. 2.8 mm, cw. 2.7 mm (AM P.12208). C, D, Teramnonotus monodi n. gen., n. sp., ovigerous female paratype cl. 4.3 mm, cw. 3.6 mm (MZUSP 10273). Note (arrows) rostrum integrated into the carapace outline in A (versus rostrum well detached from the carapace outline in C). A–D, stained with methylene blue. Scale bars: A–D, 1 mm. Mxp3 exopod long, reaching distal margin of merus, with sparse setae on inner margin. Ischium as wide as long, with dense row of stout, short setae on inner margin. Merus slightly wider than long, heart-shaped, with dense row of long setae on inner margin. Carpus articulating almost on lateral lobe of merus, with setae distally on inner margin. Dactylus longer than propodus; dactylus, propodus covered with setae, mostly on inner margin; dactylus fairly stout, bluntly pointed. Inhalant water opening bordered by long setae. Basal portion of epipod strongly curved. NEW HYMENOSOMATID CRABS Zootaxa 3905 (1) © 2015 Magnolia Press · 95 P1 slender. Ischium short, narrow proximally, with few setae, without projection over merus. Merus long, slender, without spines or tooth, sparsely setous. Carpus short, without projections, with few setae. Palm narrower than long, hardly expanded laterally, vaguely shorter than merus. Fingers slender, slightly longer than palm, rounded, with very small gap in proximal third; small, equal teeth on both fingers, evenly distributed. Pereiopods similar, slightly decreasing in size from P2–P5. Ischium rounded. Merus with long spine on anterodorsal margin, longer than remaining articles, with few setae. Carpus shorter, without anterior projections, sparsely setate. Dactylus shorter than propodus, curved distally, 2 strong subterminal spines almost of same size on ventral margin. Propodus, dactylus flattened laterally, with few long setae, sparse short hooked setae. Ventral margin of dactylus with row of setae. Female abdomen much wider than long, somites 1, 2 free, somites 3–5 fused, somite 6 separate from previous somites, fused with telson. Abdominal margins with short setae; setae sparsely distributed over abdominal surface. Pleopods 2–5 well developed, projecting laterally beyond abdomen providing greater volume for carrying eggs. Etymology. The new species is named for John S. Lucas (James Cook University, Townsville, Australia) in recognition of his contributions to the knowledge of hymenosomatid crabs. Remarks. The two ovigerous females from eastern Australia that form the basis for the description of the present new species were formerly considered by Lucas (1980: 174–175) as conspecific with Elamena gordonae Monod, 1956, from the eastern Atlantic. The type material of E. gordonae was not available at that time for him to compare. A direct comparison between the types of E. gordonae and the Australian specimens has now shown that the Australian material should be assigned to a new species in Teramnonotus n. gen., along with E. gordonae Monod, 1956 (see above). Teramnonotus johnlucasi n. gen., n. sp. can be separated from T. gordonae (Monod, 1956) n. comb. in having: (i) rostrum integrated into the carapace outline, short, not outreaching the second antennular article when fully extended in dorsal view, apex rounded (Figs. 1A; 4B; 5B) (rostrum well detached from the carapace outline, long, outreaching by far the second antennular article, apex sharper in T. gordonae; Figs. 4A; 5A); (ii) pterygostomial lobes small, first rounded, second weak (Fig. 5B) (pterygostomial lobes distinctly large, first ending in an acute tip, second much smaller but clearly discernible in T. gordonae); (iii) subhepatic lobes about the same size (Fig. 5B) (subhepatic lobes clearly of distinct sizes, first much larger in T. gordonae); (iv) antennules visible dorsally when folded (Fig. 5B) (antennules not visible dorsally in T. gordonae; Fig. 5A); and (v) no tuft of setae on metagastric region of the carapace (Fig. 4B) (small tuft of setae on metagastric region in T. gordonae). It also appears that T. johnlucasi is smaller in overall size than T. gordonae, the smallest mature female being only 2.5 mm (cl) (cl. 3.1 mm in T. gordonae). Lucas (1980: 175) had noticed its minute size: "E. gordonae [presently T. johnlucasi] is smaller than the other 'triangular' Elamena species, except E. umerata n. sp". In T. johnlucasi, the P2–P4 dactyli are armed with two subterminal teeth of about the same size. The number of subterminal teeth in the P5 seems variable. Although in the type specimens the P1–P5 are detached from the body, the P5 is readily recognizable because of its smaller size. One of the P5 pairs has only one subterminal tooth, the distal one, whereas the other pair has two subterminal teeth, though the proximal tooth is tiny and hardly recognizable. The examination of additional specimens of T. johnlucasi will be necessary to determine whether the bidentate P5 is another distinctive character between T. johnlucasi and T. gordonae or just variation. It must be noted that Monod (1956) figured the holotype of E. gordonae with tridentate P2–P4, while the P5 lack the subterminal tooth (Monod 1956: 470, fig. 630). The P5, however, is tridentate in his figure of the paratype (Monod 1956: 471, fig. 636). Monod (1956: 472) maintained in his description that the P2–P5 are tridentate: "Péréiopodes grêles, avec au bord supéro-distal du mérus une forte saillie tuberculiforme; dactyles 3-dentés (1 apicale + 2 subapicales internes)". Because the left P5 dactylus of the holotype is broken and the right P5 missing, and the dissected appendages from the paratype are also missing, it is not possible to verify if the P5 is bi- or tridentate in E. gordonae s. str. Distribution. So far known from eastern Australia (Queensland, Tyroom Roads, Gt. Sandy Strait, 18 m deep). 96 · Zootaxa 3905 (1) © 2015 Magnolia Press TAVARES & SANTANA FIGURE 2. A–D, habitus, dorsal and ventral views. A, B, Elamena mathoei (Desmarest, 1823), female (ZRC 1994.4235). C, D, Trigonoplax unguiformis H. Milne Edwards, 1853, female (RMNH). Note (arrows) part of the thoracic pleurite 8 visible between the carapace, the abdominal somites 1–2, and the coxa of P5 in A and C. A–D, stained with methylene blue. Scales: A, B, 1 mm; C, D, 2 mm. Teramnonotus monodi n. gen., n. sp. (Figs. 1C–D, 3, 4C, 5C, 6B, D, 7C–D, 8G–I) Elamena gordonae—Almeida et al. 2007: 29, figs. 7 A–I; Almeida & Coelho 2008: 197; Almeida et al. 2010: 340; Coelho & Coelho Filho 2002: 124; Coelho et al. 2008: 18. [Not Elamena (Trigonoplax) gordonae Monod, 1956] Type material. Holotype: Brazil: Rio de Janeiro, Baía de Sepetiba, Ponta da Boa Vista, Itacuruçá Island, M. Tavares coll., 4.vii. 1994, dredged from about 15 m on biogenic gravel: ovigerous female, cl. 4.0 mm, cw. 3.5 mm (MZUSP 10272). Paratypes: Brazil: Ceará, Praia do Pecém, L. E. Bezerra coll. 13.x.2010, 1 female (MZUSP 28399). Rio Grande do Norte, Areia Branca, Praia de Baixa Grande, 04°55’44.18”S, 38°30’51.840”W, rocky intertidal, P. Pachelle coll. 29.ix.2011, 1 ovigerous female (MZUSP 29814), 1 ovigerous female, 1 juvenile female (ZRC). Bahia, Baía de Todos os Santos, PROMARLAN, stn CAB1, 12°44’22.920”S, 38°30’51.840”W, i.2005, 1 female, P1–P5 detached from the body (MZUSP 24204). Baía de Todos os Santos, PROMARLAN, stn CAB1, NEW HYMENOSOMATID CRABS Zootaxa 3905 (1) © 2015 Magnolia Press · 97 12°44’22.920”S, 38°30’51.840”W, viii.2004, 1 female, P1–P5 detached from the body (MZUSP 24205). Baía de Todos os Santos, Porto da Barra, 13°00’05”S, 38°32’01”W, 4–6 m, associated with Callyspongia sp. (Porifera), C. Menegola & L. Martins coll. 20.xi.2012, 4 females (MZUSP 29178), 2 females (AM P.97355). Bahia, Baía de Camamu, stn 5, 13º54’14”S, 39º00’34”W, M. C. Cuerrazi coll., 29.viii.2004, 1 ovigerous female (UESC 719). Idem, 25.ix.2004, 1 ovigerous female (UESC 718). Caravelas, Rio Caravelas, stn 1, 17°44’39.4’S, 39°14’49.7”W, A. O. Almeida coll., 28.iii.2007, 1 ovigerous female each (MZUSP 32496, UESC 1090). Bahia, Nova Viçosa, Praia do Pontal da Barra, stn 3, 17°53’00.9”S, 39°21’48.2”W, A. O. Almeida coll., 19.iii.2007, 1 female (UESC 837). Rio de Janeiro, Baía de Sepetiba, Ponta da Boa Vista, Itacuruçá Island, M. Tavares coll., 4.vii. 1994, dredged from about 15 m on biogenic gravel: 2 ovigerous females, cl. 4.3 mm, cw. 3.6 mm and cl. 3.6 mm, cw. 2.8 mm (MZUSP 10273). Type locality. Brazil, Ponta da Boa Vista, Itacuruçá Island, Baía de Sepetiba, Estado do Rio de Janeiro. About 15 m deep. Diagnosis. Rostrum well detached from the carapace outline, long, outreached by second antennular article, curved upward, apex sharp. Small tuft of setae on metagastric region. Prominent postocular teeth. Antennules not visible dorsally when folded. Antennular basal article with acute, long anteroexternal tooth. Subhepatic region with 2 lobes, first larger, prominent, second tumescent. Mxp3 exopod nearly reaching distal margin of mxp3 merus. Mxp3 dactylus, propodus of about same length. Teeth of cutting edges of cheliped small, of different sizes. Three pterygostomian lobes. Description. Carapace pear-shaped, longer than wide, moderately convex; regions undefined, except for poorly marked gastrocardiac groove; dorsal surface with faint longitudinal ridge, with only small tuft of setae on metagastric region. Carapace with discrete anterolateral angle; posterolateral margin slightly angled; posterior margin markedly curved. Narrow, sharp edged rim around carapace, rostrum. Rostrum triangular, short, without subrostral keel, curved upward anteriorly; with minute setae on lateral margin, ventral surface. Cornea visible dorsally. Postocular tooth prominent. Carapace without orbits. Eyestalk immovable, with protuberant anteromesial lobe. Antennules short, stout, not visible dorsally when folded, 3 peduncular articles subequal in length; basal article subquadrangular, with acute anteroexternal tooth; second somewhat longer, more slender than first; distal article thinner. Interantennular septum small. Antenna conspicuously slender, short, concealed by rostrum in dorsal view. Epistome as long as wide. Subhepatic region with 2 lobes; first larger, prominent; second tumescent, undefined. Subtriangular pterygostomian region large, with 3 lobes; first conspicuously prominent, acute; second, third about the same size. Anterolateral angle of the buccal frame projected anteroventrally. Setae absent. Third maxillipeds expopods long, nearly reaching distal margin of merus, with sparse setae on inner margin. Ischium as wide as long, with dense row of stout short setae on inner margin. Merus slightly wider than long, with dense row of long setae on inner margin. Carpus articulating almost centrally between 2 distal merus lobes, setae distally on inner margin. Dactylus, propodus about same length, covered with setae, mostly on inner margin; dactylus fairly stout, bluntly pointed. Inhalant water opening bordered by long setae. Basal portion of epipod strongly curved, single row of long setae traversing anterior edge, separated by suture from lamellar distal portion. Lamellar portion of epipod as long as basal article, slender, fringed with long setae, rounded terminally. Cheliped slender. Ischium long, narrow proximally, with few setae, without projection over merus. Merus long, slender; without spines or tooth, setae sparsely. Carpus short, without projections, few setae. Palm narrower than long, hardly expanded laterally, slightly shorter than merus. Fingers slender, slightly longer than palm, rounded with small gap in proximal third; small teeth of different sizes on both fingers, evenly distributed. Pereiopods similar in size, slightly decreasing in size from P2–P5. Ischium rounded. Merus longer than remaining articles, with long spine on anterodorsal margin. Carpus shorter, without anterior projections, setae sparsely distributed. Dactylus shorter than propodus, curved distally, 2 strong subterminal spines in ventral margin, subequal in size. Propodus, dactylus flattened laterally, with few long setae, sparse short hooked setae. Ventral margin of dactylus with dense row of setae. Female abdomen much wider than long, somites 1, 2 free, somites 3–5 fused, somite 6 separated from previous somites, fused with telson. Row of slender short setae on abdominal margin, setae sparsely distributed on surface. Pleopods 2–5 well developed, projecting laterally beyond abdomen providing greater volume for carrying egg. Etymology. The present new species is dedicated to the eminent French researcher Théodore André Monod (1902–2000), chercheur d’absolu and professor at the Muséum national d’Histoire naturelle, Paris. 98 · Zootaxa 3905 (1) © 2015 Magnolia Press TAVARES & SANTANA FIGURE 3. A, Teramnonotus monodi n. gen., n. sp., ovigerous female paratype (ZRC). B, ovigerous female holotype of Elamena mexicana H. Milne Edwards, 1853, (MNHN-B655, currently MNHN-IU-2000-655), a junior synonym of Halicarcinus planatus (Fabricius, 1775). Photographs by A. Anker and T. Naruse, respectively. Scale: A, 5 mm. Remarks. Teramnonotus monodi n. gen., n. sp. closely resembles T. gordonae (Monod, 1956) n. comb., from which it can be distinguished in having: (i) both cheliped fingers markedly arcuate and deeply excavated along the mesial surface and spoon-shaped (moderately curved and laterally compressed in T. gordonae), (ii) fingers strongly NEW HYMENOSOMATID CRABS Zootaxa 3905 (1) © 2015 Magnolia Press · 99 dentate along the entire length of the cutting margins (Fig. 8G) (cutting margins edentate in T. gordonae; Fig. 8A), and (iii) the fixed finger of the cheliped ending in a strong bifid tip interlocking with the single, strong tip of dactylus (Fig. 8G) (no interlocking mechanism, finger distal end inflated, minutely serrate, denticles rounded in T. gordonae; Fig. 8A). The pterygostomial lobe in the new species is also followed by a distinctly smaller lobe continued by a low branchiostegal carina extending backwards to the level of P3/P4 but by a much stronger branchiostegal carina in T. gordonae s. str. FIGURE 4. A–D, habitus, dorsal view. A, Teramnonotus gordonae (Monod, 1956) n. gen., n. comb., holotype (after Monod 1956: fig. 629). B, Teramnonotus johnlucasi n. gen., n. sp., ovigerous female holotype cl. 2.8 mm, cw. 2.7 mm (AM P.12208). C, Teramnonotus monodi n. gen., n. sp., ovigerous female paratype cl. 4.3 mm, cw. 3.6 mm (MZUSP 10273). D, Elamena mathoei (Desmarest, 1823), female (ZRC 1994.4235). Scales: A–D, 1 mm. 100 · Zootaxa 3905 (1) © 2015 Magnolia Press TAVARES & SANTANA FIGURE 5. A–D, anteroventral view of the cephalothorax. A, Teramnonotus gordonae (Monod, 1956) n. gen., n. comb., holotype (after Monod 1956: fig. 630 bis). B, Teramnonotus johnlucasi n. gen., n. sp., ovigerous female holotype cl. 2.8 mm, cw. 2.7 mm (AM P.12208). C, Teramnonotus monodi n. gen., n. sp., ovigerous female paratype cl. 4.3 mm, cw. 3.6 mm (MZUSP 10273). D, Elamena mathoei (Desmarest, 1823), female (ZRC 1994.4235). Arrows indicate the rostrum, which is integrated into the carapace outline in B (rostrum well detached from the carapace outline in C). Scales: A–D, 1 mm. Monod (1956) figured the paratype of T. gordonae as having the merus of both chelipeds ending in a blunt tooth at its inner distal angle (Monod 1956: 471, figs. 632–633), but no tooth is visible in the illustration of the merus of the holotype (Monod 1956: 470, fig. 630). Monod (1956: 472) described the cheliped as being inermis: "Chélipèdes grêles, doigts plus longs que la paume et incurvés transversalement vers l'intérieur, jointifs, inermes". The merus of the left cheliped of the holotype of T. gordonae s. str. actually ends in a minute tooth, hardly recognizable at its inner distal angle; the right cheliped of the holotype, and the dissected appendages from the paratype are lost. Confirmation of whether the merus of the cheliped is armed with an acute spine will prove to be an additional difference between T. monodi n. sp. and T. gordonae s. str., but it await confirmation by the availability of additional material of T. gordonae (Monod, 1956) s. str. Compared with T. gordonae s. str., T. monodi n. sp. also appears to be larger in size (largest ovigerous females of both species measuring cl 4.0, cw 3.5 mm versus cl 3.1, cw 2.6, respectively). Teramnonotus monodi n. gen., n. sp. and T. johnlucasi n. gen., n. sp. can be separated from each other by the presence in the former species of: (i) rostrum well detached from the carapace outline, long, outreaching by far the second antennular article, curved upward, apex sharper (Figs. 1C, D; 3A; 4C) (rostrum integrated into the carapace outline, short, not outreaching the second antennular article when fully extended in dorsal view, straight, apex rounded in T. johnlucasi n. gen., n. sp.; Figs. 1A, B; 4B); (ii) a small tuft of setae on metagastric region (Fig. 4C) (setae absent in T. johnlucasi n. gen., n. sp.; Fig. 4B); (iii) prominent postocular teeth (Figs. 1C; 5C) NEW HYMENOSOMATID CRABS Zootaxa 3905 (1) © 2015 Magnolia Press · 101 (inconspicuous teeth in T. johnlucasi n. gen., n. sp.; Figs. 1A; 5B); (iv) antennules not visible dorsally when folded (Fig. 5C) (antennules visible dorsally in T. johnlucasi n. gen., n. sp.; Fig. 5B); (v) antennular basal article with acute, long anteroexternal tooth (Fig. 5C) (tooth still distinct but shorter in T. johnlucasi n. gen., n. sp.; (Fig. 5B); (vi) subhepatic region with two lobes, first larger and prominent, second tumescent (Fig. 5C) (both lobes pronounced and of about the same size in T. johnlucasi n. gen., n. sp.; Fig. 5B); (vii) mxp3 exopod nearly reaching the distal margin of the mxp3 merus (exopod reaching distal margin in T. johnlucasi n. gen., n. sp.; Fig. 5B); (viii) mxp3 dactylus and propodus of about the same length (dactylus longer than propodus in T. johnlucasi n. gen., n. sp.); (ix) teeth of cheliped cutting edges small and of different sizes (Fig. 8G) (teeth of same size in T. johnlucasi n. gen., n. sp.; Fig. 8D); and (x) three pterygostomian lobes (Fig. 5C) (with two pterygostomian lobes in T. johnlucasi n. gen., n. sp.; Fig. 5B). FIGURE 6. A–C, lateroanterior view of carapace. D, F, posterior view of the cephalothorax and abdomen. E, posterodorsal view of cephalothorax. A, Teramnonotus johnlucasi n. gen., n. sp., ovigerous female holotype cl. 2.8 mm, cw. 2.7 mm (AM P.12208). B, D, Teramnonotus monodi n. gen., n. sp., ovigerous female paratype cl. 4.3 mm, cw. 3.6 mm (MZUSP 10273). C, F, Elamena mathoei (Desmarest, 1823), female (ZRC 1994.4235). E, Trigonoplax unguiformis H. Milne Edwards, 1853, female (RMNH). Arrows indicate the absence of the rostral keel in A–B (distinct vertical keel on ventral surface of rostrum in C). Ab1, Ab2, first and second abdominal somites. CxP5, coxa of the pereopod 5. St 8, thoracic sternite 8 exposed. Scales: A–D, 1 mm. E–F, 2mm. Teramnonotus monodi and three other species of hymesomatid crabs are native to the Atlantic Ocean: Halicarcinus planatus (Fabricius, 1775), from Mar del Plata, Argentina southward to Tierra del Fuego, but also from localities outside the Atlantic (Peru, Chile, a number of subantarctic localities and Deception Island, Antarctica) (Melrose 1975: 34; Retamal 1981: 27; Boschi et al. 1992: 64; Boschi & Gavio 2005: 197; Boschi 2000: 84; Davie 2002: 246; Tavares 2004; Aronson et al. 2014); Hymenosoma orbiculare Desmarest, 1823, from Angola, southwest Africa, and South Africa (Monod 1956: 468; Manning & Holthuis 1981: 252); the record from Gabon is questionable (see Capart 1951: 62); and Elamena (Trigonoplax) gordonae Monod, 1956, from two West African localities between Guinea and Liberia. 102 · Zootaxa 3905 (1) © 2015 Magnolia Press TAVARES & SANTANA Two species of hymenosomatids have been recorded from the Western Hemisphere, Neorhynchoplax kempi (Chopra & Das, 1930), a freshwater species introduced into the Panama Canal (Abele 1972) and Elamena mexicana H. Milne Edwards, 1853 (as Elamene mexicana) from Mexico. Henri Milne Edwards (1853: 224) gave no illustration for E. mexicana and provided only an extremely short description: "Carapace plus étroite et à dents marginales plus marquées que dans les espèces précédentes.—Côtes du Mexique". Milne Edwards probably referred to the Pacific coast of Mexico. The presumed female holotype (by monotypy) of E. mexicana still exists as a dry specimen in the MNHN collections (Fig. 3B). Judging from this specimen, E. mexicana clearly does not belong in Elamena as now defined (see Ng & Chuang 1996). Because of the general shape of the carapace and appendages, which includes a trilobate rostrum, acute spines on the lateral carapace walls slightly below the carapace edge, approximately dorsal to base of P1 and P2, E. mexicana closely resembles Halicarcinus planatus, whose geographical distribution also includes the Eastern Pacific (Peru and Chile). Elamena mexicana H. Milne Edwards, 1853, is therefore synonymised herein with Halicarcinus planatus (Fabricius, 1775). The geographic distribution given by H. Milne Edwards (1853) for E. mexicana ("Côtes du Mexique") could have been mistaken, and the specimen was more likely originally obtained from Peru or Chile. Distribution. Teramnonotus monodi n. gen., n. sp. is known so far from Brazil (Ceará, Rio Grande do Norte, Sergipe, Bahia, and Rio de Janeiro, between about 03ºS and 22ºS). FIGURE 7. A, C. Female abdominal somites and pleotelson, dorsal view. A, B, Teramnonotus johnlucasi n. gen., n. sp., ovigerous female holotype cl. 2.8 mm, cw. 2.7 mm (AM P.12208). C, D, Teramnonotus monodi n. gen., n. sp., ovigerous female paratype cl. 4.3 mm, cw. 3.6 mm (MZUSP 10273). Ab1, Ab2, first and second abdominal somites. Ab3–Ab5, third, fourth and fifth abdominal somites fused together. PlT, pleotelson (sixth abdominal somite and telson fused together). Scales: A–D, 1 mm. NEW HYMENOSOMATID CRABS Zootaxa 3905 (1) © 2015 Magnolia Press · 103 FIGURE 8. A–C, Teramnonotus gordonae (Monod, 1956) n. gen., n. comb. A, ovigerous female holotype cl. 3.1 mm, cw. 2.6 mm (MNHN-B25803). B–C, ovigerous female paratype cl. 2.4 mm, cw. 2.2 mm (MNHN-B25802) (after Monod 1956: figs. 634 and 636). D–F, Teramnonotus johnlucasi n. gen., n. sp., ovigerous female paratype cl. 2.5 mm, cw. 2.4 mm (MZUSP 32909). G–I, Teramnonotus monodi n. gen., n. sp., ovigerous female paratype cl. 4.3 mm, cw. 3.6 mm (MZUSP 10273). A, D, G, right cheliped, lateral view. B, E, H, right P2, lateral view. C, F, I, left P5, lateral view. Scales: 1 mm. Acknowledgments We gratefully thank Alexandre Almeida (UESC), Charles Fransen (RMNH), Danièle Guinot (MNHN), Luis E. Bezerra (Universidade Federal do Ceará), Melissa Hewitt (WAM), Peter K. L. Ng (ZRC), and Stephen Keable (AM) for providing working space and/or access to their respective collections. Peter K. L. 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