Zoological Studies 46(4): 454-472 (2007) Establishment of a New Genus for Arete borradailei Coutière, 1903 and Athanas verrucosus Banner and Banner, 1960, with Redefinitions of Arete Stimpson, 1860 and Athanas Leach, 1814 (Crustacea: Decapoda: Alpheidae) Arthur Anker1,* and Ming-Shiou Jeng2 1Smithsonian 2Research Tropical Research Institute, Naos Unit 0948, APO AA 34002-0948, USA. E-mail:ankera@si.edu Center for Biodiversity, Academia Sinica, Taipei 115, Taiwan. E-mail:jengms@gate.sinica.edu.tw (Accepted October 5, 2006) Arthur Anker and Ming-Shiou Jeng (2007) Establishment of a new genus for Arete borradailei Coutière, 1903 and Athanas verrucosus Banner and Banner, 1960, with redefinitions of Arete Stimpson, 1860 and Athanas Leach, 1814 (Crustacea: Decapoda: Alpheidae). Zoological Studies 46(4): 454-472. Arete borradailei Coutière, 1903 and Athanas verrucosus Banner and Banner, 1960 are transferred to Rugathanas gen. nov., based on several unique features on the chelipeds, 3rd pereiopods, antennules, and mouthparts. The establishment of Rugathanas enables the redefinition of Athanas Leach, 1814 and Arete Stimpson, 1860, and a formal revalidation of Arete, formerly a synonym of Athanas. Two important features, the number of pereiopodal epipods and the number of carpal segments of the 2nd pereiopod, are variable within Rugathanas gen. nov., but may be used to distinguish Athanas from Arete. The distribution ranges of R. borradailei (Coutière, 1903) comb. nov. and R. verrucosus (Banner and Banner, 1960) comb. nov. are considerably extended based on recently collected material from the Ryukyu Is., Japan; Kenting, southern Taiwan; and Norfolk I., off eastern Australia. http://zoolstud.sinica.edu.tw/Journals/46.4/454.pdf Key words: Alpheidae, New genus, Athanas, Arete, Indo-Pacific. T armed with teeth, but without a fossa-tooth mechanism (e.g., Chace 1988) suggest that Athanas sensu lato is a highly heterogeneous and perhaps non-monophyletic assemblage. Banner and Banner (1960) questioned the validity of Arete Stimpson, 1860. They considered the features used by Coutière (1899 1903) to distinguish between Arete and Athanas invalid, e.g., the number of pereiopodal epipods (2 in Arete and 3 in Athanas) and the number of segments in the carpus of the 2nd pereiopod (4 in Arete and 5 in Athanas), and formally placed Arete in synonymy of Athanas. Although several workers (e.g., Holthuis 1993, Hayashi 1995, Nomura et al. 1996) continued to treat Arete as a valid genus, without discussing the synonymy proposed by Banner and Banner (1960), the majority of workers accepted he alpheid shrimp genus Athanas Leach, 1814 sensu lato (sensu Banner and Banner 1960 1973, Chace 1988) presently includes over 40 species in the eastern Atlantic and Indo-West Pacific. Most species are small (< 10 mm in total length) and inhabit coral reefs and rocky or muddy-sandy bottoms; some species live in symbiosis (commensalism) with other animals, such as thalassinideans, stomatopods, and echinoderms (Suzuki 1970, Banner and Banner 1973, Gherardi 1991, Froglia and Atkinson 1998, Anker et al. 2001, Hayashi 2002, Marin et al. 2005). Athanas species are mainly characterized by the presence of a triangular articulated plate on the 6th abdominal segment, and a well-developed, distally acute rostrum. The variously developed orbital teeth, and highly variable chelipeds, often *To whom correspondence and reprint requests should be addressed. 454 Anker and Jeng -- New alpheid genus with redefinition of Athanas and Arete this synonymy (e.g., Chace 1988). However, Banner and Banner (1982) recognized that 2 closely related species originally described as Arete borradailei Coutière, 1903 and Athanas verrucosus Banner and Banner, 1960 did not fit into either the generic diagnoses of Arete nor Athanas, and proposed reconsidering the validity of Arete, a suggestion they subsequently did not return to. Both Are. borradailei and Ath. verrucosus are rarely collected shrimps. The latter species was until now only known from the type locality, Enewetak Atoll, Marshall Is. (Banner and Banner 1960). This is partly due to the small size and cryptic lifestyles of these shrimps, which usually live in crevices of dead and living corals or in porous rocks overgrown with coralline algae. The only efficient method of collecting this type of cryptofauna is to carefully hammer these rocks apart to extract small animals from deep holes and , crevices. During the first author s (AA) field trip to Kenting National Park, southern Taiwan, 1 specimen of Ath. verrucosus was extracted by this method from dead coral rubble, collected from shallow water near the shore. We also examined several specimens of this species from Emily Bay, Norfolk I., off New South Wales, Australia, deposited in the Museum and Art Gallery of the Northern Territory, Darwin, Australia (NTM). The second author (MSJ) previously collected several specimens of Are. borradailei in Kenting. Through the courtesy of Keiichi Nomura (Kushimoto Marine Park Center, Kushimoto, Japan) we were able to study 4 specimens of Are. borradailei from the Ryukyu Archipelago. In addition, we examined 2 specimens of Are. borradailei from the southwestern Indian Ocean previously reported by Banner and Banner (1983), and numerous specimens of Athanas and Arete deposited in various museums. The morphological features of Are. borradailei and Ath. verrucosus clearly set these 2 species apart from all other species of Athanas or Arete. Therefore, a new genus, Rugathanas gen. nov., is herein established to accommodate these 2 species and formally resurrect Arete from synonymy of Athanas. Arete sensu stricto (sensu Stimspon 1860, not sensu Coutière 1903) and Athanas sensu stricto (sensu Coutière 1899 1903, not sensu Banner and Banner 1960) are redefined. MATERIAL AND METHODS The material used in this study is deposited in the collections of the Museum and Art Gallery of 455 the Northern Territory, Darwin, Australia (NTM); the Natural History Museum and Institute, Chiba, , Japan (CBM); the Muséum national d Histoire naturelle, Paris, France (MNHN); the Zoological Reference Collection, Raffles Museum of Biodiversity Research, National University of Singapore, Singapore (ZRC); Academia Sinica, Taipei, Taiwan (ASIZ); and the Naturhistorisches Museum in Wien, Vienna, Austria (NHMW). All drawings were made with the aid of a camera lucida. The carapace length (CL) and total length (TL) were measured in millimeters from the tip of the rostrum to the posterior margin of the carapace and telson, respectively. In cheliped descriptions, the terms equal, subequal, and unequal refer to size, while symmetrical, subsymmetrical, and asymmetrical refer to shape (general shape of the palm, armature of the fingers, etc.). Abbreviations used in the text include: Ar, arthrobranch; Ep, lobe-shaped epipod; Ex, exopod; Ma, mastigobranch (strap-like epipod); Mxp, maxilliped; P, pereiopod; Pd, podobranch; Pl, pleurobranch; r, rudimentary; Se, setobranch; +, present; -, absent. TAXONOMY Rugathanas gen. nov. Diagnosis: Body stout. Carapace smooth, not setose. Frontal margin with relatively short, somewhat ascendant rostrum, bearing acute tip, with or without subdistal tooth on ventral margin. Extracorneal teeth strong, acute, reaching or protruding beyond anterior margin of eyes; supra-corneal teeth usually well developed, acute; infra-corneal teeth reduced or absent. Orbital hoods absent; eyes exposed in dorsal and lateral views. Pterygostomial margin angular, protruding, not acute. Cardiac notch well developed. Second abdominal pleuron greatly expanded in females. Sixth abdominal somite with articulated plate at posteroventral angle; preanal plate rounded. Antennules with minute teeth on distal margin of first segment; ventromesial carina of 1st segment without tooth; lateral flagellum with rudimentary secondary ramus. Antenna with stout basicerite, with or without distolateral tooth; scaphocerite short, broadly oval-shaped. Mandible with palp, incisor process greatly expanded, distally with at least 15 min teeth. Maxillule with bilobed palp, ventral lobe with 1 seta, dorsal lobe unarmed. Labrum swollen, conspicuously protruding, visible in lateral view. Third maxilliped with penultimate segment as long as wide; ultimate segment with 456 Zoological Studies 46(4): 454-472 (2007) unarmed tip. First pereiopods (chelipeds) sexually dimorphic, carried flexed; coxa unarmed mesially; basis with rudimentary exopod; ischium robust, unarmed mesially; chelae without linea impressa on palm or cavity-tooth system on fingers. Male chelipeds subequal or unequal, subsymmetrical or asymmetrical (with major cheliped situated either on left or right side); ischium armed with spines on dorsal and usually ventral margin; merus swollen, distally depressed; carpus elongated, vaseshaped, excavated ventrally; chela enlarged, oval, compressed, slightly convex mesially, flattened laterally, dorsal and ventral margins rugose; linea impressa absent; fingers unarmed or armed with simple teeth. Female chelipeds equal, symmetrical; ischium usually unarmed or armed with several spines on dorsal margin; merus not or slightly inflated, flattened ventrally; carpus cup-shaped, more or less excavated ventrally; chela varying from oval-shaped to slender, with smooth or rugose margins, fingers unarmed. Carpus of 2nd pereiopod bearing 4 or 5 segments. Third pereiopod with unarmed ischium and merus; carpus with 1 distal spine on ventral margin; propodus with numerous ventral spines; dactylus simple, highly extensible dorsally. Fifth pereiopod with feebly developed propodal brush. Second pleopod with appendix interna and appendix masculina in males, with appendix interna only in females. Telson with 2 pairs of dorsal spines and 2 pairs of posterolateral spines; anal tubercles absent. Gill/exopod formula showing great variation in number of epipods (mastigobranchs/setobranchs), as summarized in following table. Pl Ar Pd Ep Ma Se Ex Mxp1 Mxp2 Mxp3 P1 P2 P3 P4 P5 + + + + -/+ + + -/+ -/+ r + -/+ -/+ r + -/+ - + - + - - - Type species: Rugathanas verrucosus (Banner and Banner, 1960) comb. nov. [Athanas verrucosus Banner and Banner 1960: 147]; type locality: Enewetak Atoll, Marshall Is. Other species included: Rugathanas borradailei (Coutière, 1903) comb. nov. [Arete borradailei Coutière 1903: 80]; type locality: Hulule Male Atoll, the Maldives. Etymology : Rugathanas is a combination of the Latin adjective ruga (rugged, uneven), referring to the characteristic rugosities on the margins of the male chelae, and Athanas (a Greek king), referring to its affinity with the genus Athanas. Distribution: Indo-West Pacific from the Red Sea and Madagascar to Japan, Australia, and Samoa. Remarks: Rugathanas may be separated from both Athanas and Arete by numerous features on the frontal margin of the carapace, antennules, antennae, chelipeds, and mandible. For instance, Rugathanas is characterized by the unique shape of the male (and sometimes female) chelipeds. The oval-shaped palm, compressed and flattened laterally and with rugose dorsal and ventral margins, and the vase-shaped, ventrally excavated carpus (Figs. 1a, 2l, m, 3c-e, 4a, 5b, c) are externally the most obvious features separating Rugathanas from Arete and Athanas. The configuration of the frontal margin of the carapace of Rugathanas is also characteristic of the genus (Figs. 2a, b, 3a, b, 5a), the most important features being the relatively short, ascendant rostrum, often with a small subdistal tooth on the ventral margin (vs. a long, straight rostrum, without a ventral tooth in Athanas and Arete); the presence of supraorbital teeth (absent in all Arete and most Athanas species, with the exceptions of Ath. djiboutensis Coutière, 1897 and Ath. areteformis Coutière, 1903, cf. Fig. 8b); the absence of infra-corneal teeth (usually absent in Arete, but often present in Athanas); and the presence of strong extra-corneal teeth reaching or sometimes protruding well beyond the anterior margin of the cornea (vs. usually not reaching far beyond this margin in Arete and Athanas, but with some exceptions, e.g., Ath. areteformis, cf. Fig. 8a). Rugathanas also differs from Arete and Athanas by the considerably expanded incisor process of the mandible (“psalistome”in Banner and Banner 1960), which is distally furnished with numerous (usually more than 15) minute teeth (Figs. 2g, h). In Arete and Athanas, the incisor process is not or only slightly expanded, distally bearing 5-12 teeth (A. Anker pers. obs.). One notable exception is again Ath. areteformis, in which the incisor process is also expanded and distally bears around 15 teeth (Figs. 8c, d). However, this species has very differently shaped chelipeds (Figs. 8e-j) and plainly belongs to the genus Athanas (see below). In species of Rugathanas, the labrum is greatly swollen (Fig. 2c), sometimes conspicuously protruding in lateral view (Fig. 3b), apparently to accommodate the expanded incisor process of the mandible. In con- Anker and Jeng -- New alpheid genus with redefinition of Athanas and Arete trast, the labrum of Ath. areteformis is only moderately swollen and not visible in lateral view. Other features distinguishing Rugathanas from Arete and Athanas are the stouter and (compared to other antennal parts) larger basicerite (Figs. 2a, 3b); the unarmed ventromesial carina on the 1st antennular segment (Fig. 2d) (vs. armed with strong tooth in all species of Arete and Athanas, partly visible in firure. 8a; see also Bruce 1990); the tip of the ultimate segment of the 3rd maxilliped without spines (vs. with spines in all Arete and most Athanas species); and the dorsally more extensible dactylus of the 3rd-5th pereiopods, which can be placed in an almost perpendicular position to the propodus (Figs. 2p, q). This unusually flexible articulation is strikingly similar to that of the unrelated alpheid genus Metalpheus Coutière, 1906, in which, however, the dactylus extension is facilitated by a small ventral sclerite (Banner 1959). Rugathanas also differs from Athanas by the strongly dentate margin of the 1st and 2nd segments of the antennular peduncle (Figs. 3a, 5a) (vs. straight or finely toothed in Athanas), and the much shorter penultimate segment of the 3rd maxilliped, which is about as long as wide in Rugathanas (Fig. 2k) (vs. distinctly longer than wide in Athanas). The number of carpal segments in the 2nd pereiopod varies from 4 in R. borradailei (Fig. 2o) to 5 in R. verrucosus (Fig. 4a). The strap-like epipods (mastigobranchs) are present on the 3rd maxilliped and 1st and 2nd pereiopods in R. verrucosus, but are absent in R. borradailei. Although these 2 important characters are variable within Rugathanas, they remain valuable for differentiating Rugathanas from Arete sensu stricto and Athanas sensu stricto (see below). Within the Alpheidae, an intrageneric variation in the number of carpal segments is not rare and also occurs in Synalpheus Bate, 1888 (4 to 5, see Banner and Banner 1975), Leptalpheus Williams, 1965 (4 to 5, see Anker et al. 2006c), Alpheopsis Coutière, 1896, and Prionalpheus Banner and Banner 1960 (3-5, see Banner and Banner 1971 1973, Bruce 1990b, Martínez-Iglesias and Carvacho 1991). Similarly, intrageneric variation in the number of pereiopodal epipods was also reported in at least 2 other genera, Alpheopsis (2-4, see Coutière 1899, Anker et al. 2005) and Athanas (see below). Rugathanas, Athanas and Arete are part of a larger generic complex (athanoid complex); other athanoid genera are Athanopsis Coutière, 1896 (5 species), Pseudathanas Bruce, 1983 (1 species), Acanthanas Anker, Poddoubtchenko and Jeng, 457 2006 (1 species), and Aretopsis De Man, 1910 (1 or 2 species). This complex forms a well-defined monophyletic clade (the clade AP of Anker et al. 2006a) within the family Alpheidae. According to Anker et al. (2006a) Athanopsis and Pseudathanas are sister genera, as are Arete and Aretopsis. Rugathanas (represented by R. borradailei) formed a sister group to Athanas. Athanas was found to be non-monophyletic; however, first, this genus was not adequately represented in this analysis, and second, the position of Ath. squillophilus as a sister to Athanopsis and Pseudathanas is rather questionable. The recently described Acanthanas was not included in Anker , et al. s (2006a) cladogram, but based on its morphological features (see Anker et al. 2006b), this genus appears to form a separate lineage somewhere within the clade Athanas - Rugathanas. Key to the genera of the athanoid generic complex and to the species of Rugathanas 1a. 1b. 2a. 2b. 3a. 3b. 4a. 4b. 5a. 5b. 6a. 6b. 7a. 7b. Rostrum distally rounded ................................................ 2 Rostrum distally acute ..................................................... 3 Chelipeds always carried extended, with dactylus in ventral position ......................................................... Aretopsis Chelipeds carried flexed (propodus fitting in a deep excavation on merus), dactylus in dorsal or dorsolateral position when chela extended ................................ Athanopsis Eyestalks with two conspicuously projecting acute spines ........................................................................ Acanthanas Eyestalks without spines ................................................. 4 Diaeresis of uropodal exopod with row of spines .................................................................... Pseudathanas Diaeresis of uropodal exopod without spines .................. 5 Male chelipeds with oval-rounded chelae, palm with rugose margins, carpus vase-shaped, ventrally excavated; labrum swollen, protruding; mandible with greatly expanded incisor process; supra-orbital teeth present ......................................................................Rugathanas 6 Male chelipeds with elongated or oval-shaped chelae, palm with non-rugose margins (sometimes with tubercles), carpus variably shaped; labrum neither swollen nor protruding; incisor process of mandible not or only slightly expanded; supra-orbital teeth present or absent ............ 7 Strap-like epipods absent; 2nd pereiopod with 4-segmented carpus; male chelipeds asymmetrical; female chelipeds only slightly enlarged ......................... R. borradailei Strap-like epipods present on Mxp3 and P1-2; 2nd pereiopod with 5-segmented carpus; male chelipeds subsymmetrical; female chelipeds greatly enlarged ....................... ..................................................................... R. verrucosus Strap-like epipods present on Mxp3 and P1-2; 2nd pereiopod with 4-segmented carpus; cheliped ischium with strong process mesially; supra-orbital teeth absent ................................................................................... Arete Strap-like epipods present on Mxp3 and P1-3 (exceptionally on P1-2 or P1-4); 2nd pereiopod with 5-segmented carpus; cheliped ischium without a strong process mesially; supra-orbital teeth absent or present .............. Athanas 458 Zoological Studies 46(4): 454-472 (2007) Rugathanas borradailei (Coutière, 1903) comb. nov. (Figs. 1-3, 6a) Arete borradailei Coutière 1903: 80, figs. 19-24; Coutière 1905: 861, fig. 133; Jacquotte 1964: 180; Banner and Banner 1966a: 152; Banner and Banner 1983: 73; Chace 1988: 61. Arete ghardaqensis Ramadan 1936: 15, pl. 1, figs. 2-8. Athanas ghardaquensis Banner and Banner 1981: 42; Banner and Banner 1983: 73. Athanas polynesia Banner and Banner 1966a: 152, fig. 7; Banner and Banner 1982: 302, fig. 93; Banner and Banner 1983: 73. Material examined: CBM-ZC 8469, 1 male (CL 2.6, TL 8.3), Japan, S Ryukyu Arch., Yaeyama Group, Kuro-shima, Nakamoto, dead coral crevices, intertidal, coll. K. Nomura, 24 Sep. 1987 [YMP-672c]; MNHN-Na 15778, 2 ovigerous females (CL 2.5, TL 8.5; CL 2.7, TL 8.5), Japan, S Ryukyu Arch., Yaeyama Group, Kuro-shima, Iko, dead coral crevices, depth 2 m, coll. K. Nomura, (a) 1 mm a (b) 1 mm b Fig. 1. Rugathanas borradailei (Coutière, 1903) comb. nov.: (a) male from Kenting, southern Taiwan (ASM); (b) ovigerous female from the Ryukyu Is., Japan (MNHN). 459 Anker and Jeng -- New alpheid genus with redefinition of Athanas and Arete 29 Aug. 1986 [YMP-163]; MNHN-Na 15779, 1 ovigerous female (CL 2.8, TL 8.1) Japan, S Ryukyu Arch., Yaeyama Group, Kuro-shima, Urabishi, dead coral crevices, depth 3 m, coll. K. Nomura, 17 Jul. 1999 [YMP-2395]; ASIZ 72761, 1 male (CL 5.3, TL 15.3), 1 female (CL, TL not mea- 0.5 mm a-r (b) (c) (a) (l) (k) (n) (d) (m) (j) (p) (q) (e) (i) (f) (g) (h) (o) (r) (s) (r) 1 mm s Fig. 2. Rugathanas borradailei (Coutière, 1903) comb. nov., ovigerous female (a-k, o-s) and male (l-n) from southwestern Madagascar (MNHN): (a) frontal region, lateral view; (b) frontal margin of carapace, dorsal view; (c) frontal region, ventral view showing bases of antennules and antennae, labrum, and position of mouthparts; (d) antennule, lateral view; (e) same, peduncle, dorsal view; (f) scaphocerite; (g, h) mandible from 2 different aspects; (i) maxillule; (j) 2nd maxilliped; (k) 3rd maxilliped; (l) left cheliped, lateral view; (m) same, chela, mesial view; (n) same, coxa and basis with exopod; (o) 2nd pereiopod; (p) 3rd pereiopod; (q) 5th pereiopod, distal carpus, propodus, and dactylus; (r) uropod; (s) abdomen and cephalothorax (frontal appendages and pereiopods not drawn). 460 Zoological Studies 46(4): 454-472 (2007) sured), S Taiwan, Kenting National Park, coll. M.S. Jeng; MNHN-Na 4604, 1 male (CL 2.7, TL 8.8), La Réunion, reef crevices, coll. M. Peyrot-Clausade; MNHN-Na 4610, 1 ovigerous female (CL 2.8, TL 8.8), Toliara (Tuléar), Madagascar, from reef crevices, coll. M. Peyrot-Clausade. Diagnosis: Male chelipeds enlarged, varying from subequal and subsymmetrical in immature individuals to unequal and asymmetrical in adults; ischium with spines; palm expanded, ventral margin rugose, without papillae; fingers unarmed or with small teeth. Female chelipeds feebly enlarged, equal, symmetrical; ischium usually without spines, palm not expanded. Second pereiopod with 4-segmented carpus. Strap-like epipods (mastigobranchs) absent. For complete description see Banner and Banner (1982 1983). Size: Small-sized shrimp; largest specimen examined, a male from Taiwan (Fig. 1a), measured 15.3 mm TL; the average TL of Australian specimens (reported as Ath. polynesia) being 8 mm (Banner and Banner 1982). Colour : Semitransparent with greenish or olive tinge and feebly marked yellowish-orange bands on abdomen tail fan and chelae; pale-brown inner organs visible due to transparence; eggs green (Fig. 6a). Habitat: Crevice-dweller on coral reefs. Most specimens from the western Indian Ocean were found in cavities of coral rocks overgrown with coralline algae, in groove and spur system on the outer reef edge to 25 m deep; in sections of reef flat and a pinnacle on the inner slope; in dead and overgrown coral heads, as well as among branches of living corals, e.g., Acropora variabilis (Klunzinger, 1879) and Stylophora mordax (Dana, 1846) (Banner and Banner 1983). Jacquotte (1964) found several specimens in boring holes of the sea urchin Stomopneustes variolaris (Lamarck, 1816). The Australian specimens (reported as Ath. polynesia) inhabited areas covered with encrusting coralline algae in 2-7 m (Banner and Banner 1982). Distribution: Maldives: Hulule Malé Atoll (type (a) (b) (d) 1 mm b-e (f) (e) (c) 0.2 mm f 0.5 mm a Fig. 3. Rugathanas borradailei (Coutière, 1903) comb. nov., male from Samoa reported as Athanas polynesia Banner and Banner, 1966 (a-e), and female from Kenting, southern Taiwan (ASIZ) (f): (a) frontal region; (b) frontal region with protruding labrum and 3rd maxilliped, lateral view; (c) major cheliped, lateral view; (d) same, carpus and chela, mesial view; (e) minor cheliped, mesial view (a-e, adapted from Banner and Banner 1966a). Anker and Jeng -- New alpheid genus with redefinition of Athanas and Arete locality of R. borradailei) (Coutière 1903 1905); Egypt: Hurghada (type locality of Are. ghardaqensis) (Ramadan 1936); Gulf of Aqaba (Banner and Banner 1981); Somalia; La Réunion; Mauritius (Banner and Banner 1983); Madagascar (Jacquotte 1964, Banner and Banner 1983); Philippines: Negros I. (Chace 1988); Australia: Lizard I. off Queensland (type locality of Ath. polynesia) (Banner and Banner 1982); Samoa (Banner and Banner 1966a); southern Taiwan: Kenting; southern Japan: Ryukyu Is. (present study). Remarks: The taxonomic statuses of Are. ghardaqensis Ramadan, 1936 and Ath. polynesia Banner and Banner, 1966 were reviewed by Banner and Banner (1983), who placed both nominal species in synonymy with Ath. borradailei (now R. borradailei). Those authors concluded that the chelipeds of R. borradailei are variable and sexually dimorphic: slender, symmetrical, with a few or no spines on the ischium in females (Coutière 1905: 461 fig. 135c, Banner and Banner 1982: fig. 93k); and robust, subsymmetrical or asymmetrical, with usually strong spines on ischium in males (Banner and Banner 1966a: figs. 7e-g, 1982: figs. 93c-e). However, other features of R. borradailei, such as the development of the orbital teeth, the shape of the rostrum, and the armature of the antennal basicerite, are also variable (Ramadan 1936, Banner and Banner 1983). Therefore, R. borradailei cannot be excluded from either being a variable species or a species complex. Rugathanas verrucosus (Banner and Banner, 1960) comb. nov. (Figs. 4, 5, 6b) Athanas verrucosus Banner and Banner 1960: 147, fig. 4; Banner and Banner 1968: 270; Banner and Banner 1983: 75. Material examined: NTM Cr001663, 2 males (a) 1 mm (b) Fig. 4. Rugathanas verrucosus (Banner and Banner, 1960) comb. nov.: (a) male from Kenting, southern Taiwan (ASIZ); (b) ovigerous female (left) and male (right) from Norfolk I. (NTM) showing sexual dimorphism in the development of the chelipeds. 462 Zoological Studies 46(4): 454-472 (2007) (CL 4.5, TL 12.7; CL 2.7, TL 7.4), 1 ovigerous female (CL 5.4, TL 15.0), Point Hunter, Emily Bay, Norfolk I., off New South Wales, Australia, 29 03.8'S 167 57.3'E, depth 0-0.25 m, Sta. HL 84-21, coll. H. Larson, det. A.J. Bruce, 18 Apr. 1984; NTM Cr001663, 2 males (CL 2.3, TL 7.2; CL 4.2, TL 11.8), 4 ovigerous females (smallest, CL 3.8, TL 10.1; largest, CL 4.6, TL 12.3), Point Hunter, Emily Bay, Norfolk I., off New South Wales, Australia, 29 03.8'S 167 57.3'E, depth 0-0.25 m, Sta. HL 84-20, coll. H. Larson, det. A.J. Bruce, 17 Apr. 2004; ASIZ 73699, 1 male (CL 3.4, TL 10.1), S Taiwan, Kenting National Park, from coral rocks, depth 0.5-2 m, coll. A. Anker, 1 Oct. 2004. Diagnosis: Male chelipeds subequal, subsymmetrical, ischium with spines; palm expanded, ventral margin rugose, distoventral portion of palm, pollex, and proximal portion of dactylus with papillae (i.e., small subacute or rounded tubercles); fingers armed with teeth. Female chelipeds enlarged, slightly less stout than male chelipeds, equal, symmetrical; ischium with spines; palm less expanded than in male, with papillae. Second pereiopod with 5-segmented carpus. Strap-like epipods (mastigobranchs) present on 3rd maxilliped and 1st and 2nd pereiopods. For complete description see Banner and Banner (1960). Size: Small-sized shrimp; type specimen measuring 3.9 mm CL and 11 mm TL (Banner and Banner 1960); the Kenting male specimen only slightly smaller, at 3.4 mm CL and 10.1 mm TL; the largest examined specimen, an ovigerous female from Norfolk I., at CL 5.4 mm and TL 15.0 mm. Colour : Semitransparent, with purple-greenish bands on abdomen; fingers of chela with golden-yellow tinge; setae fringing uropods and telson golden-brown (Fig. 6b). Habitat : In Enewetak, the habitat of R. verrucosus was described as“coralline ridge, ocean (windward) reef”and“coralline ridge, and ends of surge channels, exposed about 0.3 meters above 0.0 tides; ridge dense and hard, very irregular and without the usual smooth algal surface ... from superficial burrows, roofed over by calcareous algal sheets”(Banner and Banner 1960 1968). The single Kenting specimen was found in a crevice during process of breaking apart coral rocks and rubble collected in shallow water (0.5-2 ° ° ° ° (b) (a) (c) 1 mm a-c (e) (d) (f) 1 mm d-f Fig. 5. Rugathanas verrucosus (Banner and Banner, 1960) comb. nov., male from Kenting, southern Taiwan (ASIZ) (a-c); ovigerous female from Norfolk I. (NTM) (d-f): (a) frontal region; (b) male minor chela, lateral view; (c) male major chela, lateral view; (d) female cheliped, lateral view; (e) same, mesial view; (f) abnormal telson. Anker and Jeng -- New alpheid genus with redefinition of Athanas and Arete (a) (b) (c) (d) (e) (f) (g) (h) 463 Fig. 6. Habitus of some species of Rugathanas gen. nov. (a, b), Arete Stimpson, 1860 (c, d), and Athanas Leach, 1814 (e-h): (a) Rugathanas borradailei (Coutière, 1903) comb. nov., ovigerous female from Kenting, southern Taiwan; (b) Rugathanas verrucosus (Banner and Banner, 1960) comb. nov., male from Kenting, southern Taiwan; (c) Arete cf. dorsalis Stimpson, 1860, specimen from southern Japan; (d) Arete cf. dorsalis, shrimp in situ among spines of a sea urchin, Lord Howe I., off eastern Australia; (e) Athanas areteformis Coutière, 1903 (Ath. nitescens group), male from southern Japan; (f) Athanas nitescens (Leach, 1814) (Ath. nitescens group), male from Banyuls-sur-Mer, Mediterranean coast of France; (g) Athanas japonicus Kubo, 1936 (Ath. dimorphus group), ovigerous female from southern Japan; (h) Athanas amazone Holthuis, 1951 (Ath. dimorphus group), male from Banyuls-sur-Mer, Mediterranean coast of France (photograph credits: a, M.S. Jeng; b, A. Anker, c, e, g, M. Itoh; d, N. Coleman; f, h, P. Lecomte leg. P. Noël). 464 Zoological Studies 46(4): 454-472 (2007) m) on a near-shore, exposed reef. The Norfolk I. specimens were collected on a rocky reef, at depths reaching from low tide mark to 0.25 m. Distribution: Marshall Is.: Enewetak Atoll (type locality, Banner and Banner 1960 1968); Australia: Norfolk I.; southern Taiwan: Kenting (present study). Remarks: In R. verrucosus, the male chelipeds can be classified as subsymmetrical (Figs. 5b, c), and the female chelipeds are generally only slightly weaker than the male chelipeds (Figs. 3, 5d, e, see also Figs. 3). In contrast to this, in R. borradailei, the heterochelate condition (asymmetry) in males (Fig. 3c, e) and the sexual dimorphism (chelipeds significantly larger in males, see Figs. 1a, b, 3c-f) appear to be more pronounced. Ovigerous females of R. verrucosus from Norfolk I. carried numerous (several dozen) relatively smallsized eggs. One of the ovigerous females from Norfolk I. had an aberrant number of spines on the telson: 4 dorsal spines + 1 posterolateral spine on 1 side, and 2 dorsal and 2 posterolateral spines (typical condition) on the other side (Fig. 5f). Arete Stimpson, 1860 sensu stricto Arete Stimpson 1860: 32. Diagnosis: Body relatively stout. Carapace smooth, not setose. Frontal margin with long, straight rostrum, bearing acute tip, without subdistal tooth on ventral margin. Extra-corneal teeth acute, not protruding beyond anterior margin of eye; supra-corneal teeth absent or feebly developed; infra-corneal teeth absent. Orbital hoods absent; eyes exposed in dorsal and lateral views. Pterygostomial margin usually rounded or angular, sometimes subacute. Cardiac notch well-developed. Sixth abdominal segment with articulated plate at posteroventral angle; preanal plate acutely produced. Antennules with finely serrated distal margin of 1st segment; ventromesial carina of 1st segment with strong tooth; lateral flagellum with (a) (i) (b) (g) (h) (e) 1 mm g (d) (c) (f) 1 mm a-f, h, i Fig. 7. Arete cf. indicus Coutière, 1903, 2 males from Sesoko Beach, Okinawa (ZRC) (a-f, h, i) and male from Sri Lanka (NHMW) (g): (a) frontal region, lateral view; (b) same, dorsal view; (c) left cheliped, lateral view; (d) same, coxa to carpus, mesial view; (e) same, chela and carpus, lateral view; (f) fingers of chela of a different specimen, lateral view; (g) chela fingers of a larger specimen, mesial view; (h) 2nd pereiopod; (i) 3rd pereiopod. 465 Anker and Jeng -- New alpheid genus with redefinition of Athanas and Arete well-developed secondary ramus. Antenna with normal, not particularly stout basicerite, distolateral tooth always present; scaphocerite broadly oval or rounded. Mandible with palp, incisor process slightly expanded, distally with no more than 12 small teeth. Maxillule with bilobed palp, both ventral and dorsal lobes with setae. Labrum not swollen or protruding. Third maxilliped with distally projecting dorsal margin of antepenultimate segment; penultimate segment rectangular, about as long as or slightly longer than wide; ultimate segment with short spines on tip. First pereiopods (chelipeds) feebly sexually dimorphic, equal or subequal, usually subsymmetrical, sometimes asymmetrical (major cheliped situated on either side), stout, carried extended with dactylus in lateral or ventrolateral position; coxa unarmed mesially; ischium usually with strong tooth mesially and with lobes or teeth furnished with spines on dorsal margin; basis with rudimentary exopod; merus robust, flattened ventrally; carpus swollen, cup-shaped, embracing proximal portion of palm distally, ventral side somewhat flattened, not deeply excavated; chela smooth, without sculpturing, compressed; palm sometimes with acute distal projection on mesial side proximal to articulation with dactylus, without or with feebly marked linea impressa; fingers armed with teeth, sometimes interrupted by large hiatus, without cavity-tooth system. Carpus of 2nd pereiopod bearing 4 segments. Third pereiopod with ischium unarmed or bearing 1 spine on ventrolateral margin; merus without spines, sometimes with distoventral margin ending in angular or acute projection; carpus unarmed; propodus armed with spines; dactylus stout, biunguiculate. Fifth pereiopod with poorly developed propodal brush (at most 3 rows of setae). Second pleopod with appendix interna and appendix masculina in males and sometimes in females. Telson with 2 pairs of dorsal spines and 2 pairs of pos- (b) (a) (g) (e) 1 mm a, b, e, f, h-m (j) (i) (h) (f) (d) (k) (c) (l) 0.5 mm c-d, g (m) Fig. 8. Athanas areteformis Coutière, 1903, male from Papua New Guinea (MNHN): (a) frontal region, lateral view; (b) frontal margin of carapace, dorsal view; (c, d) mandible from 2 different aspects; (e) left (major) cheliped, mesial view; (f) same, chela and carpus, mesial view; (g) same, coxa and basis with exopod; (h) right (minor) cheliped, mesial view; (i) same, chela and carpus, mesial view; (j) same, chela fingers, lateral view; (k) 2nd pereiopod; (l) 3rd pereiopod; (m) 5th pereiopod. 466 Zoological Studies 46(4): 454-472 (2007) terolateral spines; anal tubercles lacking. Gill/exopod formula summarized in following table. Pl Ar Pd Ma Se Ex Mxp1 Mxp2 Mxp3 P1 P2 P3 P4 P5 + + + + + + + r + + + r + + - + - + - - - 1942: 82 (for synonymy see Banner and Banner 1960 1973, Miya and Miyake 1968). Arete sp. nov. 1 - Athanas indicus - Suzuki 1970: 5 (not Coutière 1903). Arete sp. nov. 2 - Athanas dorsalis - Suzuki 1970: 12 (not Stimpson 1860). Distribution: Indo-West Pacific: Red Sea, Djibouti, Madagascar, La Réunion, Maldives, Seychelles, Sri Lanka, Japan (including Ryukyu Is.), China, Taiwan, Philippines, Indonesia, Australia (including Lord Howe I.), Micronesia, Melanesia, French Polynesia. Remarks: Arete, as redefined here, may be easily separated from Rugathanas by numerous morphological features (see above), and from Athanas by the stouter chelipeds (Fig. 7c, e) with large, oval-shaped chelae (vs. slenderer in Athanas, Figs. 8e, h, 9); the ischium of the chelipeds bearing a strong mesial process (Fig. 7d) (unarmed in Athanas); the carpus of the 2nd pereiopods bearing 4 segments (Fig. 7h) (vs. 5 in Athanas, Fig. 8k); the distoventral margin of the merus of the 3rd pereiopod being angular or subacute (Fig. 7i) (vs. rounded in Athanas, Fig. 8l); the dactylus of the 3rd to 5th pereiopods stout (Fig. 7i) (vs. usually slender in Athanas, Fig. 8l); the straplike epipods or mastigobranchs present on 1st and 2nd pereiopods (vs. on 1st to 3rd or 4th pereiopods in Athanas); and the preanal plate of the 6th abdominal somite being acutely produced towards the telson (vs. rounded, not produced in Athanas). Furthermore, the antepenultimate seg- Type species: Arete dorsalis Stimpson 1860: 32; type locality: Hong Kong. Synonyms: Athanas mascarenicus Richters 1880: 164; Arete dorsalis pacificus Coutière 1903: 84; Arete maruteensis Coutière 1905: 866; Arete maruteensis salibabuensis de Man 1910: 313; ? Arete kominatoensis - Suzuki 1970: 2 (not Kubo 1942). Other species included: Three described and at least 2 undescribed species (K. Nomura, pers. obs.; A. Anker pers. obs.). Arete acanthocarpus (Miya and Miyake 1968) [Athanas acanthocarpus Miya and Miyake 1968: 157]; type locality: Okinawa, Japan. Arete amboinensis de Man 1910: 25; type locality: Ambon, Indonesia. Specific status needs confirmation. Arete indicus Coutière 1903: 84; type locality: not designated, type specimens from Djibouti and Hulule Male Atoll, Maldives. Synonyms: Arete iphianassa de Man 1910: 312; Arete intermedius Yu 1931: 513; ? Athanas kominatoensis Kubo (a) (b) 1 mm b 1 mm a (c) (e) (d) 1 mm c-f (f) Fig. 9. Chelipeds of 3 species from the Athanas nitescens group: (a) Athanas grimaldii Coutière, 1911, male from Senegal (MNHN), major cheliped, carpus, and chela, mesial view; (b) Athanas granti Coutière, 1908 from southern Australia, type (MNHN), major cheliped, carpus, and chela, mesial view; (c-f) Athanas cf. nitescens (Leach, 1814) from Senegal (MNHN), (c) male cheliped, lateral view; (d) same, chela and carpus, mesial view; (e) female cheliped, lateral view; (f) same, chela and carpus. Anker and Jeng -- New alpheid genus with redefinition of Athanas and Arete ment of the 3rd maxilliped usually projects distodorsally in Arete (see Bruce 1990a), but not in Rugathanas or Athanas. The propodal brush on the 5th pereiopod is poorly developed in both Arete (cf. Bruce, 1990a) and Rugathanas (Fig. 2q), but is well-developed in Athanas (Fig. 8m). In Arete, both males and females bear enlarged, stout chelipeds, with broadened chelae that are carried extended forward, with the dactylus situated in lateral or ventrolateral position (Fig. 6c, d). The chelipeds of Athanas are highly variable in shape (cf. Figs. 6e-h), and may be carried either extended (Ath. nitescens (Leach, 1814) species group, cf. Figs. 6e, f), or folded (Ath. dimorphus Ortmann, 1894 species group, cf. Figs. 6g, h). The Ath. nitescens group may have some affinities with Arete, but in none of the species of this group, the chelae are as broad as in Arete (compare Figs. 6c, d, 7c-g and Figs. 6e, f, 8e-h, 9), which actually look like“mini-lobsters”(Figs. 6c, d). All Arete species are characterized by a 4segmented carpus in the 2nd pereiopod and the presence of strap-like epipods on the 1st and 2nd pereiopods. In the vast majority of Athanas spp., the carpus of the 2nd pereiopod is composed of 5 , segments (A. Anker pers. obs.). Kemp s (1915) reported specimens of Ath. polymorphus Kemp, 1915 from Chilka Lake (India) that had either 4 or 5 segments, but in 7 examined specimens of Ath. polymorphus from Singapore, the carpus had 5 segments (Anker 2003). In Ath. squillophilus Hayashi, 2002, the number of carpal segments varies from 5 to 6 (Hayashi 2002). One of us (AA) also examined a specimen of Ath. dimorphus with 6 instead of the typical 5 carpal segments. In most species of Athanas, the strap-like epipods are present on the 1st-3rd pereiopods, although Ath. ohsimai Yokoya, 1936 appears to have a complete epipod set, i.e., in this species, the strap-like epipods are present on the 1st-4th pereiopods (Yokoya 1936). Banner and Banner (1960) reported an infraspecific variation in the number of pereiopodal epipods in a population of Ath. djiboutensis Coutière, 1897, with most specimens having epipods on the 1st and 2nd pereiopods, and only some on the 1st-3rd pereiopods. Despite all the aforementioned exceptions, the combination of the cheliped features (e.g., chelae broad, oval-shaped in Arete, more or less elongated in Athanas); the number of carpal segments in the 2nd pereiopod (4 in Arete, 5 in Athanas, exceptionally 4 or 6); and the number of pereiopodal epipods (on P1 and P2 in Arete, on 467 P1-3 in Athanas, exceptionally on P1 and P2 or P1-4) enables Arete to be distinguished from Athanas. Arete is also ecologically distinct from Rugathanas and Athanas. All Arete species are obligate symbionts of sea urchins (Fig. 6d), mostly species of Echinometra, Anthocidaris, Heterocentrotus, Heliocidaris, Echinothrix, Tripneustes, and Centrostephanus (Suzuki 1970, Banner and Banner 1973), and at least 2 species exhibit protandric hermaphroditism (Suzuki 1970, Gherardi and Calloni 1993), which explains the presence of an appendix masculina in some ovigerous specimens. Within Athanas, only Ath. granti Coutière, 1908 occasionally associates with sea urchins of the genus Centrostephanus (Banner and Banner 1973). Remarkably, this species shows some resemblance to Arete, but differs in many other respects and clearly belongs to Athanas (see below). Protandric hermaphroditism has yet to be documented in Athanas, although it is suspected in Ath. ornithorhynchus Banner and Banner, 1973, a remarkable species associated with brittle stars (Marin et al. 2005). As already mentioned, Arete requires revision at the species level. The taxonomic status of de , Man s“Arete maruteensis salibabuensis”placed in synonymy of Are. dorsalis, as well as the taxonomic statuses of Are. iphianassa, Are. intermedius, and Are. kominatoensis (originally described as Ath. kominatoensis), placed in synonymy with Are. indicus (Banner and Banner 1973), should be reconsidered. All these forms should be contrasted to 2 species misidentified as Are. indicus and Are. dorsalis by Suzuki (1970), and which probably represent undescribed species (K. Nomura, pers. comm.). Therefore, many of the previous identifications of Arete species (e.g., Banner and Banner 1960 1973, Miya and Miyake 1968, Suzuki 1970, Bruce 1989 1990a), including , Bruce s (1990a) redescription of Are. dorsalis, are questionable. As such a key to the genus Arete is considered premature. Athanas Leach, 1814 sensu stricto Athanas Leach 1814: 401. Diagnosis: Body usually slender. Carapace usually smooth, sometimes setose. Frontal margin with long, straight, slightly ascendant, or descendant rostrum, with acute tip, without subdistal tooth on ventral margin, rarely with teeth on dorsal margin. Extra-corneal teeth present, acute, sometimes protruding beyond anterior margin of eye; 468 Zoological Studies 46(4): 454-472 (2007) supra- and infra-corneal teeth absent or present. Orbital hoods absent; eyes exposed in dorsal and lateral views. Pterygostomial margin usually rounded, rarely sharply protruding or with small acute tooth. Cardiac notch well-developed. Sixth abdominal segment with articulated plate at posteroventral angle; preanal plate rounded. Antennules with straight, not serrated distal margin of 1st segment; ventromesial carina of 1st segment with strong tooth; lateral flagellum with welldeveloped secondary ramus. Antenna with normal, not particularly stout basicerite, distolateral tooth present; scaphocerite variably oval to subrectangular, usually more or less elongate. Mandible with palp, incisor process usually not expanded, distally with 5-8 medium-sized teeth, rarely expande, with 12-15 min teeth. Maxillule with bilobed palp, both ventral and dorsal lobes with setae. Labrum not swollen or protruding. Third maxilliped without distal tooth on dorsal margin of antepenultimate segment; penultimate segment elongate, distinctly longer than wide; ultimate segment often with spines on tip. First pereiopods (chelipeds) highly variable in shape, size, and degree of asymmetry, often sexually dimorphic or polymorphic, carried folded or extended with dactylus in lateral position; coxa usually with apressed subtriangular tooth mesially; basis with rudimentary exopod; ischium unarmed mesially, often with spines on dorsal and ventral margins; chelae without linea impressa on palm and without fossa-plunger on fingers. Male chelipeds usually enlarged and/or elongate, equal, subequal, or unequal, symmetrical, subsymmetrical, or asymmetrical (with major cheliped situated either on left or right side); ischium sometimes with lobes furnished with spines on dorsal margin; merus usually stout, depressed or excavated ventrally; carpus elongate or cup-shaped, sometimes flattened ventrally; chela highly variable, usually conspicuously swollen or elongate, sometimes with tubercles along ventral margin; fingers usually armed with teeth. Female chelipeds variable from slender, slightly elongate, to greatly enlarged, equal, subequal, or unequal, symmetrical, subsymmetrical, or asymmetrical (with major cheliped situated on left or right side); merus variable, slender to stout, flattened or excavated ventrally; carpus elongate or cup-shaped, sometimes flattened ventrally; chelae variable from slender to oval or elongate, sometimes with tubercles along ventral margin; fingers unarmed or armed with teeth. Carpus of 2nd pereiopod with 5 segments, exceptionally 4 or 6. Third pereiopod with ischium unarmed or bearing 1 spine on ventrolateral margin; merus unarmed, distoventral margin never acute or angular; carpus unarmed; propodus usually with ventral spines, rarely with spine-like setae; dactylus more or less slender, simple or biunguiculate. Fifth pereiopod with well-developed propodal brush. Second pleopod with appendix interna and appendix masculina in males, appendix interna in females, exceptionally with appendix masculina in ovigerous specimens. Telson with 2 pairs of dorsal spines and 2 pairs of posterolateral spines; anal tubercles absent. Gill formula with somewhat variable number of epipods, as summarized in following table (untypical condition indicated in parentheses). Pl Ar Pd Ma Se Ex Mxp1 Mxp2 Mxp3 P1 P2 P3 P4 P5 + + + + + + + r + + + r + +(-) + - + -(+) +(-) - + -(+) - Type species: Athanas nitescens Leach, 1814 [Palaemon nitescens Leach 1814: 401]; type locality: England. Most important synonyms: Palaemon laevirhincus Risso 1816: 108; Athanas var. laevirhincus (Risso, 1816), Holthuis and Gottlieb 1958: 32; Arete diocletiana Heller 1862: 404; Athanas transitans Czerniawsky 1884: 25; Athanas nitescens var. suchumicae Czerniawsky 1884: 24; Athanas nitescens forma rotundicauda Czerniawsky 1884: 24; Athanas transitans var. longispina Czerniawsky 1884: 25; Athanas transitans var. pontica Czerniawsky 1884: 26; Athanas veloculus Bate 1888: 529. The taxonomic statuses of Ath. laevirhincus and Ath. veloculus requires confirmation (see below). Other species included: Athanas includes all the remaining species described mostly under the generic name Athanas. Athanas amazone Holthuis 1951: 111; type locality: Nigeria. Athanas anatidactylus Anker and Marin 2007: 163; type locality: Vietnam. Athanas areteformis Coutière 1903: 79; type locality: Hulule Male Atoll, Maldives. Synonyms: Athanas naifaroensis Coutière 1903: 77; Athanas erythraeus Ramadan 1936: 13; Athanas dubius Banner 1956: 322 (see Banner and Banner 1973). The taxonomic statuses of Ath. naifaroensis, Ath. erythraeus and Ath. dubius require confirmation. Athanas crosslandi Tattersall 1921: 372; type Anker and Jeng -- New alpheid genus with redefinition of Athanas and Arete locality: Khor Dongonab, Sudanese Red Sea. Athanas dentirostris Anker, Jeng and Chan 2001: 1049; type locality: Haiphong, Vietnam. Athanas dimorphus Ortmann 1894: 12; type locality: Dar es Salaam, Tanzania. Synonyms: Alpheus monoceros (Heller 1862: 274, nomen dubium); Athanas setoensis Kubo 1951: 265; Athanas dimorphus seedang Banner and Banner 1966b: 28 (see Banner and Banner 1960 1973). The taxonomic statuses of Ath. setoensis and Ath. dimorphus seedang require confirmation. Athanas djiboutensis Coutière 1897: 233; type locality: Djibouti. Synonym: Athanas sulcatipes Borradaile 1898: 1011 (see Banner and Banner 1960 1973). Athanas esakii Kubo 1940: 93; type locality: “ Kusaie”, possibly Kosrae, Caroline Is., Micronesia. Athanas gracilipes Banner and Banner 1978: 234; type locality: South China Sea. Athanas gracilis Boone 1935: 122; type locality: Raiatea, Society Is., French Polynesia. Athanas granti Coutière 1908: 192; type locality: Adelaide, South Australia. Athanas grimaldii Coutière 1911: 1; type locality: not designated, type specimens from Bay of Biscay, France and Cape Verde Is. Athanas haswelli Coutière 1908: 192 (amended from the original spelling Ath. hasswelli by Banner and Banner 1973); type locality: Adelaide, South Australia. Athanas hongkongensis Bruce 1990b: 624; type locality: Hong Kong, China. Athanas japonicus Kubo 1936: 43; type locality: Yamaguchi, Japan. Synonyms: Athanas lamellifer Kubo 1940: 102 (see Miya and Miyake 1968). The taxonomic status of Ath. lamellifer needs clarification. Athanas jedanensis de Man 1910: 313; type locality: Pulau Djedan, eastern Indonesia. Athanas locincertus Banner and Banner 1973: 311; type locality:“Panchoran Buoy”, possibly Western Australia. Athanas marshallensis Chace 1955: 17; type locality: Marshall Is. Athanas minikoensis Coutière 1903: 76; type locality: Minikoy Atoll, Laccadives, India. Athanas naga Banner and Banner 1966b: 26; type locality: Gulf of Thailand. Athanas nouvelae Holthuis 1951: 104; type locality: Cape Verde Is. Athanas ohsimai Yokoya 1936: 129; type locality: Sagami Bay, Japan. Athanas orientalis Pearson 1905: 88; type 469 locality: Gulf of Manaar, Sri Lanka. Athanas ornithorhynchus Banner and Banner 1973: 319; type locality: Northern Territory, Australia. Athanas parvus de Man 1910: 315; type locality: Timor, Indonesia. Synonym: Athanas sibogae de Man 1910: 314 (see Banner and Banner 1973, Chace 1988). Athanas phyllocheles Banner and Banner 1983: 152; type locality: off La Réunion, Indian Ocean. Athanas polymorphus Kemp 1915: 295; type locality: Chilka Lake, eastern India. Athanas rhothionastes Banner and Banner 1960: 142; type locality: Canton, Phoenix Is., Kiribati. Athanas squillophilus Hayashi 2002: 396; type locality: southern Hokkaido, Japan. Athanas stebbingi de Man 1920: 106; type locality: Pulau Bawean, Indonesia. Athanas tenuipes de Man 1910: 316; type locality: Sulawesi, Indonesia. Athanas sp. nov. 1 aff. dimorphus (A. Anker pers. obs.), Pulau Bintan, Indonesia. Athanas sp. nov. 2 aff. parvus (A. Anker pers. obs.), Vietnam. Athanas sp. nov. 3 aff. amazone (Anker and Ahyong in press), Ivory Coast. Athanas sp. nov. 4 aff. phyllocheles (Anker and Ahyong in press), New South Wales, Australia. Distribution: Indo-West Pacific from the Red Sea to South Africa, Australia, French Polynesia, and Japan; eastern Atlantic from southern Norway and Ireland to southeastern Africa, including the Mediterranean and Black Seas (absent from central and western Atlantic and eastern and northcentral Pacific). Remarks: Athanas, as redefined here, differs from Rugathanas and Arete in several morphological features (see descriptions of Rugathanas and Arete above). The presence of a subtriangular projection on the mesial face of the cheliped coxa, a previously unconsidered feature, may turn out to be a further diagnostic feature of Athanas. This projection was present in all 12 personally examined species of Athanas (A. Anker pers. obs.), including A. dimorphus, A. nitescens, and A. areteformis (Fig. 8g), but is absent in all species of Arete and Rugathanas. With over 30 species, Athanas remains a relatively heterogeneous genus. It was subdivided into 2 informal species groups by Coutière (1899), the Ath. nitescens group (chelipeds carried extend- 470 Zoological Studies 46(4): 454-472 (2007) ed) and the Ath. dimorphus group (chelipeds carried folded). The Ath. nitescens group appears to be relatively homogenous, except for the somewhat aberrant Ath. areteformis and Ath. granti. However, these 2 species clearly belong to the Ath. nitescens group based on the features of the chelipeds (see below). The Ath. dimorphus group is much more heterogeneous and may be further subdivided into several smaller groups mainly defined by the features of the chelipeds, frontal margin, and walking legs (A. Anker pers. obs.). However, the complex intrageneric structure of Athanas is beyond the scope of this study and will be examined elsewhere. The placement of Ath. areteformis within the Ath. nitescens group is based mainly on the features of the male chelipeds. The shape of the frontal margin of the carapace (Figs. 8a, b) and the expanded incisor process of the mandible (Figs. 8c, d) of this species are superficially similar to those of Rugathanas, but the chelipeds (Figs. 8ej), in particular the stout major chela (Fig. 8f), greatly differ from the chelipeds of Rugathanas, and more closely resemble those of Ath. grimaldii (Fig. 9a) or Are. indicus (Fig. 7c). The specific name Ath. areteformis (resembling Arete) given by Coutière (1903) refers to the general resemblance of the chelipeds of this species to the chelipeds of Arete. Banner and Banner (1960) misinterpreted , this as Coutière s doubts about the distinction between Arete and Athanas. Despite some similarities with Rugathanas and Arete, Ath. areteformis plainly belongs to the genus Athanas, as redefined above, because of the general shape of the chelipeds, the non-extensible dactylus of the 3rd-5th pereiopods, the well-developed row of setae on the propodus of the 5th pereiopod, as well as the presence of 3 strap-like epipods on the pereiopods (P1-3), a typical condition in Athanas. Another species that shows some morphological and ecological resemblance to Arete is Ath. granti. This species also has a robust major cheliped (Fig. 9b), stout biunguiculate dactylus on the 3rd-5th pereiopods, and is furthermore, facultatively associated with the sea urchin Centrostephanus rodgersii (Agassiz, 1863) (Banner and Banner 1973). However, the shape of the frontal margin; the marked asymmetry of the chelipeds (the minor cheliped being much slenderer, see Banner and Banner 1973); the 5-segmented carpus of the 2nd pereiopod; the number of pereiopodal epipods; and the absence of tooth on the mesial face on the cheliped ischium, all indicate that Ath. granti is a member of Athanas. Furthermore, in none of the species of the Ath. nitescens group, including Ath. nitescens (Figs. 9c, d), Ath. grimaldii (Fig. 9a), Ath. aretefromis (Fig. 8f), and Ath. granti (Fig. 9b), does the palm of the chela reach the same broadness as in Arete (Fig. 7). Banner and Banner (1973) reported the presence of a median flap on the 5th abdominal sternite in Ath. granti and Are. sp. cf. dorsalis. Interestingly, this flap was absent in the closely related Are. sp. cf. indicus (both Are. dorsalis and Are. indicus were probably misidentified by Banner and Banner (1973), see above). The function of this odd flap is unknown; it may have evolved independently in some Arete species and in Ath. granti, and perhaps plays a role in the symbiosis with the sea urchins. On the other hand, it is also possible that the Arete clade evolved from ancestors within Athanas, in which case the latter genus would be paraphyletic and require further splitting. The morphological complexity of Athanas and the absence of a phylogeny of the Athanas complex (based on molecular and/or morphological characters) make further conclusions impossible. Acknowledgments: The 1st author (AA) is indebted to Dr. A. Richard Palmer (University of Alberta, Edmonton, Canada) for his support in form of an NSERC operating grant (A7245). Mr. Gavin Dally (Museum and Art Gallery of the Northern Territory, Darwin, NT, Australia) arranged the loan of the NTM specimens. Mr. Keiichi Nomura (Kushimoto Marine Park Center, Japan) sent us additional specimens from Japan. 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