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Alpheus nipa Banner & Banner, 1985 (Figs. 44, 45, 52D) Alpheus nipa Banner & Banner 1985: 21, fig. 2. Type material. Paratypes, 5 males (cl 6.2–7.5 mm, tl 16.0 –20.0 mm, all missing chelipeds, but detached major and minor chelipeds present in the vial), 1 ov. female (cl 9.5 mm, tl 23.5 mm, chl 12.0 mm), ZMUC CRU-7526, Indonesia, Strait of Malacca off Sumatra, vicinity of Medan, Galathea Expedition sta. 325, 4°20’N 98°53’E, palm fronds floating on water surface (depth at collection site: 40 m), 10.05.1951. Description. See Banner & Banner (1985) for detailed description and illustrations; complementary illustrations of one of the paratypes are provided in Figs. 44, 45. Colour pattern. Unknown. Type locality. Off Sumatra, Indonesia. Distribution. Indo-West Pacific: presently known only from the type locality in the Straits of Malacca off Sumatra, western Indonesia (Fig. 52D). Common name proposed. Nipa palm snapping shrimp. Ecology and biology. All known specimens of A. nipa were found on floating fronds of nipa palm (Nypa fructicans Wurmb, Arecaceae), suggesting that they may inhabit estuarine muddy banks fringed by nipa palms, a habitat that is both difficult to access and dangerous to sample due to the presence of saltwater crocodiles [Crocodylus porosus (Schneider)]. Taxonomic remarks. Banner & Banner (1985) provided a reasonably detailed description of A. nipa, accompanied by illustrations of all major diagnostic features. Two important features of A. nipa not mentioned nor illustrated by Banner & Banner (1985) are the unarmed antennal basicerite (Fig. 45B) and the presence of a faint mesial subdistal ridge on the major cheliped pollex (Figs. 44B, 45G), which may link this species to the A. euphrosyne — A. microrhynchus complex. Another previously unknown feature of A. nipa is the presence of three small, gill-like leaflets at the base of the third maxilliped exopod; this feature was observed on two dissected third maxillipeds and one maxilliped still in situ. This gill-like structure does not appear to be homologous to the “rudimentary pleurobranch at the base of the third maxilliped” in A. pontederiae (Christoffersen 1984) and “une petite pleurobranchie de forme très spéciale sur le 3 e mxp.” in several other species of Alpheus, including A. euphrosyne, A. microrhynchus and A. macrodactylus (see discussion in Coutière 1899: 279–281). As noted by Banner & Banner (1985), A. nipa can be separated from A. euphrosyne, A. richardsoni, A. paludicola and A. microrhynchus by the dactyli of the third and fourth pereiopods being short and conical rather than spatulate (in the first three species) or trigonal-subspatulate (in the latter species). This distinguishing character is also valid for A. eurydactylus, A. nomurai sp. nov., A. mangalis sp. nov., A. takla sp. nov. and A. songkla, all of which with spatulate dactyli on the ambulatory pereiopods. In addition, A. nipa differs from A. microrhynchus by the markedly different ratio of the first two carpal subarticles of the second pereiopod (1: 0.8 in A. nipa vs. 1: 0.3 in A. microrhynchus); from both A. microrhynchus and A. cyanoteles by the much stronger sculpture of the male minor chela, with both dorsal and ventral transverse grooves well pronounced and bordered by shoulders (vs. feebly delineated, if it all, and without shoulders, in A. microrhynchus and A. cyanoteles); and the scaphocerite blade much shorter than the distolateral tooth (vs. reaching slightly or far beyond the distolateral tooth in A. microrhynchus and A. cyanoteles). Another diagnostic character of A. nipa is the presence of several long erect setae on the rostrum (Fig. 45A, B; see also Banner & Banner 1985: fig. 2a, b); these setae are absent in the other species being dealt with in this study (or if present, not nearly as conspicuous as in A. nipa). Banner & Banner (1985) suggested that A. nipa “is derived from one of these euryhaline mud-or soft sedimentdwelling species, possibly from the most common and widespread A. euphrosyne * and has left the mud and silt where the other three species [A. euphrosyne *, A. richardsoni, A. paludicola] dwell and has invaded a unique habitat, the nipa palms. In so doing, it has lost the spatulate dactyli on the walking legs, so necessary for digging in soft sediments” [*note: A. euphrosyne not sensu De Man (1897)]. This is indeed an interesting hypothesis to be tested in the future, and to be contrasted to an alternative hypothesis of A. nipa not being closely related to the A. euphrosyne — A. microrhynchus complex and being derived, for instance, from the morphologically similar A. lobidens complex, in which the walking leg dactyli are also conical. Whatever the case may be, the origin and phylogenetic affinites of A. nipa are difficult to ascertain without a comprehensive phylogenetic analysis of the entire A. edwardsii group, with inclusion of as many members as possible of the A. euphrosyne — A. microrhynchus and A. lobidens complexes., Published as part of Anker, Arthur, 2023, Revision of Alpheus euphrosyne De Man, 1897 and A. microrhynchus De Man, 1897, with description of three new species and taxonomic remarks on several other morphologically and ecologically similar snapping shrimps (Malacostraca: Decapoda: Alpheidae), pp. 1-115 in Zootaxa 5282 (1) on pages 83-85, DOI: 10.11646/zootaxa.5282.1.1, http://zenodo.org/record/7912292, {"references":["Banner, D. M. & Banner, A. H. (1985) The alpheid shrimp of Indonesia, based upon J. G. de Man's \" The Decapoda of the Siboga Expedition, Part II. Family Alpheidae \" (1911). Marine Research in Indonesia, 25, 1 - 79. https: // doi. org / 10.14203 / mri. v 25 i 0.431","Christoffersen, M. L. (1984) The western Atlantic snapping shrimps related to Alpheus heterochaelis Say (Crustacea, Caridea) with the description of a new species. Papeis Avulsos de Zoologia, 35, 189 - 208.","Coutiere, H. (1899) Les \" Alpheidae \". Morphologie externe et interne, formes larvaires, bionomie. Annales des Sciences Naturelles, Zoologie et Paleontologie, Serie 8, 9, 1 - 559, pls. 1 - 6. Masson, Paris. https: // doi. org / 10.5962 / bhl. title. 13143","De Man, J. G. (1897) Bericht u ¨ ber die von Herrn Schiffscapitan Storm zu Atjeh, an den westlichen Ku ¨ sten von Malakka, Borneo und Celebes sowie in der Java-See gesammelten Decapoden und Stomatopoden. Fu ¨ nfter Theil. Z oologische Jahrbu ¨ cher. Abtheilung fu ¨ r Systematik, Geographie und Biologie der Thiere, 9, 725 - 790, pls. 34 - 37 (in volume 10)."]}

Alpheus nomurai sp. nov. (Figs. 25–28, 51F) Alpheus bis-incisus (or A. bisincisus).— Dôtu 1961: 136, fig. 1; Kim & Park 1972: 198; Kim 1976: 142; Kim 1977: 247, figs. 100, 101, pl. 24, fig. 44 [not A. bisincisus De Haan, 1844]. Alpheus euphrosyne.— Jeng & Chang 1985: 248, fig. 23 [not A. euphrosyne De Man, 1897]. Alpheus euphrosyne richardsoni.— Miya 1995: 275, fig. 2; Yang & Kim 1996: 106, figs. 1-3; Hayashi 1998: 292, figs. 353c, g, 354c, 355c, e [not A. richardsoni Yaldwyn, 1971]. Alpheus richardsoni.— Yoshigou 2009: 249, fig. 12m, 13m, 14m, 15m,. 16m, 17m, 18m, pl. III 1-4; Koyama et al. 2017: 132 [not A. richardsoni Yaldwyn, 1971]. Alpheus sp. — Kirihara et al. 2021: 99, fig. 1B. Type material. Holotype, male (cl 8.8 mm, tl 23.0 mm, chl 13.0 mm), MNHN-IU-2018-5590, Japan, Wakayama, Tanabe, intertidal, leg. I. Sakamaki, 27.11.1993 [SMP-940]. Paratypes: 1 male (cl 8.5 mm), MNHN-IU-2018-5588, Japan, Wakayama, Tanabe, intertidal, leg. K. Nomura, 25.08.1995 [SMP-1220]; 1 ov. female (cl 8.8 mm), MNHN-IU-2018-5744, same collection data as for previous paratype [SMP-1220a]; 1 male (cl 9.6 mm), OUMNH. ZC. 2011.06.6, same collection data as for previous paratypes [SMP-1220b]; 1 ov. female (cl 11.9 mm), MNHN-IU-2018-5589, Japan, Kochi, Shimanto River, intertidal, leg. K. Nomura, 12.07.2002 [SMP-1748]. Additional material. Japan: 1 ov. female (cl 9.6 mm, tl 27.5 mm, chl 12.7), MNHN-IU-2018-5748, OkinawaJima, Sajiki, intertidal, leg. K. Nomura, 17.10.1994 [SMP-1334]; 1 male (cl 6.5 mm), 1 ov. female (7.7 mm), OUMNH. ZC. 2019.06.58, Okinawa, Sajiki, intertidal, leg. A. Kikukawa, 08.08.1994 [SMP-1332]. South Korea: 1 male (8.7 mm), 1 ov. female (cl 8.2 mm), MNHN-IU-2018-5587 (ex-SNU-220610-012), Garorim Bay, Yecheon-dong, Seosan-si, Chungcheongnam-do, 36°53’5.74”N – 126°23’23.46”E, collector unknown (probably H.J. Yang), 22.06.2002. Taiwan: 1 male (cl 11.8 mm), 2 females (cl 5.5, 8.5 mm), OUMNH. ZC. 2019.06.59, Chang Hua, mudflat, leg. T. Y. Chan et al., 12.1996; 1 juv. male (cl 4.7 mm), ZRC 2014.0676, Shinzhu, mangrove, leg. Y. Cai & C.W. Lin, 28.11.1997. Description. See Figs. 25–27. Medium-sized species of Alpheus (maximal cl 12.1 mm, tl ~ 36.3 mm; present study; Miya 1995). Carapace glabrous, finely pitted, without pubescence, with slight grooves. Rostrum short, triangular, distally acute, typically reaching 0.3 length of first article of antennular peduncle, sometimes its mid-length; rostral carina present as low elevation, fading posterior to eye level; rostro-orbital furrows poorly demarcated or indistinct. Pterygostomial angle broadly rounded; cardiac notch deep. Pleon smooth, scarcely pitted, without pubescence. Telson moderately broad, ovate-rectangular, gently tapering towards posterior margin, about 1.7–1.8 times as long as maximal width near proximal margin; lateral margins with broad concavity on proximal half; dorsal surface finely pitted, without trace of median depression, with two pairs of short spiniform setae inserted in deep pits at considerable distance from lateral margins, first pair near telson mid-length, second pair between 0.7 and 0.8 of telson length; posterior margin broadly rounded, with two pairs of spiniform setae at posterolateral angles, lateral shorter than mesial. Antennular peduncle with stylocerite broad, markedly convex laterally, with acute tip, latter reaching, but not overreaching distal margin of first article; ventromesial carina with anteriorly directed, distally subacute tooth; second article about 2.3–2.4 times as long as wide. Antennal peduncle with basicerite armed with small, sharp tooth; carpocerite reaching far beyond scaphocerite; scaphocerite with lateral margin nearly straight to very shallowly concave; blade moderately broad, separated from broad distolateral tooth by deep cleft; anterior margin of blade broadly rounded, exceeded by stout distolateral tooth. Third maxilliped with antepenultimate article not particularly broadened; penultimate article elongate, about four times as long as wide, with long setae on ventral margin; ultimate article with dense transverse rows of short serrulate setae on mesial surface and longer setae on dorsal and lateral surfaces, especially near or on apex; coxal lateral plate acutely protruding dorsally; exopod reaching end of antepenultimate article. Major cheliped of A. edwardsii - type. Merus stout, about twice as long as wide; ventromesial margin unarmed distally. Carpus very short, cup-shaped. Chela large, massive, with fingers about 0.7 length of palm. Mesial face of palm largely smooth, except for small field of minute granules dorsal to ventral shoulder, and with weak mesial transverse ridge in proximal half, extending from proximal edge of subtriangular mesial longitudinal groove to ventral surface of palm. Lateral face of palm smooth; lateral longitudinal groove deep, subrectangular; proximal half without distinct transverse ridge. Dorsal shoulder smooth, gently sloping into broad transverse groove with angle of about 45°. Ventral shoulder pronounced, slightly protruding in lateral view, broadly rounded, smooth. Pollex with mesial face finely granulated over most of its surface, with short subdistal mesial ridge near margin, without proximal protuberance; area ventral to mesial subdistal ridge not grooved; distomesial angle distinctly superior to 90°, blunt; lateral face smooth; distolateral angle superior to 90°. Dactylus with weak dorsal ridge, latter slightly twisted mesially, not protruding proximally; mesial and lateral surfaces smooth; dactylar plunger very stout, large, with anterior margin well delineated, clearly separated from ventral margin of dactylus; distal surface of plunger somewhat truncate. Adhesive discs small. Male minor cheliped with chela strongly balaeniceps. Merus slightly slenderer than that of major cheliped, about 2.5 times as long as wide; ventral margin with distomesial angle unarmed; distomesial margin slightly rugose. Chela not particularly swollen, slender in lateral view, with fingers about as long as palm; all surfaces smooth, without granulation. Palm subcylindrical, compressed, feebly swollen; both mesial and lateral longitudinal grooves well marked, deep; dorsal shoulder distinct, rounded, sloping almost perpendicularly into dorsal transverse groove; ventral transverse groove present, relatively deep, marked by distinct ventral shoulder on lateral surface. Pollex with mesial surface bearing row of balaeniceps setae on low crest extending from base to about mid-length of pollex; lateral surface with similar row of balaeniceps setae as on mesial surface. Dactylus with conspicuous lateral expansion (balaeniceps expansion) in proximal half, about 1.7 as long as maximal width; lateral and mesial dactylar ridges each with dense rows of balaeniceps setae. Cutting edges of both fingers shallowly excavated on each side of sharp, blade-like ridge. Female minor cheliped simple, not balaeniceps. Chela slender, not particularly swollen, with fingers subequal to, or slightly longer than palm. Palm with surface smooth, not granulated; mesial longitudinal groove feebly developed; lateral longitudinal groove distinct, but shallow; dorsal transverse groove usually distinct, but dorsal shoulder poorly demarcated; ventral transverse groove present in form of deep sinus bordered by feebly developed ventral shoulder, latter sometimes not distinct (especially in younger females or regenerated chelipeds). Fingers simple, with sharp cutting edges, slightly gaping when closed; fingertips strongly curved and crossing. Second pereiopod with ratio of carpal subarticles approximately equal to 4: 2.3: 1: 1: 1.7. Third pereiopod moderately robust; ischium armed with short, stout spiniform seta on ventrolateral surface; merus about 5.2 times as long as maximal width, unarmed; propodus with three or four short spiniform setae on ventral margin and one spiniform seta on distoventral margin adjacent to dactylus; dactylus about half-length of propodus, distinctly spatulate, expanded, wider than propodus in dorsal view, with two rows of setae along ridges. Fourth pereiopod generally similar to third pereiopod, shorter and slenderer; dactylus comparatively shorter, about 0.4 length of propodus. Fifth pereiopod much slenderer than third and fourth pereiopods; ischium unarmed; propodus apparently without spiniform setae, with well-developed setal brush; dactylus subspatulate, much less expanded compared to that of third or fourth pereiopod, about half-length of propodus. Male second pleopod with appendix masculina 0.8 length of appendix interna. Uropod with each protopodal lobe ending in subacute tooth distally; exopod and endopod broad, ovate; diaeresis of exopod complete, almost straight, with broadly subtriangular lobe near small distolateral spiniform seta and adjacent distolateral tooth; lateral margin of exopod broadly convex. Eggs numerous (> 100 in larger females), small; egg diameter about 0.65 mm. Colour pattern. Carapace greyish, semi-opaque, with two complete transverse bands of dark brown to dark grey-blue colour, one broad complete band near posterior margin and one narrower, laterally broken band at about carapace mid-length; two smaller, incomplete (dorsally not closing) and more diffuse bands present in anterior half of carapace, one along its anterolateral margin and one lateral to cardiac area; pleon with six moderately broad, dark blue-grey, blue-green or red-brown transverse bands, one across each pleonite and near its posterior margin; broad white spaces between dark bands with small but conspicuous neon-yellow spots, most of them fringing dark bands on carapace and pleon; antennular peduncles and scaphocerite patchily marbled with brown and yellow; antennular and antennal flagella pale yellow or pale greenish; mesial face of major chela olive-green, with paler greenish or orange-yellowish areas, especially on dorsal and ventral transverse grooves; dactylus dark olive-green; distal portion of both fingers pale orange; minor chela olive-green mottled with brown, fingers darker; second to fifth pereiopods pale yellowish; telson and uropods with patchy brown mottling and yellow spots, darker distally; posterior half of uropodal exopod dark blue (Fig. 28; see also Jeng & Chang 1985: fig. 23; Yoshigou 2009: pl. III, figs. 1–4). Type locality. Tanabe, Wakayama Prefecture, Honshu, Japan (collection locality of the holotype). Distribution. Temperate and subtropical north-western Pacific (Fig. 51F): Japan (Honshu, Kyushu, Shikoku, Okinawa), South Korea (Garorim Bay, Byeonsan = Pyonsan) and Taiwan (Chang Hua, Shinzhu, Tamsui River). Etymology. This species is named after the author’s dear colleague, Keiichi Nomura (Sabiura Marine Park Research Station, Japan), who contributed to alpheid taxonomy early in his career and made important material available for the present study. Common name proposed. Nomura’s snapping shrimp. Ecology and biology. Alpheus nomurai sp. nov. is a marine / estuarine snapping shrimp largely confined to intertidal mudflats, especially those situated inside or close to estuaries. The shrimps usually live in pairs (rarely as singles) in complex tridimensional burrows made in soft mud, with side branches extending from the burrow entrance in a radius of about 25 cm and penetrating about 10–20 cm deep into the mud (Miya 1995, as A. euphrosyne richardsoni, based on Dôtu 1961: fig. 1). The burrows of A. nomurai sp. nov. are typically composed of a main tunnel with one or two or three openings and multiple branches, either interconnected or ending blindly (see Kirihara et al. 2021: figs. 2, 3, as Alpheus sp.). They often serve as shelter for small gobies, such as Apocryptodon punctatus Tomiyama, 1934, Gymnogobius cylindricus (Tomiyama, 1936) and Acentrogobius sp. (Miya 1995; Suzuki & Wada 1999; Yoshigou 2009; Koyama et al. 2017; Kirihara et al. 2021, as Alpheus sp., A. euphrosyne richardsoni or A. richardsoni). It must be noted that the association between the snapping shrimp identified as A. richardsoni and the goby Drombus ocyurus (Jordan & Seale, 1907) in Kunishima et al. (2022, as Acentrogobius ocyurus) involves a species from the A. malabaricus complex, possibly A. dolichodactylus Ortmann, 1890 (see under the synonymy of A. richardsoni). Whether or not A. ocyurus also associates with A. nomurai sp. nov. remains to be shown. Some shrimps were found in burrows made under large intertidal rocks partly embedded in mud (Jeng & Chang 1985, as A. euphrosyne). The large number of eggs and their small size suggest an extended larval development in A. nomurai sp. nov. (Yang & Kim 1996, as A. euphrosyne richardsoni). Taxonomic remarks. Alpheus nomurai sp. nov. was previously confused with two other species of Alpheus, viz. the distantly related A. bisincisus De Haan, 1844 (Dôtu 1961) and the presumably more closely related A. richardsoni (Miya 1995, as A. euphrosyne richardsoni). The morphological differences between A. nomurai sp. nov. and A. richardsoni are relatively subtle, which explains Miya’s (1995) identification of the Japanese and Korean material as A. euphrosyne richardsoni, despite the resulting disjunct, antitropical distribution for A. richardsoni. Nevertheless, A. nomurai sp. nov. and A. richardsoni can be separated from each other by the shape of the dactylar plunger of the major chela, which appears to broader and stouter in A. nomurai sp. nov. (cf. Fig. 27D; Miya 1995: fig. 2C; Banner & Banner 1982: fig. 74e); the presence of a small subdistal mesial ridge in A. nomurai sp. nov., which is not distinct in A. richardsoni (cf. Fig. 27C; Banner & Banner 1982: fig. 74c); the size and position of the tooth on the distolateral margin of the antennal basicerite, which is more developed and more ventral in A. nomurai sp. nov. (cf. Fig. 26D; Miya 1996: fig. 2A; Banner & Banner 1982: fig. 74b); and the armature of the third pereiopod propodus, which has only three to four spiniform setae in A. nomurai sp. nov. vs. at least six in A. richardsoni (cf. Fig. 26G; Miya 1995: fig. 2F; Banner & Banner 1982: fig. 74n). In addition, A. nomurai sp. nov. with the maximal total length not exceeding 36.5 mm is a much smaller species compared to A. richardsoni, which may reach 65 mm. The most obvious and reliable distinguishing feature between these two species involves the banding pattern of the pleon: in A. nomurai sp. nov., each pleonite has a simple dark transverse band (Fig. 28, disregarding diffuse bands formed by neon-yellow spots), whereas in A. richardsoni, each pleonite has a double or even triple band (Figs. 17, 18). Of course, the two species are also geographically widely separated, with A. nomurai sp. nov. and A. richardsoni each restricted to temperate and subtropical north-eastern Asia and Australia / New Zealand, respectively (Fig. 51C, F). Alpheus nomurai sp. nov. can be separated from A. euphrosyne and A. eurydactylus essentially by the same criteria as A. richardsoni (see above), and from A. microrhynchus and A. cyanoteles by the conspicuously broadened, spatulate dactylus of the third to fifth pereiopods, which in the other two species is trigonal-subspatulate in shape (cf. Miya 1995: fig. 2F; Yeo & Ng 1996: fig. 6b, d). For separation of A. nomurai sp. nov. from other morphologically similar species see below and refer to Table 1., Published as part of Anker, Arthur, 2023, Revision of Alpheus euphrosyne De Man, 1897 and A. microrhynchus De Man, 1897, with description of three new species and taxonomic remarks on several other morphologically and ecologically similar snapping shrimps (Malacostraca: Decapoda: Alpheidae), pp. 1-115 in Zootaxa 5282 (1) on pages 50-57, DOI: 10.11646/zootaxa.5282.1.1, http://zenodo.org/record/7912292, {"references":["Dotu, Y. (1961) The bionomics and life history of the gobioid fish, Apocryptodon bleekeri (Day). Bulletin of the Faculty of Fisheries, Nagasaki University, 10, 133 - 139, pl. 28. [in Japanese]","Kim, H. S. & Park, K. B. (1972) Faunal studies on the macrurans in Korea. In: Floral studies on some taxa of plants and faunal studies on some taxa of animals in Korea. R- 72 - 82. Ministry of Science and Technology, pp. 185 - 216, pls. 1 - 6. [in Korean with English summary]","Kim, H. S. (1976) A checklist of the Macrura (Crustacea, Decapoda) of Korea. Proceedings of the College of Natural Science, Seoul National University, 1, 131 - 152.","Kim, H. S. (1977) Macrura. Illustrated Flora and Fauna of Korea. Vol. 19. Samwha Publishing Co., Seoul, 414 pp., 56 pls. [in Korean]","De Haan, W. (1844) [1833 - 1850] Crustacea. In: von Siebold, P. F. (Ed.), Fauna Japonica sive Descriptio Animalium, quae in Itinere per Japoniam, Jussu et Auspiciis Superiorum, qui Summum in India Batava Imperium Tenent, Suspecto, Annis 1823 - 1830 Collegit, Notis, Observationibus et Adumbrationibus Illustravit. Apud Auctorem, Lugduni-Batavorum, pp. i - xxxi + ix - xvi + 1 - 243, pls. A - J + L - Q + 1 - 55.","Jeng, M. S. & Chang, K. H. (1985) Snapping shrimps (Crustacea: Decapoda: Alpheidae) of Taiwan. Bulletin of Institute of Zoology, Academica Sinica, 24, 241 - 256.","De Man, J. G. (1897) Bericht u ¨ ber die von Herrn Schiffscapitan Storm zu Atjeh, an den westlichen Ku ¨ sten von Malakka, Borneo und Celebes sowie in der Java-See gesammelten Decapoden und Stomatopoden. Fu ¨ nfter Theil. Z oologische Jahrbu ¨ cher. Abtheilung fu ¨ r Systematik, Geographie und Biologie der Thiere, 9, 725 - 790, pls. 34 - 37 (in volume 10).","Miya, Y. (1995) Four species of Alpheus from intertidal and shallow water mudflats in the Sea of Ariake, Kyushu, Japan (Crustacea, Decapoda, Alpheidae). Bulletin of the Faculty of Liberal Arts, Nagasaki University, Natural Science, 35, 271 - 288.","Yang, B. P. & Kim, C. H. (1996) Zoeal stages of Alpheus euphrosyne richardsoni Yaldwyn, 1971 (Decapoda: Macrura: Alpheidae) reared in the laboratory. Korean Journal of Zoology, 39, 106 - 114.","Hayashi, K. I. (1998) Prawns, shrimps and lobsters from Japan (101). Family Alpheidae - Genus Alpheus (10). Aquabiology, 117 (20), 289 - 293.","Yaldwyn, J. C. (1971) Preliminary descriptions of a new genus and twelve new species of natant decapod Crustacea from New Zealand. Records of the Dominion Museum, 7, 85 - 94.","Yoshigou, H. (2009) A collection of alpheid shrimps (Crustacea; Decapoda; Alpheidae) from the mouth of rivers and anchialine caves of Japan. Miscellaneous Reports of the Hiwa Museum for Natural History, 50, 221 - 273. [in Japanese with English abstract]","Koyama, A., Inui, R., Sawa, K. & Onikura, N. (2017) Symbiotic partner specificity and dependency of two gobies (Apocryptodon punctatus and Acentrogobius sp. A) and four alpheid shrimps inhabiting the temperate estuary of southern Japan. Ichthyological Research, 64, 131 - 138. https: // doi. org / 10.1007 / s 10228 - 016 - 0535 - 6","Kirihara, S., Itani, G., Nunobe, J. I., Nomoto, A., Aldea, K. Q., Murakami, R., Sakata, H. & Henmi, Y. (2021) Burrow morphology of an alpheid shrimp at muddy tidal flats in western Japan. Kuroshio Science, 14, 99 - 102.","Tomiyama, I. (1934) Four species of gobies of Japan. Journal of the Tokyo Imperial University, Faculty of Science, Section IV, 3, 325 - 334.","Tomiyama, I. (1936) Gobiidae of Japan. Japanese Journal of Zoology, 7, 37 - 112.","Suzuki, T. & Wada, K. (1999) Record of a gobiid fish, Apocryptodon punctatus from the Uchinoura Inlet in Wakayama Prefecture, Japan. Nankiseibutu, 41, 61 - 63.","Jordan, D. S. & Seale, A. (1907) Fishes of the islands of Luzon and Panay. Bulletin of the Bureau of Fisheries, 26, 1 - 48.","Kunishima, T., Palla, H. P., Tachikara, K. & Maeda, K. (2022) First records of an estuarine goby Acentrogobius ocyurus (Gobiiformes: Gobiidae) from Japan and the Sulu Sea in the Philippines. Species Diversity, 27, 129 - 138. https: // doi. org / 10.12782 / specdiv. 27.129","Ortmann, A. (1890) Die Decapoden-Krebse des Strassburger Museums, mit besonderer Beru ¨ cksichtigung der von Herrn Dr. D ˆ derlein bei Japan und bei den Liu-Kiu-Inseln gesammelten und z. Z. im Strassburger Museum aufbewahrten Formen. I. Theil. Die Unterordnung Natantia (Boas) (Abtheilungen: Penaeidae und Eucyphidea = Caridae der Autoren). Zoologische Jahrbu ¨ cher. Abtheilung fu ¨ r Systematik, Geographie und Biologie der Thiere, 5, 437 - 542, pls. 36, 37.","Banner, D. M. & Banner, A. H. (1982) The alpheid shrimp of Australia. Part III: The remaining alpheids, principally the genus Alpheus and the family Ogyrididae. Records of the Australian Museum, 34, 1 - 357. https: // doi. org / 10.3853 / j. 0067 - 1975.34.1982.434","Yeo, D. C. J. & Ng, P. K. L. (1996) A new species of freshwater snapping shrimp, Alpheus cyanoteles (Decapoda: Caridea: Alpheidae) from Peninsular Malaysia and a redescription of Alpheus paludicola Kemp, 1915. Raffles Bulletin of Zoology, 44, 37 - 63."]}

Alpheus takla sp. nov. (Figs. 2F, G, 29–33, 34B–E, 35A, B, D, E, 52A) (?) Alpheus microrhynchus.— Johnson 1965: 9 (part.) [not A. microrhynchus De Man, 1897]. Alpheus euphrosyne euphrosyne. — Banner & Banner 1982: 232 (part.?), fig. 73 (part., not fig. 73i); Chace 1988: 27 (part.?) [not A. euphrosyne De Man, 1897]. (?) Alpheus euphrosyne. — Banner & Banner 1966: 130 (part.?, not fig. 49); Wada 2013: 184, fig. 5 [not A. euphrosyne De Man, 1897]. (?) Alpheus sp. cf. audouini .— Johnson 1962: 53 (part.). Alpheus macellarius. — Aaron-Cabrillos 2019: 22, fig. 3 [not A. macellarius Chace, 1988]. Alpheus sp. — Baobao et al. 2015: 337. Type material. Holotype, male (cl 25.5 mm, tl ~70.0 mm, chl 41.0 mm), NMCR 99110, Philippines, Bohol, from fishermen, leg. Y. Cai, 18.12.2000. Paratypes: 4 males (cl 24.1–27.0 mm), 1 female (cl 24.6 mm), 1 ov. female (cl 27.2 mm), ZRC 2014.0668, same collection data as for holotype; 2 males (cl 23.9, 25.0 mm), OUMNH. ZC. 2019.06.66, same collection data as for holotype; 2 males (cl 23.3, 24.4 mm), FLMNH UF 68697, same collection data as for holotype; 3 males (cl 24.0–25.0 mm), 1 female (cl 27.6 mm), MNHN-IU-2022-2939, same collection data as for holotype. Additional material. Philippines: 1 female (cl 22.0 mm), OUMNH. ZC. 2019.06.67, Cebu, Cebu City, fish market, leg. P.K.L. Ng et al., 14.12.2000; 2 males (cl 23.5, 25.7 mm), 1 female (cl 23.0 mm), OUMNH. ZC. 2019.06.68, unknown locality, fisheries import, leg. K. Nomura, 01.1998; 1 female (cl 26.4 mm), CBM 17105, same collection data as for previous specimens [YMP-1667h]; 1 female (cl 19.3 mm, missing both chelipeds), NMCR 99111, Mindanao, mangrove forest along coastal road between Dipolog and Dapitan, leg. J.C.Y. Lai & N.K. Ng, 29.01.2005; 1 male (cl 22.6 mm), USNM 252030, Luzon, Manila, market, no further data, 04.01.1908; 1 male (cl 23.5 mm), 2 females (cl 21.9, 25.4 mm), USNM 128409, Mactan, Calawisan, sta. 20578, leg. Santos, 28.05.1968. Singapore: 1 male (cl 24.7 mm), 1 female (cl 26.7 mm), ZRC 1979.4.4.7–8, Jurong, prawn ponds, sta. B26, leg. S. R.F. R.S., 29.06.1954 [J7225–7226]; 1 female (cl 21.7 mm, missing major cheliped), ZRC 1979.4.4.9, Jurong, prawn ponds, sta. B26, leg. S. R.F. R.S., 29.06.1954 [J7994]; 1 male (cl 20.7 mm), 4 females (cl 19.8–29.7 mm), ZRC 1979.4.4.1–5, Jurong, prawn ponds, sta. B17, leg. S. R.F. R.S., 1– 2.06.1954; 1 female (cl 15.9 mm) + 1 major cheliped, ZRC 2014.0680, Jurong prawn pond, no further data [J8049]; 1 male (cl 21.5 mm), 1 female (15.8 mm), ZRC 2014.0673, Singapore, no further data [ex cat. nr. 530, Nanyang University Department of Biology, det. “ Callianassidae, Alpheus bisincisus ”]; 1 male (cl 26.4 mm), NHM 1975.457.1, unknown locality, leg. Singapore Fisheries Research Station (det. A.H. Banner as A. euphrosyne). Thailand: 1 ov. female (cl 32.5 mm, tl 86.0 mm, chl 52.0 mm), ZRC 1999.0850, Thailand, Ranong, 08.09.1929. Indonesia: 1 male (cl 21.0 mm), OUMNH. ZC. 2019.06.69, Papua, Ajkwa Island, mangrove, leg. A. Darmawan et al., 20.07.2012; 1 female (cl 17.3 mm), MNHN-IU-2018-5596, Papua, Kamora, mangrove, leg. A. Darmawan et al., 07.08.2012; 1 female (cl indet., tl 85.0 mm), ZMA Crust.De. 202792, Sulawesi, Makassar, Siboga-Expedition sta. 71, mud, sand with mud, coral, depth: to 32 m (?), 10.05.1899 – 07.06.1899. Papua New Guinea: 1 male (cl 15.7 mm), USNM 1172134, Madang Harbour, Naui Island, mangrove, poison station BBC1492, depth: 0–1 m, leg. B. Collette, 28.05.1970. Australia: 1 male (cl 24.8 mm), 2 females (cl 27.2, 29.0 mm), NTM Cr. 002639, Queensland, Gladstone, Calliope River, sta. 151, leg. P. Saenger, 1982; 1 female (cl 31.0 mm), NTM Cr. 000865, Queensland, Townsville, mangrove swamp, don. P. Sammarco, 15.02.1984; 1 male (cl 27.4 mm, missing major cheliped), AM P 28129, Queensland, Mangrove Creek south of Ross River, 19°22’S 146°44E, leg. B. Campbell, 03.1962. Tentative identification. Alpheus cf. takla sp. nov. Indonesia: 1 female (cl indet., tl 61.0 mm), ZMA Crust. De. 202791, eastern Java, Siboga-Expedition sta. 4, eastern Java, anchorage off Jangkar (= Djangkar), 7°42’S, 114°12.6’E, shore, 09.03.1899. Description. See Figs. 2F, G, 29–31. Very large species of Alpheus (maximal cl 32.5 mm, tl 86.0 mm in present material, but see below). Carapace glabrous, without pubescence, with slight grooves. Rostrum short, subtriangular, distally acute, not reaching 0.3 length of first article of antennular peduncle; rostral carina present as faint elevation extending at most to eye level; rostro-orbital furrows poorly demarcated. Pterygostomial angle broadly rounded; cardiac notch deep. Pleon smooth, without pubescence. Telson very broad, subrectangular, very slightly tapering towards posterior margin, about 1.7 times as long as maximal width near proximal margin; lateral margins almost straight; dorsal surface with shallow longitudinal median depression (sometimes barely distinct or absent) and two pairs of short spiniform setae inserted in deep pits at some distance from lateral margins, first pair near telson mid-length, second pair between 0.7 and 0.8 of telson length; posterior margin rounded, with two pairs of small spiniform setae at posterolateral angles, lateral shorter than mesial. Antennular peduncle with stylocerite broad, markedly convex laterally, with acute tip, latter slightly overreaching distal margin of first article; ventromesial carina with low, blunt tooth; second article about 2.1–2.2 times as long as wide. Antennal peduncle with basicerite armed with stout, sharp tooth; carpocerite reaching distinctly beyond scaphocerite; scaphocerite with lateral margin nearly straight to slightly convex; blade very broad, separated from broad distolateral tooth by deep cleft; anterior margin of blade broadly rounded, greatly exceeding tip of distolateral tooth. Third maxilliped with antepenultimate article not particularly broadened; penultimate article elongate, about 3.5 times as long as wide, slightly widening distally, with some elongate setae on ventral margin; ultimate article with dense transverse rows of short serrulate setae on mesial surface and longer setae on dorsal and lateral surfaces, especially near or on apex; coxal lateral plate bluntly protruding dorsally; exopod overreaching end of antepenultimate article. Major cheliped of A. edwardsii - type. Merus stout, about twice as long as wide; ventromesial margin unarmed distally. Carpus very short, cup-shaped. Chela large, massive, with fingers about 0.6 length of palm. Mesial face of palm largely smooth, with slight mesial transverse ridge (or rather elevation) in proximal half, running obliquely from central area towards ventral surface of palm. Lateral face of palm smooth; lateral longitudinal groove deep, subrectangular; proximal half without clear transverse ridge. Dorsal shoulder smooth, gently sloping into broad transverse groove with angle of about 45°. Ventral shoulder pronounced, not protruding in lateral view, broadly rounded, smooth. Pollex with mesial face usually finely granulated, especially near distomesial angle, with well-developed subdistal mesial ridge near margin, marked by proximal protuberance; area ventral to mesial subdistal ridge deeply depressed; distomesial angle distinctly about 90°, blunt; lateral face smooth; distolateral angle superior to 90°. Dactylus with short, dorsally protruding dorsal ridge; mesial surface with barely detectable granulation; lateral surface smooth; dactylar plunger very stout, large, with anterior margin well delineated, separated from ventral margin of dactylus; distal surface of plunger rounded anteriorly, remaining area obliquely truncate.Adhesive discs small. Male minor cheliped with chela strongly balaeniceps. Merus distinctly slenderer than that of major cheliped, about 2.5 times as long as wide; ventral margin with distomesial angle unarmed. Chela feebly swollen, slender in lateral view, with fingers about as long as palm. Palm subcylindrical, compressed, feebly swollen; all surfaces smooth, without granulation; both mesial and lateral longitudinal grooves well marked; dorsal shoulder and transverse groove poorly marked, former present only in form of slight, gently sloping elevation; ventral transverse groove poorly developed, marked by smooth ventral shoulder on lateral surface. Pollex with mesial surface bearing row of balaeniceps setae on low crest extending from base to mid-length of pollex or beyond it; lateral surface with similar row of balaeniceps setae as on mesial surface. Dactylus with conspicuous lateral expansion (balaeniceps expansion) in proximal half, about 1.7 as long as maximal width; lateral and mesial dactylar ridges each with dense rows of balaeniceps setae. Cutting edges of both fingers shallowly excavated on each side of sharp, blade-like ridge. Female minor cheliped simple, not balaeniceps. Chela slender, not particularly swollen, with fingers slightly longer than palm. Palm with surface smooth, not granulated; mesial longitudinal groove not distinct; lateral longitudinal groove feebly developed, very shallow; dorsal and ventral transverse groove and shoulders not distinct, ventral shoulder indicated by very faint sinus laterally. Fingers simple, with sharp cutting edges, slightly gaping when closed; dactylus with low longitudinal ridge laterally; fingertips strongly curved and crossing. Second pereiopod with ratio of carpal subarticles approximately equal to 4.3: 2: 1: 1: 1.5. Third pereiopod moderately robust; ischium unarmed or armed with very small spiniform seta on ventrolateral surface; merus about four times as long as maximal width, unarmed; propodus with six or so stout spiniform setae on ventral margin and at least one stout spiniform seta on distoventral margin adjacent to dactylus; dactylus about 0.4 length of propodus, spatulate, expanded, wider than propodus in dorsal view, with two rows of setae (or tufts of setae) along ridges. Fourth pereiopod generally similar to third pereiopod, shorter and slenderer. Fifth pereiopod much slenderer than third and fourth pereiopods; ischium usually unarmed; propodus with three small spiniform setae on ventral margin and one spiniform seta close to dactylus, and well-developed setal brush, latter extending to proximal half of propodus; dactylus subspatulate, less expanded compared to that of third or fourth pereiopod, about 0.35 length of propodus. Male second pleopod with appendix masculina almost as long as appendix interna. Uropod with each protopodal lobe ending in subacute tooth distally; exopod and endopod broad, ovate; diaeresis of exopod complete, broadly concave towards posterior end, with broadly subtriangular lobe near small distolateral spiniform seta and adjacent distolateral tooth; lateral margin of exopod broadly convex. Eggs numerous (> 100 in larger females), small; egg diameter about 0.6 mm. Colour pattern. Body (carapace and pleon) ranging from uniform olive-brown to greenish blue, without transverse bands; orbital hoods conspicuously whitish translucent, contrasting with dark frontal area of carapace; antennules and antennae uniform brown to blue-green; flagella pale brownish to greenish; mesial face of major chela largely dark olive-brown or olive-green; lateral face of major chela palm whitish with blue markings, especially bright along lateral longitudinal depression, and with olive-brown area dorsally and orange area distally; distolateral portion of pollex orange; lateral face of dactylus orange with some olive tinge; distal portion of both fingers greypurplish; plunger bright red-orange; minor chela olive-brown or brown-green; second to fifth pereiopods reddish or reddish green; pleopods greenish; telson and uropods dark uniform olive-brown to greenish blue; uropodal exopod with dusky or dark blue area posterior to transverse suture (Figs. 32, 33, 35A). Type locality. Bohol, Philippines (collection locality of the holotype and several paratypes). Distribution. Tropical Indo-West Pacific from Thailand to northern Australia (Fig. 52A); specific localities include the Philippines (e.g., Bohol, Panglao, Cebu, Luzon, Mindanao, Mactan), Singapore (Jurong), Thailand (Ranong), Indonesia (Sulawesi, Papua, record of A. cf. takla sp. nov. from Java), Papua New Guinea (Madang), Australia (Northern Territory, Queensland). Etymology. The specific epithet refers to “takla”—the local Tagalog name of this large and conspicuous snapping shrimp harvested for food in rural areas of Bohol, the type locality of the new species. Used as a noun in apposition. Common names proposed. Takla; giant snapping shrimp. Ecology and biology. Alpheus takla sp. nov. lives in spacious, deep burrows built in very soft, soggy mud among mangrove roots (Fig. 35B–F). Each burrow is typically occupied by a male-female pair. The species appears to be adapted to marine to slightly brackish conditions, which are typical for mangrove forests and creeks. The large number of eggs and their small diameter suggest an extended larval development in A. takla sp. nov. Alpheus takla sp. nov. appears to be the largest snapping shrimp in the world, with most specimens examined in this study exceeding 60 mm tl and largest individuals reaching almost 90 mm tl (Figs. 32H, 33). In the largest specimens, the impressive major chela typically exceeds 45 mm in length (palm + fingers) and in one ovigerous female from Thail and (86 mm tl, ZRC 2014.0723), its length reaches 52 mm. The species also has an unusually large, stout, strongly calcified ocellar beak (bec ocellaire), protruding between the eyestalks. Using quadrat sampling in the mangroves of Bohol, Philippines, Baobao et al. (2015) determined the mean density of A. takla sp. nov. (as Alpheus sp.) at seven individuals per m 2; the body weight of the shrimps ranged from 3.87 g to 12.86 g; and the total length from 47.8 mm to 74.4 mm (although, as noted above, the maximal total length of the species is closer to 90 mm). In dense mangroves of northern Queensland, Australia, the burrows of A. takla sp. nov. and A. malabaricus (sensu lato, see below) are sometimes inhabited by small gobies (Fig. 35E). In Bohol, Panglao and probably some other parts of the Philippines, A. takla sp. nov. is known as “takla” (see etymology) and is harvested for consumption by local fishermen or “gatherers” (Baobao et al. 2015; Aaron-Cabrillos 2019, misidentified as A. macellarius Chace, 1988; see also Fig. 32). Many gatherers use small crabs known in Bohol as “asan” [Parasesarma cf. eumolpe (De Man, 1895), Sesarmidae] as bait to lure the taklas from their burrows (M. Gabe, pers. comm.). The usual technique is to remove all crab’s appendages, tie it to an abaca (Musa textilis Née) fiber, drop the bait on the fiber into the takla’s burrow (filled with water), wait until the shrimp comes up and grasps the bait and then pull quickly and with force and try to secure the shrimp with the hands. Trying to defend itself, the takla may deliver a powerful and painful snap. The harvested taklas are then cooked in Sinigang broth with vegetables or in coconut cream sauce (Fig. 32C–E); see also links to two videos showing the takla harvest and preparation in Calape, Bohol, at the end of the reference section). According to Aaron-Cabrillos (2019), the bait used for collecting taklas in Clarin, Bohol, is a “hermit crab”. In 2014, the average daily catch per gatherer in Bohol was 75 shrimps, which is significantly lower compared to the average catch in the 1980–1990s, ranging from 150 to 300 shrimps per day (Baobao et al. 2015). The possible causes of the population decline of A. takla sp. nov. in Bohol identified by Baobao et al. (2015) are the unregulated gathering of snapping shrimps and general loss of mangrove swamps due to fishpond conversion and commercial firewood production. It is possible that the ecologically and morphologically very similar A. eurydactylus is also harvested for food in the Philippines, together with A. takla sp. nov. Taxonomic remarks. The Singaporean material reported as A. microrhynchus by Johnson (1962, 1965) contained at least two species: the larger specimens (tl> 60 mm) were reidentified as A. takla sp. nov., whereas the smaller ones were A. eurydactylus. At least some of the previous records of A. euphrosyne or A. euphrosyne euphrosyne (e.g., Banner & Banner 1966, 1982; Chace 1988) may be referable to A. takla sp. nov. For instance, the illustrations of a 78 mm male of A. euphrosyne euphrosyne from Australia in Banner & Banner (1982: fig. 73, except 73i) probably represents A. takla sp. nov., although the entire Australian material examined by these authors may also contain A. eurydactylus and possibly other species. Therefore, three large specimens from Gladstone (Queensland, Australia, NTM Cr-002639) identified by A.H. Banner & D.M. Banner as A. euphrosyne euphrosyne were reexamined by the present author and confirmed as A. takla sp. nov. (see also Fig. 35A). The two females from Jangkar, Java, and Makassar, Indonesia, reported by De Man (1911) under A.microrhynchus (deposited in RMNH under ZMA.Crus.D.202791 and ZMA Crus.D.202792, respectively), were photographed and briefly examined at the author’s request by Charles H.J.M. Fransen (RMNH). The smaller female from Jangkar (ZMA.Crus.D.202791) is generally similar to A. takla sp. nov., but differs from it mainly by the unarmed antennal basicerite, as noted by De Man (1911), and the presence of a distinct mesial transverse ridge on the major chela palm, similar to that of A. cyanoteles and A. paludicola (Yeo & Ng 1996). In addition, in this specimen, the dorsal surface of the telson is smooth, without a trace of median depression; the mesial surface of the major chela, including the distomesial angle of the pollex, presents no granulation; and the rostrum is longer than the rostrum of the syntype of A. microrhynchus (Fig. 2A) and more similar to that of A. takla sp. nov. (Fig. 30A). The proportions of the carpal subarticles of the second pereiopod and the broadened spatulate dactylus of the third and fourth pereiopods do not allow to identify this specimen as A. microrhynchus sensu De Man (1897), whereas the absence of granulation on the major chela excludes it from A. eurydactylus. Therefore, it is here tentatively assigned to A. cf. takla sp. nov., awaiting collection of fresh specimens throughout central Indonesia, especially males. On the other hand, the larger female from Makassar (ZMA Crus.D.202792) can be more reliably assigned to A. takla sp. nov., based on the well-developed rostrum (with a blunt carina); the shallow but distinct adrostral furrows; the prese, Published as part of Anker, Arthur, 2023, Revision of Alpheus euphrosyne De Man, 1897 and A. microrhynchus De Man, 1897, with description of three new species and taxonomic remarks on several other morphologically and ecologically similar snapping shrimps (Malacostraca: Decapoda: Alpheidae), pp. 1-115 in Zootaxa 5282 (1) on pages 57-67, DOI: 10.11646/zootaxa.5282.1.1, http://zenodo.org/record/7912292, {"references":["Johnson, D. S. (1965) A review of brackish water prawns of Malaya. Bulletin of the National Museum of Singapore, 33, 7 - 11.","De Man, J. G. (1897) Bericht u ¨ ber die von Herrn Schiffscapitan Storm zu Atjeh, an den westlichen Ku ¨ sten von Malakka, Borneo und Celebes sowie in der Java-See gesammelten Decapoden und Stomatopoden. Fu ¨ nfter Theil. Z oologische Jahrbu ¨ cher. Abtheilung fu ¨ r Systematik, Geographie und Biologie der Thiere, 9, 725 - 790, pls. 34 - 37 (in volume 10).","Banner, D. M. & Banner, A. H. (1982) The alpheid shrimp of Australia. Part III: The remaining alpheids, principally the genus Alpheus and the family Ogyrididae. Records of the Australian Museum, 34, 1 - 357. https: // doi. org / 10.3853 / j. 0067 - 1975.34.1982.434","Chace, F. A. Jr. (1988) The caridean shrimps (Crustacea: Decapoda) of the Albatross Philippine Expedition, 1907 - 1910, Part 5: Family Alpheidae. Smithsonian Contributions to Zoology, 466, 1 - 99. https: // doi. org / 10.5479 / si. 19436696.391.1","Banner, A. H. & Banner, D. M. (1966) The alpheid shrimp of Thailand. The Siam Society Monograph Series, 3, 1 - 168.","Wada, K. (2013) Benthic animals in mangrove swamp: a review. Global Environmental Research, 17 / 2013, 183 - 188.","Johnson, D. S. (1962) A synopsis of the Decapoda Caridea and Stenopodidea of Singapore, with notes on their distribution and a key to the genera of Caridea occuring in Malayan waters. Bulletin of the National Museum of Singapore, 30, 44 - 79, pl. 2.","Aaron-Cabrillos, J. L. (2019) Density, biomass and distribution of Alpheus macellarius (Chace, 1988) in Clarin, Bohol. International Journal of Agriuculture and Biological Sciences, November / December 2019, 1 - 33.","Baobao, J. G., Rabia, M. D. S. & Rulida, E. C. (2015) Preliminary assessment of the abundance and fishery of snapping shrimp (Alpheus sp.) in Calape, Bohol, Philippines. In: Romana-Eguia, M. R. R., Parado-Estepa, F. D., Salayo, N. D. & Lebata- Ramos, M. J. H. (Eds.), Resource Enhancement and Sustainable Aquaculture Practices in Southeast Asia: Challenges in Responsible Production of Aquatic Species: Proceedings of the International Workshop on Resource Enhancement and Sustainable Aquaculture Practices in Southeast Asia 2014 (RESA). Aquaculture Department, Southeast Asian Fisheries Development Center, Tigbauan, Iloilo, pp. 337. [abstract]","De Man, J. G. (1895) [1895 - 1898). Bericht ¸ ber die von Herrn Schiffscapitan Storm zu Atjeh, an den westlichen K ¸ sten von Malakka, Borneo und Celebes sowie in der Java-See gesammelten Decapoden und Stomatopoden. Zoologische Jahrb ¸ cher, Abtheilung f ¸ r Systematik, Geographie und Biologie der Thiere, 8 - 10, 485 - 609, 75 - 218 + 339 - 386 + 459 - 514 + 725 - 790 + pls. 12 - 14, 677 - 708 + pls. 28 - 38.","De Man, J. G. (1911) The Decapoda of the Siboga Expedition. Part II. Family Alpheidae. Siboga-Expeditie, 39 a 1 (2), 133 - 465; (1915) Supplement - Explanations of plates of Alpheidae, pls. 1 - 23. E. J. Brill, Leiden. https: // doi. org / 10.5962 / bhl. title. 10512","Yeo, D. C. J. & Ng, P. K. L. (1996) A new species of freshwater snapping shrimp, Alpheus cyanoteles (Decapoda: Caridea: Alpheidae) from Peninsular Malaysia and a redescription of Alpheus paludicola Kemp, 1915. Raffles Bulletin of Zoology, 44, 37 - 63."]}

Alpheus microrhynchus De Man, 1897 (Figs. 2A–E, 19–22, 51D) Alpheus microrhynchus De Man 1897: 754, pl. 36, fig. 65 (as Alpheus sp.). Alpheus microrhynchus. — De Man 1898: 318, pl. 4, fig. 3; Nobili 1900: 479; Johnson 1965: 9 (part.?); Banner & Banner 1966: 133 (part.?), fig. 50; Johnson 1976: 39 (part.?); Banner & Banner 1985: 24 (footnote); Naiyanetr 2007: 173. Crangon microrhynchus.— Suvatti 1937: 48. (?) Alpheus paludicola. — Kemp 1918: 273 (not A. paludicola Kemp, 1915). Not Alpheus microrhynchus. — De Man 1911: 413 (part.), pl. 23, fig. 99 [= A. takla sp. nov.]; Johnson 1965: 9 (part.); Johnson 1976: 39 (part.?) [possibly = A. takla sp. nov. and/or A. mangalis sp. nov.]; Yeo & Ng 1996: 56, fig. 8a, b [= A. takla sp. nov.]. Type material. Neotype, male (cl 16.3 mm, tl 37.0 mm, chl 21.0 mm), ZRC 2000.2193, Indonesia, Kalimantan Barat, Kabupaten Sambas, Sambas Pasar, fish market, leg. H.H. Tan, 18.04.1998 [THH-9833]. Additional material. Thailand: 1 male (cl 15.0 mm), 1 female (cl 14.9 mm, in poor condition), OUMNH. ZC. 2019.06.49, Bangkok, aquarium trade market, leg. P.K.L. Ng, 06.2001; 1 male (cl 13.4, in poor condition), 1 male (16.8 mm, tl 45.0 mm, chl 28.0 mm), MNHN-IU-2018-5739, Bangkok, aquarium trade market, leg. F. Fasquel, 06.2001; 3 males (cl 12.6–13.0 mm), 1 ov. female (13.1 mm), ZRC 2012.0399, between Bangkok and Chonburi, river mouth, local collectors, leg. P.K.L. Ng, 04.2012. Vietnam: 1 male (cl 17.1 mm), MNHN-IU-2013-14789, near Ho Chi Minh City, mangrove, water brackish (“very slightly salty”), in burrow in mud, leg. N. Ngoc-Ho, 05.1993. Malaysia: 2 females (cl 14.3, 14.7 mm), ZRC 2014.0670, Sarawak, Kuching prawn farm, no further data. Tentative identification. Alpheus cf. microrhynchus. Thailand: 1 male (cl 7.4 mm), 1 female (cl 8.7 mm), USNM 65477 /1, Bang Pakong River, leg. H. Smith, 01.07.1923. Redescription. See Figs. 19, 20 and Banner & Banner (1966: fig. 50). Medium-sized species of Alpheus (maximal cl 16.8 mm, tl ~45.0 mm). Carapace glabrous, minutely and sparsely pitted, without pubescence, with weak grooves. Rostrum very short, subtriangular, broader than long, narrowing towards apex, distally subacute, not reaching 0.3 length of first article of antennular peduncle, obliquely ascendant or pointing straight-forward (see remarks below); rostral carina present as faint broad elevation flattening posterior to orbital hoods; rostro-orbital furrows very shallow; anterior margin of orbital hoods very shallowly concave, almost perpendicular to rostrum. Pterygostomial angle broadly rounded; cardiac notch deep. Pleon smooth, finely pitted, without pubescence. Telson broad, ovate-rectangular, gently tapering towards posterior margin, about 1.3 times as long as maximal width near proximal margin; lateral margins with weak convexity at about mid-length; dorsal surface finely pitted, without longitudinal median depression, with two pairs of stout spiniform setae inserted in deep pits at some distance from lateral margins, first pair at about telson mid-length, second pair at about 0.7 of telson length; posterior margin broadly rounded, with two pairs of minute spiniform setae at posterolateral angles, lateral shorter than mesial. Antennular peduncle with stylocerite slightly convex laterally, with acute tip, latter not overreaching distal margin of first article; ventromesial carina with broadly triangular tooth, latter longer than high and without acute point; second article about 1.8–1.9 times as long as wide. Antennal peduncle with distoventral margin of basicerite unarmed or with minute tooth; carpocerite reaching slightly beyond scaphocerite; scaphocerite with lateral margin nearly straight; blade broad, separated from broad strong distolateral tooth by deep cleft; anterior margin of blade rounded and reaching slightly beyond distolateral tooth. Third maxilliped with antepenultimate article not particularly broadened, rather slender; penultimate article almost 3.2 times as long as wide, with long setae on lateroventral and mesioventral margins, latter not surpassing ultimate article; ultimate article with dense transverse rows of short serrulate setae on mesial surface and some much longer setae on dorsal and lateral surfaces, especially near or on apex; coxal lateral plate bluntly protruding dorsally; exopod reaching end of antepenultimate article. Major cheliped of A. edwardsii - type. Merus very stout, about 1.8 as long as wide; ventromesial margin unarmed distally. Carpus very short, cup-shaped. Chela large, massive, with fingers about 0.6–0.7 length of palm. Mesial face of palm with feebly developed mesial transverse ridge in proximal half, extending from proximal edge of subtriangular mesial longitudinal groove to ventral surface of palm; surface smooth, without granules. Lateral face of palm smooth, without granulation; lateral longitudinal groove moderately deep, subrectangular; proximal half of palm without transverse ridge. Dorsal shoulder rounded, sloping gently with angle inferior to 45° into broad transverse groove, not overhanging. Ventral shoulder pronounced, slightly protruding in lateral view, rounded. Pollex with mesial face smooth over most of its surface, with prominent mesial subdistal ridge marked by low protuberance proximally and extending to subdistal angle of pollex; area ventral to mesial subdistal ridge noticeably depressed; distomesial angle distinctly superior to 90°, blunt; lateral face smooth; distolateral angle about 90°, rounded. Dactylus slightly longer than pollex, with moderately developed dorsal ridge, latter twisted mesially, rounded, not protruding; both mesial and lateral surfaces smooth; dactylar plunger moderately large, stout, clearly distinct from ventral dactylar margin; distal surface of plunger with obliquely flattened ventromesial area, without deep transverse groove. Adhesive discs small. Male minor cheliped with chela strongly balaeniceps.Merus somewhat slenderer than that of major cheliped,about 2.5 times as long as wide; ventral margin with distomesial angle unarmed. Chela moderately swollen, with fingers about 0.9 length of palm; all surfaces smooth, without granulation. Palm subcylindrical, slightly swollen; mesial face with subdistal swelling; mesial longitudinal and lateral longitudinal groove indistinct or poorly demarcated; dorsal shoulder absent; dorsal transverse groove noticeable as shallow sinus; ventral transverse groove reduced to shallow sinus; ventral shoulder poorly marked as low bump or indistinct. Pollex with row of balaeniceps setae on low crest extending from base to just before mid-length of pollex; lateral surface with similar row of balaeniceps setae as on mesial surface. Dactylus with moderate lateral expansion (balaeniceps expansion) in proximal half, about twice as long as maximal width; lateral and mesial dactylar ridges each with dense rows of balaeniceps setae. Cutting edges of both fingers shallowly excavated on each side of sharp, blade-like ridge. Female minor cheliped simple, not balaeniceps. Chela slender, not particularly swollen, with fingers of about same length as palm. Palm surface smooth, without granulation, without noticeable sculpture, i.e., without mesial and lateral longitudinal grooves, and without shoulders. Fingers simple, with sharp cutting edges, not gaping when closed; fingertips curved and crossing. Second pereiopod with ratio of carpal subarticles ranging from 4: 1.7: 1: 1: 1.5 to 5: 1.8: 1: 1: 1.6. Third pereiopod moderately robust; ischium armed with small spiniform seta on lateroventral surface; merus about four times as long as maximal width, unarmed; propodus with about five spiniform setae on ventral margin and one pair of spiniform setae on distoventral margin adjacent to dactylus; dactylus close to 0.3 of propodus length, trigonal-conical, tapering, not broader than propodus in dorsal view. Fourth pereiopod generally similar to third pereiopod, slenderer; merus about 4.8 times as long as maximal width; dactylus about 0.3 length of propodus. Fifth pereiopod much slenderer than third and fourth pereiopods; ischium unarmed; propodus with at least two slender spiniform setae (or robust setae) and well-developed setal brush; dactylus trigonal-subspatulate, about 0.3 length of propodus. Male second pleopod with appendix masculina ~1.2 length of appendix interna. Uropod with each protopodal lobe ending in sharp tooth distally; exopod and endopod broad, ovate; diaeresis of exopod complete, almost straight, with two subacute teeth fringing small distolateral spiniform seta (occasionally two spiniform setae); lateral margin convex. Eggs not numerous (10 in one ovigerous female examined, ZRC 2012.0399, less than 40 in ovigerous females in Fig. 21A, D), very large; egg diameter about 1.4 mm. Colour pattern. Carapace pale olive-brown, sometimes with purplish or bluish tinge; posterior margin of carapace and each pleonite with one narrow, transverse, dark grey-blue or dusky band; antennular peduncles and scaphocerite pale brown-green or greenish blue; antennular and antennal flagella bluish or greenish; mesial face of major chela olive-green to bluish green or bluish grey, with some olive-beige, pale orange or yellowish areas; fingers pinkish distally; minor chela dark olive-grey to brown or bluish green; walking legs reddish; telson greyish blue distally; uropods mostly greyish blue with some reddish tinge; exopod with conspicuous brick-red patch above distolateral spiniform seta; area posterior to transverse suture intense deep blue (Figs. 21, 22). Type locality. Pontianak, Kalimantan (west coast of Borneo). Neotype from Kabupanten Sambas, Kalimantan Barat (west coast of Borneo), Indonesia. Distribution. South-East Asia (Fig. 51D): Thailand (vicinity of Bangkok and possibly Songkhla Lake and Bang Pakong), Indonesian and Malaysian parts of Borneo (Kalimantan Barat, Sarawak) (De Man 1897, 1898; Nobili 1900; present study); records from eastern Java, Indonesia (De Man 1911), and Singapore (Johnson 1965, 1976) require confirmation. Common name proposed. Klang snapping shrimp. Ecology and biology. Alpheus microrhynchus can be qualified as an stenohaline snapping shrimp, confined to oligohaline habitats, such as rivers, creeks, lagoons and artificial canals (man-made for navigation or irrigation of rice fields) under slight tidal influence, living in deep burrows dug into muddy bottom or riverbanks. The small number of eggs and their large size suggest a highly abbreviated larval development in this species (H. Coutière in De Man 1898), although it remains to be studied in more details. Alpheus microrhynchus is occasionally sold on fish or pet markets, mainly in Thailand (e.g., Chatuchak market in Bangkok). According to observations of F. Fasquel (pers. comm.), the species needs a low percentage of salt added to freshwater since all shrimps kept in pure freshwater unexpectedly died after only a few weeks. Taxonomic remarks. The neotype of A. microrhynchus from western Borneo corresponds to De Man’s (1897) description in all but a few minor details. The first small discrepancy is the direction of the rostrum, which was described as pointing obliquely upwards (= “schräg nach oben gerichtetes Spitzchen”). In most examined specimens of A. microrhynchus, including the neotype, the rostrum is directed straight forwards (Fig. 20A), although in one male specimen from Thailand (ZRC 2012.0399), it is directed slightly obliquely upwards. In addition, a clearly obliquely upwards-pointing rostrum can be seen in the illustrated (but not preserved) specimen from Thailand (Fig. 21A). Some variation in the direction of the rostrum is also present in the closely related A. cyanoteles (see below). The second difference is the armature of the antennal basicerite, which according to De Man (1897) has a rudimentary, hardly 1.0 mm long tooth (= “rudimentäres, kaum mm langes Stachelchen”). In the neotype of A. microrhynchus from Kalimantan and all specimens from Thailand, the distoventral margin of the basicerite is smoothly rounded, i.e., has no tooth, exactly as in the material from Thailand reported by De Man (1898) and Banner & Banner (1966). Because of its small size, this tooth may break off or get blunter after each moult, to the point of eventually becoming obsolete in full-grown adults. However, the development of the tooth on the basicerite may be simply variable in A. microrhynchus. Noteworthy here is that a similar variation in the armature of the basicerite was observed in A. euphrosyne (see above). Two further characters of A. microrhynchus, viz. the proportions of the carpal subarticles of the second pereiopod (P2) and the shape of the dactyli of the third to fifth pereiopods need to be discussed in more detail. In the original description of A. microrhynchus (material from Pontianak, as Alpheus sp.), De Man (1897) described the proportions of the carpal subarticles of the second pereiopod as 3.5: 1.3: 0.6: 0.6: 0.96, which is roughly similar to his observations (De Man 1898) of the material from Bangkok, 4.3: 1.6: 0.8: 0.72: 1.04 (right P2), and 4.0: 1.5: 0.8: 0.72: 1.1 (left P2). These numbers show that the ratio of the first to second subarticle is proportionally higher in the material from Thailand, 2.7–3.0, as illustrated by Banner & Banner (1966: fig. 50F), compared to the material from Borneo, where this ratio is at most 2.4. Regarding the condition of the dactylus of the ambulatory pereiopods, De Man (1897: 754) stated that the third to fifth pereiopods are somewhat similar to those of A. euphrosyne (“gleichen ungefähr denen von A. euphrosyne ”). No specific details on the dactylus were provided in the complementary description, where De Man (1898: 321) only noted that the ambulatory pereiopods agree with those of the material from Pontianak. However, Banner & Banner (1966) noted that the dactylus of A. microrhynchus, at least in the material from Thailand, is not spatulate, as in A. euphrosyne, but rather “conical, tapering”. Yeo & Ng (1996) examined several specimens identified as A. microrhynchus in the ZRC holdings and concluded that the dactylus in this species is “subconical to subspatulate”, which is essentially the same as trigonal-subspatulate in the above description. Alpheus microrhynchus can be separated from A. euphrosyne, A. eurydactylus and A. richardsoni by the ischium of the third and fourth pereiopods unarmed (vs. usually armed with a spiniform seta in the other three species); the much larger size of eggs, with the diameter around 1.4 mm (vs. equal to or less than 0.7 mm in the other species); the less expanded, trigonal-conical (at most weakly subspatulate) dactyli of the third to fifth pereiopods (vs. spatulate, much more expanded, in the other species; see also below); the perfectly smooth mesial surface of the major chela (vs. with a small distal field of granules in A. richardsoni and with large granulated areas in A. euphrosyne and A. eurydactylus); and the absence of a median longitudinal depression on the dorsal surface of the telson (present in the other species, especially well demarcated in A. euphrosyne and A. eurydactylus). In addition, A. microrhynchus differs from A. eurydactylus and A. richardsoni by the much shorter rostrum; from A. euphrosyne and A. eurydactylus by the absence of dorsal and ventral transverse grooves on the palm of the male minor chela (well developed in A. euphrosyne and A. eurydactylus); specifically from A. euphrosyne by the dorsal shoulder of the major chela smoothly sloping into the adjacent transverse groove (vs. protruding and overhanging the transverse groove in A. euphrosyne); and specifically from A. richardsoni by the much stronger distomesial ridge on the mesial face of the major chela pollex (which is feebly developed in A. richardsoni). In life, A. microrhynchus (Figs. 21, 22) can be distinguished from A. euphrosyne (Fig. 8–10) and A. richardsoni (Fig. 17, 18) by its colour pattern, which is more similar to that of A. eurydactylus (Fig. 16). There also seem to be an ecological separation between A. microrhynchus, A. euphrosyne, A. eurydactylus and A. richardsoni, with only the former species being confined to oligohaline habitats (sections of rivers, lagoons and canals under weak tidal influence). For separation of A. microrhynchus from other species treated in this study see below or refer to Table 1. Kemp’s (1918) record of A. paludicola from Thale Sap (Songkhla Lake) in southern Thailand most likely refers to A. microrhynchus. Kemp noted that the Thale Sap specimens had a distinctly longer rostrum, “extending considerably beyond the end of the orbital hoods”. They also had a different colour pattern, described by the collector, Dr. N. Annandale, as “translucent, without definite markings, but tinged... with reddish brown,... palm and fingers of both chelae were deeply tinged with blue, especially on the dorsal surface... eggs were pale green”. All females had very large eggs, 1.3–1.4 mm in diameter. This combination of characters agrees more with the diagnosis of A. microrhynchus than with that of A. paludicola, although in the latter species, females bear equally large eggs (Kemp 1915). Johnson’s (1965, 1976) records of A. microrhynchus from Singapore are probably based on material containing at least two species. All large-sized specimens (tl> 50 mm) identified as A. microrhynchus by D.S. Johnson were herein reidentified as either A. eurydactylus or A. takla sp. nov. (see below), whereas two smaller problematic females (tlA. cf. eurydactylus B. (see above). Although the historical occurrence of A. microrhynchus in Singapore cannot be excluded, it is rather unlikely that the species occurs there today, after most of transitional freshwater-to-brackish habitats were lost to urban development. Two specimens from Bang Pakong River east of Bangkok, Thailand (USNM 65477/1), were tentatively identified as A. cf. microrhynchus, because they deviate even more from the present redescription of the species. The third pereiopod merus of these specimens is somewhat broader, reaching the proportions of its homolog in A. eurydactylus. Most importantly, the minor chela of the male specimen has a more broadened, balaeniceps-shaped dactylus, almost as broad as in A. eurydactylus. On the other hand, the major chela has no trace of granulation (which is present in A. eurydactylus) and its dorsal shoulder is sloping very gently into the adjacent transverse groove (vs. sloping with a stronger angle in A. eurydactylus). The antennal basicerite of these specimens is unarmed, as in A. eurydactylus, which contradicts De Man’s (1897) original diagnosis of A. mic, Published as part of Anker, Arthur, 2023, Revision of Alpheus euphrosyne De Man, 1897 and A. microrhynchus De Man, 1897, with description of three new species and taxonomic remarks on several other morphologically and ecologically similar snapping shrimps (Malacostraca: Decapoda: Alpheidae), pp. 1-115 in Zootaxa 5282 (1) on pages 38-43, DOI: 10.11646/zootaxa.5282.1.1, http://zenodo.org/record/7912292, {"references":["De Man, J. G. (1897) Bericht u ¨ ber die von Herrn Schiffscapitan Storm zu Atjeh, an den westlichen Ku ¨ sten von Malakka, Borneo und Celebes sowie in der Java-See gesammelten Decapoden und Stomatopoden. Fu ¨ nfter Theil. Z oologische Jahrbu ¨ cher. Abtheilung fu ¨ r Systematik, Geographie und Biologie der Thiere, 9, 725 - 790, pls. 34 - 37 (in volume 10).","De Man, J. G. (1898) Note sur quelques especes du genre Alpheus Fabr., appartenant a la section dont l' Alpheus Edwardsii est le representant. Memoires de la Societ Zoologique de France, 11, 309 - 325, pl. 4.","Nobili, G. (1900) Decapodi e stomatopodi Indo-Malesi. Annali del Museo Civico di Storia Naturale di Genova, 40, 473 - 523.","Johnson, D. S. (1965) A review of brackish water prawns of Malaya. Bulletin of the National Museum of Singapore, 33, 7 - 11.","Banner, A. H. & Banner, D. M. (1966) The alpheid shrimp of Thailand. The Siam Society Monograph Series, 3, 1 - 168.","Johnson, D. S. (1976) Prawns of the Malacca Straits and Singapore waters. Journal of the Marine Biological Association of India, 18, 1 - 54.","Banner, D. M. & Banner, A. H. (1985) The alpheid shrimp of Indonesia, based upon J. G. de Man's \" The Decapoda of the Siboga Expedition, Part II. Family Alpheidae \" (1911). Marine Research in Indonesia, 25, 1 - 79. https: // doi. org / 10.14203 / mri. v 25 i 0.431","Naiyanetr, P. (2007) Checklist of crustacean fauna in Thailand (Decapoda, Stomatopoda, Anostraca, Myodocopa and Isopoda). Office of Natural Resources and Environmental Policy and Planning, Bangkok, 196 pp.","Suvatti, C. (1937) A check-list of aquatic fauna of Siam (excluding fishes). Bureau of Fisheries, Bangkok, 116 pp.","Kemp, S. (1918) Crustacea Decapoda of the Inle Lake Basin. Records of the Indian Museum, 14, 81 - 102, pls. 24 + 25. https: // doi. org / 10.26515 / rzsi / v 14 / i 4 / 1918 / 163580","Kemp, S. (1915) Fauna of the Chilka Lake. No. 3. Crustacea Decapoda. Memoirs of the Indian Museum, 5, 201 - 325, pls. 12 - 13. https: // doi. org / 10.5962 / bhl. title. 10414","De Man, J. G. (1911) The Decapoda of the Siboga Expedition. Part II. Family Alpheidae. Siboga-Expeditie, 39 a 1 (2), 133 - 465; (1915) Supplement - Explanations of plates of Alpheidae, pls. 1 - 23. E. J. Brill, Leiden. https: // doi. org / 10.5962 / bhl. title. 10512","Yeo, D. C. J. & Ng, P. K. L. (1996) A new species of freshwater snapping shrimp, Alpheus cyanoteles (Decapoda: Caridea: Alpheidae) from Peninsular Malaysia and a redescription of Alpheus paludicola Kemp, 1915. Raffles Bulletin of Zoology, 44, 37 - 63."]}

Alpheus euphrosyne De Man, 1897 (Figs. 1, 4–10, 51A, 53A, lower row) Alpheus euphrosyne De Man 1897: 745, pl. 36, fig. 64; De Man 1898: 317, pl. 4, fig. 2; De Man 1911: 413; Ahmad 1957: 16; Banner & Banner 1966: 130 (part.?, not fig. 49); Thomas 1976: 667; Holthuis, 1980: 119 (part.?); (?) Becker 1996: 95; (?) Becker et al. 2000: 195; Naiyanetr 2007: 173 (part.?, referring to Banner & Banner 1966); Harikrishnan & Kurup 2008: 111, fig. 1 (part., specimen on the left photograph); Paras et al. 2019: 1695 (part., not description), figs. 1–3. Crangon euphrosyne.— Suvatti 1937: 48 (part.?). Alpheus euphrosyne euphrosyne. — Li et al. 2008, fig. 2A; Harikrishnan et al. 2010: 195. Alpheus euphrosyne sensu lato.— Anker & De Grave 2016: 355 [specimen dredged off the eastern Straits of Johor; most Singaporean specimens are A. mangalis sp. nov. and A. eurydactylus; at least one specimen is A. takla sp. nov., see below]. Not Alpheus euphrosyne. — Johnson 1976: 37 [= Alpheus spp., most probably A. eurydactylus De Man, 1920 or A. mangalis sp. nov.]; Frith et al. 1976: 18 [= Alpheus spp., most probably A. eurydactylus De Man, 1920 or A. takla sp. nov.]; Yeo & Ng 1996: 58 [= Alpheus spp., including A. eurydactylus and A. mangalis sp. nov., fig. 8d = A. mangalis sp. nov.]; Bruce & Coombes 1997: 321 [= Alpheus spp., most probably A. eurydactylus De Man, 1920 or A. mangalis sp. nov.]; Edgar 1997: 194 (colour photo only) [= Synalpheus sp.]; Shokita 2004: 82 [= Alpheus spp., most probably A. eurydactylus De Man, 1920 or A. takla sp. nov.]; Wada 2013: 184, fig. 5 [= Alpheus spp., most probably A. takla sp. nov.]; Paras et al. 2019: 1695 (part., text, not figs. 1–3) [= A. euphrosyne euphrosyne sensu Banner & Banner (1982), see below]; Sha et al. 2019: 87, fig. 2.48 [= Alpheus spp., possibly A. eurydactylus De Man, 1920, A. nomurai sp. nov. or other taxa, see below]. Not Alpheus euphrosyne euphrosyne.— (?) Banner & Banner 1981: 23, 92; Banner & Banner 1982: 232, fig. 73, tab. 5 (part.?); (?) Banner & Banner 1983: 30; Banner & Banner 1985: 16 (part.); Chace 1988: 27 (part.) [= Alpheus spp., possibly A. eurydactylus De Man, 1920, A. takla sp. nov. and A. mangalis sp. nov.]. Not Alpheus euphrosyne langi.— Banner & Banner 1982: 238 [= A. pontederiae de Rochebrune, 1883]. Alpheus tirmiziae Kazmi 1974: 170, figs. 1–4; Banner & Banner 1978: 233, fig. 2i–m; Sha et al. 2019: 122, fig. 2.67. Alpheus tirmizae (lap. cal.).— Kazmi & Kazmi 1979: 154. Type material. Neotype: male (cl 16.6 mm, tl 48.5 mm, chl 26.0 mm), RMNH Crus.D. 49669, Indonesia, Java, Surabaya, leg. P. Buitendijk, 02.1929. Additional material examined. Indonesia: 1 female (cl indet., missing both chelipeds), ZMA Crus.D. 202727, Siboga-Expeditie, Postillon Islands (Sulawesi Selatan), leg. Jacquin, no date; 1 male (cl indet.), 1 ov. female (cl indet., missing both chelipeds), ZMA Crus.D. 202726, Sumatra, Bagansi Api-Api, leg. J.D.F. Hardenberg, 01.1929, det. H. Balss; 1 female (cl 15.5 mm, missing both chelipeds), 1 ov. female (cl 16.2 mm, missing major cheliped, identification somewhat tentative), ZMA Crus.D. 202793, Nias, leg. Kleiweg de Zwaan, no date. China: 1 male (cl 14.3 mm), ZRC 2014.0677, Guangdong, Nanao County, Houzhai, fish market, leg. Y. Cai & N.K. Ng, 12.11.1998; 1 male (cl 23.0 mm), USNM 62106, locality not specified, don. S.F. Light; 1 male (cl 20.5 mm), USNM 62107, Fujian, “Santu north of Foochow” (possibly Shantou, south of Fuzhou), no date, don. S.F. Light; 1 female (cl 13.7 mm), USNM 62109, Fujian, Xiamen, Amoy, outer harbour, net trap, leg. & don. S.F. Light, 16.04.1923; 1 male (cl 17.5 mm), MCZ 1743, Hong Kong, leg. Capt. Putnam, 03.1861. Taiwan: 1 female (cl 10.0 mm), ZRC 1999.0723, Kaohsiung County, Tungkang, fish port, leg. P.K.L. Ng & K. Lim, 05.1999; 1 female (cl 13.8 mm), USNM 59044, Yensui River, leg. M. Maki, don. M. Oshima, 01.1923. Vietnam: 4 males (cl 12.5–18.0 mm), MNHN-IU-2018-5730, Ha Tien, Sau, depth: 20–30 m, leg. H. Duy, 21.08.1995. Cambodia: 1 male (cl 18.9 mm), ZRC 2013.0458, Kampot and Prek Romeas fish landings, leg. T. R. Roberts, 29.03.2006 – 03.04.2006. Thailand: 5 specimens, including 3 males (cl 16.0–16.9 mm), ZRC 2000.2150, Gulf of Thailand, off Khanom, trawl, fish port, leg. P.K.L. Ng, 12.1984; 1 male (cl 19.3 mm), MNHN-IU-2018-5752, same collection data as for previous specimens; 1 male (cl 14.0 mm), ZRC 2003.0619, Songkhla Lake (Thale Sap), Sakom crab landing, leg. P.K.L. Ng et al., 15– 17.10.2003; 1 male (cl indet.), ZRC 2000.2137, Gulf of Thailand, Siracha, fish market, leg. P.K.L. Ng, 02.2000; 3 males (cl 20.2–21.7 mm), 1 ov. female (cl 21.9 mm), ZRC 2000.0957, Gulf of Thailand, Chonburi, Angsila and Siracha, fish ports, leg. P.K.L. Ng et al., 20.02.2000; 3 males (cl 15.0–16.9 mm), OUMNH. ZC. 2019.06.42 [ex-ZRC 2000.0956], Gulf of Thailand, Chonburi, Angsila and Siracha, fish ports, leg. P.K.L. Ng et al., 20.02.2000;> 20 specimens of both sexes (cl indet.), ZRC 1998.1066, Gulf of Thailand, Chonburi, Angsila, fish port, leg. P.K.L. Ng, 25.09.1998; 1 male (cl 16.4 mm), 1 female (cl 18.0 mm), MNHN-IU-2018-5593 (ex-ZRC 1998.1066), same collection data as for previous specimens; 2 males, 1 female (cl indet.), RMNH Crus.D.42009, Gulf of Thailand, Chonburi, ca. 100 km SE of Bangkok, between Si Racha and Naklua, fishermen, 14.03.1985, leg. A.C.J. Burgers & L.B. Holthuis; 1 male, 1 female (cl indet.), RMNH Crus.D.42426, Gulf of Thailand, Surat Thani, off river mouth of Don Sak, depth: 4 m, 20.10.1992, leg. S. Chaitiamvong; 1 male (cl 21.9 mm), 1 female (cl 13.2 mm), MNHN-IU-2018-5751 (ex ZRC 1999.2006), Gulf of Thailand, Siracha, fish port, leg. P.K.L. Ng et al., 10.11.1999; 1 male (cl 16.1 mm), 1 female (cl 14.5 mm), ZRC 1999.2005, Gulf of Thailand, Angsila, fish port, leg. P.K.L. Ng et al., 10.11.1999; 2 males (cl 15.5 mm, 17.5 mm), 1 female (cl 16.0 mm), USNM 95528, Gulf of Thailand, off Lem Sing, 14.06.1926; 1 male (cl 21.5 mm), ZRC 2008.0600, Andaman Sea off Phuket, Phuket fish port, leg. P.K.L. Ng et al., 17.02.2001; 1 male (cl 20.2 mm), OUMNH. ZC. 2019.06.43 [ex-ZRC 1998.1152], Andaman Sea off Phuket, Phuket fish port, leg. S. Chiatiamuong et al., 12.1998. Malaysia: 1 male (cl indet., missing major cheliped), RMNH Crus.D. 49740, Penang, Kuala Jalan Baru, 02.06.1961, leg. E. R. Alfred; 1 female (cl 21.5 mm), MNHN-IU-2018-5594, Lumut, no further data, don. V. C. Chong, 1999. Singapore: 1 male (cl 23.6 mm), ZRC 2008.0632, Siglap (Bedok area), 06.1933; 1 male (cl 6.7 mm, missing both chelipeds), ZRC 2014.0671, south of Pulau Tekong, MPA grid 0422, depth: 11.9–12 m, mud, clay, rectangular dredge, leg. S.C. Lim, A. Anker & C.K. Chim, 25.03.2014. India: 1 male (cl indet.), RMNH Crus.D.49747, Kochi (Cochin), 28.03.1979, leg. J.C. Miquel; 1 female (cl indet.), RMNH Crus.D.49748, same collection data as for previous specimen. Pakistan. 1 female (cl 11.4 mm), USNM 63515, Pakistan, Kurrachee Bay (= Karachi), no further data [identified by H. Coutière as A. euphrosyne]. Redescription. See Figs. 4–7. Large-sized species of Alpheus (maximal cl 23.6 mm, tl ~72.0 mm). Carapace glabrous, finely pitted, especially around cardiac region, without pubescence, with slight grooves. Rostrum short, triangular, distally acute, reaching at most to mid-length of first article of antennular peduncle; rostral carina absent or present as low broad elevation; rostro-orbital furrows indistinct or extremely shallow. Pterygostomial angle broadly rounded; cardiac notch deep. Pleon smooth, finely and scarcely pitted, without pubescence. Telson very broad, ovate-rectangular, slightly tapering towards posterior margin, about 1.3–1.4 times as long as maximal width near proximal margin; lateral margins with shallow concavity on proximal half; dorsal surface finely pitted, with longitudinal median depression densely furnished with setae along its margins, and with two pairs of short spiniform setae inserted in deep pits at considerable distance from lateral margins, first pair near telson mid-length, second pair between 0.7 and 0.8 of telson length, and with numerous erect simple setae; posterior margin broadly rounded, with two pairs of small spiniform setae at posterolateral angles, lateral shorter than mesial (often broken or missing). Antennular peduncle with stylocerite broad, markedly convex laterally, with acute tip, latter not overreaching distal margin of first article; ventromesial carina with small, anteriorly directed, sharp tooth; second article about 2.2–2.5 times as long as wide. Antennal peduncle with distoventral margin of basicerite unarmed or with minute, blunt or subacute tooth; carpocerite not reaching beyond scaphocerite; scaphocerite with lateral margin almost straight; blade very broad, separated from broad distolateral tooth by deep cleft; anterior margin of blade broadly rounded and reaching somewhat beyond distolateral tooth. Third maxilliped with antepenultimate article somewhat broadened; penultimate article elongate, about three times as long as wide, with long setae on ventrolateral margin, some surpassing ultimate article; ultimate article with dense transverse rows of short serrulate setae on mesial surface and much longer setae on dorsal and lateral surfaces, especially near or on apex; coxal lateral plate bluntly protruding dorsally; exopod fleshy, broad, reaching end of antepenultimate article. Major cheliped of A. edwardsii - type. Merus stout, about twice as long as wide; ventromesial margin unarmed distally. Carpus very short, cup-shaped. Chela large, massive, with fingers about 0.6–0.7 length of palm; surface partly heavily granulated. Mesial face of palm with distinct mesial transverse ridge in proximal half, extending from proximal edge of subtriangular mesial longitudinal groove to ventral surface of palm, although not reaching it; large areas of mesial surface, including mesial longitudinal groove and part of mesial transverse ridge, densely covered with granules. Lateral face of palm mostly smooth, with relatively small field of granules present on low ridge between lateral longitudinal groove and dorsal edge of ventral shoulder; lateral longitudinal groove deep, subrectangular; proximal half with low transverse ridge. Dorsal shoulder overhanging transverse groove, i.e., distinctly protruding above groove in form of blunt tooth. Ventral shoulder pronounced, slightly protruding in lateral view, broadly rounded; surface covered with granules. Pollex with mesial face distinctly granulated over most of its surface; both mesial subdistal ridge and adjacent proximal protuberance prominent, extending subdistal angle of pollex; area ventral to mesial subdistal ridge conspicuously grooved; distomesial angle distinctly superior to 90°, blunt; lateral face distinctly granulated, except close to ventral edge; distolateral angle about 90°, broadly rounded. Dactylus with prominent dorsal ridge, latter distinctly twisted mesially and protruding proximally; mesial surface strongly granulated; lateral surface moderately granulated, mostly proximally so, with deep depression above plunger; dactylar plunger relatively short, with anterior margin gently sloping and merging into (i.e., almost in continuation with) ventral margin of dactylus; distal surface of plunger obliquely flattened, its mesial surface with deep transverse channel. Adhesive discs small. Male minor cheliped with chela strongly balaeniceps. Merus noticeably slenderer than that of major cheliped, about three times as long as wide; ventral margin with distomesial angle unarmed. Chela moderately swollen, slender in lateral view, with fingers about as long as palm. Palm subcylindrical, slightly compressed, feebly swollen; mesial face smooth proximally, distinctly granulated distally, with low bump-like elevation subdistally; mesial longitudinal groove well marked, deep; lateral face of palm with small field of granules distally, below lateral longitudinal groove; lateral longitudinal groove well marked, deep, with lower margin slightly elevated; dorsal shoulder pronounced, rounded, in form of blunt tooth slightly overhanging deep, broad dorsal transverse groove; ventral transverse groove moderately deep, bordered by distinct ventral shoulder on lateral surface. Pollex with mesial surfaces granulated proximally and with row of balaeniceps setae on low crest extending from base to mid-length of pollex; lateral surface mostly smooth, with similar row of balaeniceps setae as on mesial surface. Dactylus with well-developed proximal balaeniceps expansion, about 1.7–1.8 as long as maximal width; lateral and mesial dactylar ridges each with dense rows of balaeniceps setae. Cutting edges of both fingers shallowly excavated on each side of sharp, blade-like ridge. Female minor cheliped simple, not balaeniceps. Chela slender, not particularly swollen, with fingers 1.5–2.0 times as long as palm. Palm granulated, more on mesial than on lateral side, without mesial or lateral longitudinal grooves. Fingers simple, with sharp cutting edges, slightly gaping when closed; fingertips strongly curved and crossing. Second pereiopod with ratio of carpal subarticles approximately equal to 4: 3: 1: 1: 1.5. Third pereiopod moderately robust; ischium armed with short, stout spiniform seta on ventrolateral surface near proximal margin, sometimes unarmed; merus about 4.5 times as long as maximal width, unarmed; propodus with two to four more or less stout spiniform setae on ventral margin and one pair of spiniform setae on distoventral margin adjacent to dactylus; dactylus about 0.3 length of propodus, spatulate, conspicuously expanded, slightly wider than propodus in dorsal view, with dense row of setae and tufts of setae along median ridge. Fourth pereiopod generally similar to third pereiopod, slightly slenderer; merus about 5.0 times as long as maximal width; dactylus about 0.3 length of propodus. Fifth pereiopod much slenderer than third and fourth pereiopods; ischium unarmed; propodus with four slender spiniform setae and well-developed setal brush; dactylus spatulate, less expanded compared to that of third or fourth pereiopod, about 0.4 length of propodus. Male second pleopod with appendix masculina 0.6–0.8 length of appendix interna. Uropod with each protopodal lobe ending in sharp tooth distally; exopod and endopod broad, ovate; diaeresis of exopod complete, straight for most part, with small subacute or blunt tooth near small distolateral spiniform seta and adjacent distolateral tooth; lateral margin of exopod broadly convex. Eggs numerous (> 200 in large females), small; egg diameter less than 0.6 mm. Colour pattern. Body pale beige to greyish yellow; carapace with dark grey-blue anterolateral margin and two diffuse, dark brown, transverse bands, one across mid-length and one near posterior margin; each pleonite with dark brown to grey-brown band running transversely near posterior margin; antennular flagella bright blue (lateral) or reddish purple (mesial); antennal flagella bright blue; mesial face of major chela olive-green with large pale yellow or pale brown-orange areas, and two large, more or less conspicuous, violet-blue patches on palm; fingers distally pale orange-pink; dactylar plunger with whitish blue area; minor chela pale olive-green, darker and with purplish tinge on fingers; walking legs pink to reddish; pleopods reddish; telson pale greenish yellow, darker posteriorly; uropods pale yellow proximally, dark blue distally (Figs. 8–10; see also black-and-white photograph in Li et al. 2008: fig. 2A, and colour photographs in Harikrishnan & Kurup 2008: fig. 1 and Paras et al. 2019: figs. 1–3, specimen with faded colours). Type locality. Java Sea (De Man 1897). Neotype from Surabaya, Java, Indonesia. Distribution. Tropical Indo-West Pacific from the northern Arabian Sea to South China Sea and Java Sea (Fig. 51A), including Pakistan (Karachi), India (Kochi, Vembanad Lake, Gulf of Mannar, Palk Bay), Bangladesh, Thailand (Andaman Sea: Phuket; Gulf of Thailand: Pattani, Angsila, Khanom, Surat Thani, Satun, Songkhla Lake), Cambodia (Kampot), Vietnam (Nhatrang Bay), Malaysia (Andaman Sea: Penang), Indonesia (Java, Sulawesi, Sumatra, Nias?), Singapore (Bedok, Pulau Tekong), southern China (Guangdong, Zhejiang, Hong Kong), and Taiwan (Yensui, Kaohsiung) (De Man 1897; Kazmi 1974; Banner & Banner 1978; Li et al. 2008; Harikrishnan et al. 2010; Paras et al. 2019; present study). Common names proposed. Euphrosyne snapping shrimp; De Man’s estuarine snapping shrimp. Ecology and biology. Alpheus euphrosyne appears to be mainly a marine / estuarine snapping shrimp associated with subtidal sandy, sandy-muddy and muddy bottoms.Therefore, it can be considered as a polyhaline and mesohaline species, typically encountered within or near large estuarine areas. However, A. euphrosyne is capable of tolerating very low salinities and even freshwater conditions, at least in laboratory experiments (Harikrishnan & Kurup 2008; Harikrishnan et al. 2010). The exact depth range of A. euphrosyne is unknown as most specimens examined herein came from crab/prawn/fish landings or fish markets. Others were dredged from a depth of 8.5–27 m (Thomas 1976; Banner & Banner 1978; Paras et al. 2019; present study) or collected with indigenous stake nets (Harikrishnan et al. 2010). However, the identity of A. euphrosyne in Harikrishnan & Kurup (2008) and Harikrishnan et al. (2010) needs confirmation (see below). In the lower Kochi (Cochin) estuary in southern India, Harikrishnan et al. (2010) observed a high preponderance of A. euphrosyne in post- and pre-monsoon months, i.e., in salinity conditions varying from polyhaline in pre-monsoon months to mesohaline (5–18 0 / 00) in post-monsoon months, suggesting an inward migration of primarily marine species to highly fertile estuarine areas for feeding and breeding. These authors also studied some aspects of the reproductive biology of A. euphrosyne, examining 1309 females with cl 8.5–21.5 mm. The large number of eggs (141–1553 eggs per brood, mean fecundity 480 272) and their small size (diameter range: 0.12–0.53 mm) suggest an extended larval development in this species. Alpheus euphosyne is one of the largest snapping shrimps known to date, with the carapace length (cl) reaching 22.6 mm and total body length (tl) up to 71 mm (Thomas 1976). The largest specimen in the present material is a female from Thailand (ZRC 2000.0957) with cl 22 mm and tl 66 mm, with several other adults exceeding cl 20 mm and tl 60 mm; in the largest specimens (Fig. 8), the major chela length (chl) overreaches 35 mm. Due to its large size, A. euphro, Published as part of Anker, Arthur, 2023, Revision of Alpheus euphrosyne De Man, 1897 and A. microrhynchus De Man, 1897, with description of three new species and taxonomic remarks on several other morphologically and ecologically similar snapping shrimps (Malacostraca: Decapoda: Alpheidae), pp. 1-115 in Zootaxa 5282 (1) on pages 12-23, DOI: 10.11646/zootaxa.5282.1.1, http://zenodo.org/record/7912292, {"references":["De Man, J. G. (1897) Bericht u ¨ ber die von Herrn Schiffscapitan Storm zu Atjeh, an den westlichen Ku ¨ sten von Malakka, Borneo und Celebes sowie in der Java-See gesammelten Decapoden und Stomatopoden. Fu ¨ nfter Theil. Z oologische Jahrbu ¨ cher. Abtheilung fu ¨ r Systematik, Geographie und Biologie der Thiere, 9, 725 - 790, pls. 34 - 37 (in volume 10).","De Man, J. G. (1898) Note sur quelques especes du genre Alpheus Fabr., appartenant a la section dont l' Alpheus Edwardsii est le representant. Memoires de la Societ Zoologique de France, 11, 309 - 325, pl. 4.","De Man, J. G. (1911) The Decapoda of the Siboga Expedition. Part II. Family Alpheidae. Siboga-Expeditie, 39 a 1 (2), 133 - 465; (1915) Supplement - Explanations of plates of Alpheidae, pls. 1 - 23. E. J. Brill, Leiden. https: // doi. org / 10.5962 / bhl. title. 10512","Ahmad, N. (1957) Prawn and prawn fishery of East Pakistan. Directorate of Fisheries, Government of East Pakistan, 31 pp.","Banner, A. H. & Banner, D. M. (1966) The alpheid shrimp of Thailand. The Siam Society Monograph Series, 3, 1 - 168.","Thomas, M. M. (1976) New records of four alpheid shrimp from the Indian waters. Journal of the Marine Biological Association of India, 18, 666 - 669.","Holthuis, L. B. (1980) FAO species catalogue. Vol. 1. Shrimps and prawns of the world. An annotated catalogue of species of interest to fisheries. FAO Fisheries Synopsis 125. FAO, Rome, xvii + 271 pp.","Becker, K. (1996) Epibionts on carapaces of some malacostracan crustaceans from the Gulf of Thailand. Journal of Crustacean Biology, 16, 92 - 104. https: // doi. org / 10.2307 / 1548934","Becker, K., Hormchong, T. & Wahl, M. (2000) Relevance of crustacean carapace wettability for fouling. Hydrobiologia, 426, 193 - 201. https: // doi. org / 10.1023 / A: 1003918512565","Naiyanetr, P. (2007) Checklist of crustacean fauna in Thailand (Decapoda, Stomatopoda, Anostraca, Myodocopa and Isopoda). Office of Natural Resources and Environmental Policy and Planning, Bangkok, 196 pp.","Harikrishnan, M. & Kurup, B. M. (2008) Potential of snapping shrimps as ornamentals. In: Kurup, B. M., Boopendranth, M. R., Ravindran, K., Sairabanu & Nair, A. G. (Eds.), Ornamental fish, breeding, farming and trade. Department of Fisheries, Government of Kerala, Kerala, pp. 1 - 280.","Paras, N. J., Purushothaman, P., Chinnadurai, S., Renjith R. K., Remesan, M. P., Baiju, M. V. & Edwin, L. (2019) Occurrence of Alpheus euphrosyne de Man, 1897 (Crustacea: Decapoda: Alpheidae) from the south-eastern Arabian Sea, India. Indian Journal of Geo Marine Sciences, 48, 1695 - 1698.","Suvatti, C. (1937) A check-list of aquatic fauna of Siam (excluding fishes). Bureau of Fisheries, Bangkok, 116 pp.","Li, X., Shen, A. & Xu, Z. (2008) Report on some shrimps from Dongtou Island, Zhejiang, China (Decapoda, Natantia). Crustaceana, 81, 1371 - 1330. https: // doi. org / 10.1163 / 156854008 X 369500","Harikrishnan, M., Unnikrishnan, U., Smija Maju, M., Reena Greeshma, A. R. & Madhusoodana Kurup, B. (2010) Size at sexual maturity, egg number and reproductive output of the snapping shrimp Alpheus euphrosyne euphrosyne De Man, 1897. Invertebrate Reproduction & Development, 54, 195 - 202. https: // doi. org / 10.1080 / 07924259.2010.9652333","Anker, A. & De Grave, S. (2016) An updated and annotated checklist of marine and brackish caridean shrimps of Singapore (Crustacea, Decapoda). Raffles Bulletin of Zoology, Supplement 34, 343 - 454.","Johnson, D. S. (1976) Prawns of the Malacca Straits and Singapore waters. Journal of the Marine Biological Association of India, 18, 1 - 54.","De Man, J. G. (1920) Diagnoses of some new species of Penaeidae and Alpheidae with remarks on two known species of the genus Penaeopsis A. M. - Edw. from the Indian Archipelago. Zoologische Mededelingen, Leiden, 5, 103 - 109.","Frith, J. W., Tantanasiriwong, R. & Bhatia, O. (1976) Zonation of macrofauna on a mangrove shore, Phuket Island, southern Thailand. Phuket Marine Biology Center Research Bulletin, 17, 1 - 14.","Yeo, D. C. J. & Ng, P. K. L. (1996) A new species of freshwater snapping shrimp, Alpheus cyanoteles (Decapoda: Caridea: Alpheidae) from Peninsular Malaysia and a redescription of Alpheus paludicola Kemp, 1915. Raffles Bulletin of Zoology, 44, 37 - 63.","Bruce, A. J. & Coombes, K. E. (1997) An annotated check-list of the caridean shrimps (Crustacea: Decapoda) of Darwin Harbour, with descriptions of three new species of Periclimenes [Palaemonidae: Pontoniinae]. In: Hanley, J. R., Caswell, G., Megirian, D. & Larson, H. K. (Eds.), Proceedings of the sixth international marine biological workshop. The marine fauna and flora of Darwin Harbour, Northern Territory, Australia. Museums and Art Galleries of the Northern Territory and the Australian Marine Sciences Association, Darwin, pp. 301 - 337.","Edgar, G. J. (1997) Australian marine life. The plants and animals of temperate waters. Reed, Victoria, 544 pp. [revised edition published in 2001]","Shokita, S. (2004) Chapter 5. The role of aquatic organisms in mangrove ecosystems. In: Vannucci, M. (Ed.), Mangrove management and conservation: present and future. United Nations University Press, Tokyo, pp. 76 - 110.","Wada, K. (2013) Benthic animals in mangrove swamp: a review. Global Environmental Research, 17 / 2013, 183 - 188.","Banner, D. M. & Banner, A. H. (1982) The alpheid shrimp of Australia. Part III: The remaining alpheids, principally the genus Alpheus and the family Ogyrididae. Records of the Australian Museum, 34, 1 - 357. https: // doi. org / 10.3853 / j. 0067 - 1975.34.1982.434","Sha, Z., Wang, Y. & Cui, D. (2019) The Alpheidae from China seas. Crustacea: Decapoda: Caridea. Springer, Singapore, 323 pp. https: // doi. org / 10.1007 / 978 - 981 - 15 - 0648 - 2 _ 2","Banner, D. M. & Banner, A. H. (1981) Annotated checklist of the alpheid shrimp of the Red Sea and Gulf of Aden. Zoologische Verhandelingen, Leiden, 190, 1 - 99.","Banner, A. H. & Banner, D. M. (1983) An annotated checklist of the alpheid shrimp from the western Indian Ocean. Travaux et Documents de l'ORSTOM, 158, 1 - 164.","Banner, D. M. & Banner, A. H. (1985) The alpheid shrimp of Indonesia, based upon J. G. de Man's \" The Decapoda of the Siboga Expedition, Part II. Family Alpheidae \" (1911). Marine Research in Indonesia, 25, 1 - 79. https: // doi. org / 10.14203 / mri. v 25 i 0.431","Chace, F. A. Jr. (1988) The caridean shrimps (Crustacea: Decapoda) of the Albatross Philippine Expedition, 1907 - 1910, Part 5: Family Alpheidae. Smithsonian Contributions to Zoology, 466, 1 - 99. https: // doi. org / 10.5479 / si. 19436696.391.1","Rochebrune, A. T. de (1883) Diagnoses d'arthropodes nouveaux propres la Senegambie. Bulletin de la Societe Philomathique de Paris, Serie 7, 7, 167 - 182.","Kazmi, M. A. (1974) A new species of the genus Alpheus from the Bay of Bengal (Decapoda, Caridea). Crustaceana, 27, 170 - 174. https: // doi. org / 10.1163 / 156854074 X 00389","Banner, D. M. & Banner, A. H. (1978) Annotated checklist of alpheid and ogyridid shrimp from the Philippine Archipelago and the South China Sea. Micronesica, 14, 215 - 257.","Kazmi, M. A. & Kazmi, Q. B. (1979) A check-list of marine caridean prawns of Pakistan. Biologia, 25, 151 - 157."]}

Alpheus eurydactylus De Man, 1920 (Figs. 3H, I, 11–16, 51B) [see also Fig. 53B, C for A. cf. eurydactylus] Alpheus eurydactylus De Man 1920: 109; De Man 1924: 48, fig. 17. Alpheus euphrosyne.— Banner & Banner 1966: 130 (part.?), fig. 49; Johnson 1976: 37 (part.) [not A. euphrosyne De Man, 1897]. (?) Alpheus euphrosyne euphrosyne.— Banner & Banner 1981: 23, 92 (part.?); Banner & Banner 1982: 232, fig. 73 (part.?); Banner & Banner 1983: 30 (part.?); Banner & Banner 1985: 16 (part.?); Chace 1988: 27 (part.?) [not A. euphrosyne De Man, 1897]. Alpheus cf. euphrosyne . — Marin 2021: 381, fig. 1a [not A. euphrosyne De Man, 1897]. (?) Alpheus sp. cf. audouini .— Johnson 1962: 53 (part.?) [not A. audouini Coutière, 1905]. Type material. Lectotype, male (cl 13.8 mm, tl ~40.0 mm, chl 21.0 mm), paralectotype, ov. female (cl 15.0 mm, tl ~44.0 mm, chl 21.5 mm), RMNH Crus.D.49670 = ZMA Crus.D.102536, Indonesia, Java, Surabaya, leg. P. Buitendijk, no date. Additional material examined. Indonesia: 1 male (cl 19.4 mm), MNHN-IU-2018-5738, Papua, Kamora, mangrove, leg. A. Darmawan et al., 07.08.2012; 1 female (cl 12.6 mm), 1 ov. female (cl 17.3 mm), OUMNH. ZC. 2019.06.44, Papua, Ajkwa Island, mangrove, leg. A. Darmawan et al., 20.07.2012; 1 female (cl 21.5 mm), ZMA Crus.D.202791, Siboga Expedition sta. 4, “Strand” (beach), no further details; 1 ov. female (cl indet.), RMNH Crus. D.49671, Madura Island, leg. P. Buitendijk, 03.1930; 2 males, 1 ov. female (cl indet.), RMNH Crus.D.49660, Java, Tanjong Priok (near Jakarta), leg. P. Buitendijk, 1907; 4 males, 1 ov. female (cl indet.), RMNH Crus.D.49666, Java, Tanjong Priok (near Jakarta), leg. P. Buitendijk, 07.1927; 1 ov. female (cl indet.), RMNH Crus.D.49667, Java, Probolinggo, leg. P. Buitendijk, 04.1928; 1 male, 2 females (cl indet.), RMNH Crus.D.49661, Sumatra, Pulau Weh, leg. P. Buitendijk, 1910; 1 male (cl indet.), RMNH Crus.D.49668, East Java, Passeroean, leg. P. Buitendijk, 1929; 1 male (cl indet.), RMNH Crus.D.49665, East Java, Panaroekan, leg. P. Buitendijk, 08.1927; 1 female (cl indet.), RMNH Crus.D.49408, West Java, Banten Baai, Pulau Pamudjan Besar, 5°56’S 106°12’E, “laboratorium voor Onderzoek der zee, Batavia”, no further details; 1 female (cl 21.3 mm), RMNH Crus.D.49664, Java, Surabaya, leg. P. Buitendijk, 02.1927; 3 chelipeds, RMNH Crus.D.49672, Java, Bay of Jakarta, Alkmaar Island, leg. P. Buitendijk, 1906. Malaysia: 1 male (cl 11.2 mm), 1 ov. female (cl 17.5 mm), OUMNH. ZC. 2019.06.45, Penang (mainland part), Nibong Tebal, mudflat, leg. Z. Jaafar, 22.11.2001; 2 males (cl 11.5, 15.2 mm), 2 females (cl 10.3, 11.7 mm), MNHN-IU-2018-5680, same collection data as for previous specimens; 1 male (cl 15.1 mm), MNHN-IU-2018-5599, Johor, Muar, mudflat, soggy mud, leg. N.K. Ng & J.C.Y. Lai, 10.02.2009; 1 ov. female (cl indet.), MNHN-IU-2018-5598, same collection data as for previous specimen; 1 male (cl 15.1 mm), 1 ov. female (cl 13.8 mm), ZRC 2009.0306, Terengganu, Telok Tebrau, mudflats, with gobies, leg. Z. Jaafar, 05.11.2002; 1 male (cl 13.9 mm), ZRC 1979.4.3.7, Selangor, Port Klang (Port Swettenham), mudflats, leg. S. Kumar, 27.08.1968; 1 male (cl 14.7 mm), MNHN-IU-2018-5595, Selangor, Port Klang, Taman Sri Sementa, leg. N.K. Ng & J.C.Y. Lai, 10.02.2009. Singapore: 1 male (cl 18.0 mm), ZRC 2000.1220, East Coast, mudflat, leg. P.K.L. Ng, 18.04.1996; 1 female (cl 10.6 mm), ZRC 1996.0463, Sungei Punggol, sta. D1, mudflat, leg. R.E.S.T., 28.06.1995; 1 female (cl 11.0 mm), ZRC 1996.47, Mandai, mangrove, burrow in soft mud, leg. P.K.L. Ng, 18.05.1984; 1 male (cl 10.8 mm), ZRC 2000.2145, Sungei Buloh, mangrove, mud, leg. NUS Zoology Class, 28.07.1992; 1 male (cl 15.0 mm, missing minor cheliped), ZRC 1979.4.4.14, Geylang River, sta. B71, mud bottom, depth: 2 m, leg. S.R.F.R.S., 03.03.1955 [J6851, det. D.S. Johnson as A. microrhynchus]; 1 female (cl 10.7 mm, identification tentative), ZRC 1991.10140, Singapore River, mud, leg. R.E.S.T., 03.10.1988. Philippines: 2 males (cl 13.8, 14.2 mm), 1 female (cl 18.3 mm), ZRC 1999.0235, Manila (?) fish market, leg. S.H. Tan, 26.03.1996; 1 female, RMNH Crus.D. 49750, Panay, Ilo Ilo, leg. J.C. Miquel, 19.06.1979; 1 female (cl 16.9 mm), ZRC 2014.0725, Panay, Ilo Ilo, market near fishing port, leg. N.K. Ng & J.C.Y. Lai, 22.01.2005; 1 female? (cl 21.4 mm, pleopods damaged, missing minor cheliped), USNM 252028, Palawan, Malampaya River, 26.12.1908; 1 male (cl 18.4 mm), USNM 1199121, Luzon, Manila Bay, Malolos, Guzman Island, Bulacan, fish pond, 02.09.1927. Thailand: 1 male (cl 20.3 mm), OUMNH. ZC. 2019.06.46, Songkhla Lake (Thale Sap), leg. S. Hajisamae, 17.08.2013; 2 males (cl 13.3, 15.6 mm), 2 females (cl 10.1, 13.2 mm), ZRC 2014.0675, same collection data as for previous specimen; 1 male (cl 16.8 mm), ZRC 2014.0672, Pattani Bay, Pattani, leg. S. Hajisamae, 01.2013; 1 male (cl 16.0 mm), OUMNH. ZC. 2019.06.47, same collection data as for previous specimen; 4 females (cl 16.8–20.8 mm), USNM 65478, Tachin River, leg. H.M. Smith, 30.10.1923; 1 female (cl 20.0 mm), USNM 65557, Paknam, Menam Chao Phraya near Bangkok, leg. H. Smith, 21.07.1925; 1 female (cl 22.0 mm), NHM 1976.559, Phuket, leg. N. Ngoc-Ho, 1973 [det. D.M. Banner as A. euphrosyne]; 2 males (cl 21.6, 24.0 mm), 2 females (cl 22.5, 24.5 mm), NHM 1976.560, Phuket, leg. N. Ngoc-Ho, no date (possibly 1973) [det. A. Anker as A. euphrosyne ?]. Cambodia: 1 male (cl 18.4 mm), 3 ov. females (cl 16.0– 20.7 mm), MNHN-IU-2014-10287, locality not specified, “Sac 200 / 7C”, leg. M. Fily; 2 males, 3 ov. females (cl indet.), MNHN-IU-2018-5727, locality not specified, “Sac 200 / 7B”, leg. M. Fily. Vietnam: 1 female (cl 11.9 mm), OUMNH. ZC. 2019.06.48, Haiphong, Cam River, back swamp, leg. K. Wada, 12.09.1995 [YMP-1557]; 1 female (cl 17.8 mm), MNHN-IU-2018-5737, Haiphong, Cam River, back swamp, leg. K. Wada, 12.09.1995 [YMP-1558]; 3 males (cl 13.0– 14.2 mm, pinned dried specimens), MNHN-IU-2000-1124, Tonkin, locality not specified, leg. H. Muhllk / coll. J. Bourgeois, 1911; 1 ov. female (cl 18.7 mm), MNHN-IU-2018-5729, Ha Tien, Sau, 20–30 m (?), leg. H. Duy, 21.08.1995. China: 2 females (cl 18.0, mm, 18.5 mm), 1 ov. female (cl 13.4 mm), USNM 62108, locality unknown, don. S.F. Light. Australia: 1 ov. female (cl 16.2 mm, missing minor cheliped), AM P 28130, Queensland, Cairns, Barron River, 16°55’S 145°46’E, among crab burrows (Sesarma sp.), deep within riverbank, leg. J.C. Yaldwyn, 02.11.1963. Tentative identification. Alpheus cf. eurydactylus. Singapore: 1 male (cl 10.5 mm, tl 28.0 mm, rostrum damaged, missing minor cheliped), ZRC 2014.0724, Lim Chu Kang, mangrove, sta. Y595, 04.04.1989; 1 female (cl 15.7 mm, missing both chelipeds), ZRC 2014.0674, Jurong prawn pond, no further data [J8048]; 1 female (cl 9.6 mm, tl 26.0 mm), ZRC 1979.4.4.6, Jurong, prawn ponds, leg. S. R.F. R.S., 1– 2.06.1954 [J8096]; 1 female (cl 12.5 mm, tl 35.0 mm, missing major cheliped), ZRC 1992.8064, Sungei Buloh, mangrove swamp, leg. P.K.L. Ng, 03.04.1991. India: 1 ov. female (cl indet.), RMNH Crus.D. 49749, Kochi (Cochin), leg. J.C. Miquel, 28.03.1979 (det. Y. Miya as A. euphrosyne). Redescription. See Figs. 11–15. Large-sized species of Alpheus (maximal cl 20.3 mm, tl ~65.0 mm). Carapace glabrous, finely pitted, especially around cardiac region, without pubescence, with slight grooves. Rostrum short, subtriangular, narrowing towards apex, distally acute, typically falling short of mid-length of first article of antennular peduncle; rostral carina feebly developed, low, flattening posterior to orbital hoods; rostro-orbital furrows distinct, shallow; anterior margin of orbital hoods shallowly concave. Pterygostomial angle broadly rounded; cardiac notch deep. Pleon smooth, finely pitted, without pubescence. Telson broad, ovate-rectangular, slightly tapering towards posterior margin, about 1.3–1.4 times as long as maximal width near proximal margin; lateral margins very shallowly concave in proximal half almost straight in distal half; dorsal surface finely pitted, with shallow, longitudinal, median depression (without setae on edges) and two pairs of stout spiniform setae inserted in deep pits at some distance from lateral margins, first pair near telson mid-length, second pair at about 0.7 of telson length; posterior margin broadly rounded, with two pairs of small spiniform setae at posterolateral angles, lateral shorter than mesial (sometimes broken or missing). Antennular peduncle with stylocerite broad, moderately convex laterally, with acute tip, latter not overreaching distal margin of first article; ventromesial carina with anteriorly rounded tooth, without acute point; second article about 2.2–2.3 times as long as wide. Antennal peduncle with distoventral margin of basicerite unarmed; carpocerite not reaching beyond scaphocerite; scaphocerite with lateral margin straight to very shallowly concave; blade very broad, separated from broad distolateral tooth by deep cleft; anterior margin of blade broadly rounded and reaching distinctly beyond distolateral tooth. Third maxilliped with antepenultimate article not particularly broadened; penultimate article about 2.5 times as long as wide, with long setae on ventrolateral margin, latter surpassing ultimate article; ultimate article with dense transverse rows of short serrulate setae on mesial surface and some longer setae on dorsal and lateral surfaces, especially near or on apex; coxal lateral plate bluntly produced dorsally; exopod fleshy, broad, reaching end of antepenultimate article. Major cheliped of A. edwardsii - type. Merus very stout, about 1.8 times as long as wide; ventromesial margin unarmed distally. Carpus very short, cup-shaped. Chela large, massive, with fingers about 0.6–0.7 length of palm. Mesial face of palm with low mesial transverse ridge in proximal half, extending from proximal edge of subtriangular mesial longitudinal groove to ventral surface of palm; surface smooth, largely without granules, except for very few near ventral shoulder. Lateral face of palm smooth, without granulation; lateral longitudinal groove deep, subrectangular; lateral half with low transverse ridge. Dorsal shoulder pronounced, rounded, sloping with angle of 45º–80º into broad transverse groove, not overhanging. Ventral shoulder pronounced, slightly protruding in lateral view, broadly rounded. Pollex with mesial face granulated over most of its surface, with prominent mesial subdistal ridge marked by conspicuous protuberance proximally and extending to subdistal angle of pollex; area ventral to mesial subdistal ridge conspicuously grooved; distomesial angle distinctly superior to 90º, blunt; lateral face smooth, not granulated; distolateral angle about 90º, broadly rounded. Dactylus slightly longer than pollex, with prominent dorsal ridge, latter twisted mesially and protruding proximally; mesial surface granulated in proximal half; lateral surface largely smooth; dactylar plunger large, stout, clearly distinct from ventral dactylar margin; distal surface of plunger with obliquely flattened ventromesial area, its mesial surface with feebly developed transverse channel. Adhesive discs small. Male minor cheliped with chela strongly balaeniceps. Merus somewhat slenderer than that of major cheliped, about 2.5 times as long as wide; ventral margin with distomesial angle unarmed. Chela moderately swollen, with fingers about as long as palm. Palm subcylindrical, somewhat swollen; mesial face with surface largely smooth, except for very small field of granules on subdistal swelling below mesial longitudinal groove; mesial longitudinal groove well marked, deep; lateral longitudinal groove well marked, deep; dorsal shoulder pronounced, rounded, sloping with 45º or smaller angle into deep dorsal transverse groove; ventral transverse groove moderately deep, bordered by distinct ventral shoulder on lateral surface. Pollex with largely smooth surfaces; mesial surface with row of balaeniceps setae on low crest extending from base to slightly beyond mid-length of pollex; lateral surface with similar row of balaeniceps setae as on mesial surface. Dactylus with well-developed proximal balaeniceps expansion, about 2.0–2.2 times as long as maximal width; lateral and mesial dactylar ridges each with dense rows of balaeniceps setae. Cutting edges of both fingers shallowly excavated on each side of sharp, blade-like ridge. Female minor cheliped simple, not balaeniceps. Chela slender, not particularly swollen, with fingers about 1.1–1.2 times length of palm. Palm surface smooth, without granulation (at most with some granules on distomesial surface), with shallow (sometimes traces of) mesial and lateral longitudinal grooves, and with shallow dorsal transverse groove and poorly demarcated ventral shoulder (sometimes not distinct). Fingers simple, with sharp cutting edges, not or slightly gaping when closed; fingertips curved and crossing. Second pereiopod with ratio of carpal subarticles approximately equal to 4.2: 2.5: 1: 1: 1.5. Third pereiopod fairly robust; ischium armed with small, stout spiniform seta on ventrolateral surface near proximal margin, sometimes unarmed (especially in larger individuals); merus about 4.2 times as long as maximal width, unarmed; propodus with five or six stout spiniform setae on ventral margin and one pair of spiniform setae (sometimes one seta) on distoventral margin adjacent to dactylus; dactylus about 0.4 and sometimes close to half-length of propodus, spatulate, expanded, as wide as propodus in dorsal view, with row of tufts of setae along mesial ridge. Fourth pereiopod generally similar to third pereiopod, slightly slenderer; merus about five times as long as maximal width; dactylus about 0.4 length of propodus, spatulate. Fifth pereiopod much slenderer than third and fourth pereiopods; ischium unarmed; propodus with three small spiniform setae and well-developed setal brush; dactylus subspatulate, about 0.4 length of propodus. Male second pleopod with appendix masculina ~0.8 length of appendix interna. Uropod with each protopodal lobe ending in sharp tooth distally; exopod and endopod broad, ovate; diaeresis of exopod complete, almost straight, with small subacute or blunt tooth near small distolateral spiniform seta and adjacent distolateral tooth; lateral margin of exopod broadly convex. Eggs numerous (> 200 in large females), small; egg diameter less than 0.5 mm. Colour pattern. Body uniform dark green, green-brown or brown, without marked banding on carapace or pleon; antennular flagella greenish, bluish proximally; antennal flagella olive-greenish with some bluish tinge proximally; mesial face of major chela olive or greenish brown with large pale orange area, especially near ventral transverse groove; fingers darker olive-green, distally purplish or pinkish; dactylar plunger with mesial side white and blue proximally and bright orange distally and anteriorly; minor chela olive-green with darker fingers; second pereiopods reddish proximally, bright blue distally; ambulatory legs reddish; pleopods pinkish; telson and uropods darker than pleon; uropods with dark, dusky blue patch distally (Fig. 16). One of the Singaporean females (ZRC 1996.47) was accompanied by a short colour note: “overall dark green, chela dactylus purple”. Type locality. Surabaya, Java, Indonesia. Distribution. Tropical Indo-West Pacific from the southern Andaman Sea and Malacca Straits to South-China Sea and northern Australia (Fig. 51B), including coasts of Indonesia (Sumatra, Java, Papua), Singapore (East Coast, Mandai, Sungei Buloh, Punggol), Malaysia (Penang, Selangor, Johor), Thailand (Songkhla Lake, Pattani), Cambodia, Vietnam (Haiphong, Nha Trang, Ho Chi Minh City, Can Gio), China, Philippines (Manila Bay, Panay), and Australia (Queensland) (De Man 1920, 1924; Marin 2021; present study); possibly extending to southern India (Kochi, see discussion below and Fig. 51B). Common name proposed. De Man’s mangrove snapping shrimp. Ecology and biology. Alpheus eurydactylus is a largely intertidal and shallow subtidal snapping shrimp (maximal depth probably not exceeding 3 m) inhabiting estuarine, mangrove mudflats and brackish lagoons, where it lives in deep burrows dug in thick, soft mud, sometimes under large rocks or mangrove debris, among mangrove roots, or in the muddy banks of mangrove creeks. The unusual depth range (20—30 m) given for the female from Ha Tien, Vietnam (MNHN-IU-2018-5729), is somewhat questionable. Like most euryhaline decapods present in these habitats, A. eurydactylus is a resilient shrimp capable of tolerating very low salinities. The ovigerous females of A. eurydactylus carrying a large number of eggs of relatively small size suggest an extended larval development in this species. This large snapping shrimp is occasionally seen on fish markets and crab / shrimp landings, for instance, in the Philippines (Manila). Interestingly, one of the specimens from Thailand (USNM 65557) was accompanied by a note: “kung deet = snapping shrimp, not eaten but used as bait for catfish and drum”. Taxonomic remarks. Alpheus eurydactylus is morphologically very close to A. euphrosyne, as redefined above. The two species can be most easily separated from each other by the dorsal shoulder of the major chela sloping with an angle of>45º into the adjacent transverse groove in A. eurydactylus (Figs. 11C, D, 15A, C) vs. forming a rounded tooth overhanging the transverse groove in A. euphrosyne (Figs. 5C, D, 7A, B). In A. eurydactylus, the mesial palmar surface of the major chela is granulated only distally and subdistally (Fig. 15C), whereas in A. euphrosyne, several fields of granules are also present on the central portion of the palmar surface (Fig. 7A). The dactylar plunger of A. eurydactylus is larger, heavier and not continuous with the ventral edge of the dactylus, as in A. euphrosyne (cf. Figs. 7D, 15D). The rostral carina and rostro-orbital furrows are more demarcated in A. eurydactylus than in A. euphrosyne, where they are both not or barely distinct (cf. Figs. 6A, 14A). Some differences between the two species also exist in the sculpture and proportions of the female minor chelae (cf. Figs. 7H, 15H). The colour patterns of A. eurydactylus (Fig. 16) and A. euphrosyne (Fig. 8–10) are markedly different. In addition, A. eurydactylus is a largely intertidal and shallow subtidal species commonly encountered in soggy mud in mangroves and on estuarine mudflats, whereas A. euphrosyne appears to be a subtidal snapping shrimp preferring sandy-muddy bottoms in deeper water (8–27 m), in brackish lagoons and near estuaries. For separation of A. eurydactylus from other morphologically similar species (especially A. takla sp. nov.) see below or refer to Table 1. Some previous records of A. euphrosyne or A. euphrosyne euphrosyne, for instance those in Banner & Banne, Published as part of Anker, Arthur, 2023, Revision of Alpheus euphrosyne De Man, 1897 and A. microrhynchus De Man, 1897, with description of three new species and taxonomic remarks on several other morphologically and ecologically similar snapping shrimps (Malacostraca: Decapoda: Alpheidae), pp. 1-115 in Zootaxa 5282 (1) on pages 24-33, DOI: 10.11646/zootaxa.5282.1.1, http://zenodo.org/record/7912292, {"references":["De Man, J. G. (1920) Diagnoses of some new species of Penaeidae and Alpheidae with remarks on two known species of the genus Penaeopsis A. M. - Edw. from the Indian Archipelago. Zoologische Mededelingen, Leiden, 5, 103 - 109.","De Man, J. G. (1924) On a collection of macrurous decapod Crustacea chiefly Penaeidae and Alpheidae from the Indian Archipelago. Archiv fu ¨ r Naturgeschichte, Berlin, Series A, 90, 1 - 60.","Banner, A. H. & Banner, D. M. (1966) The alpheid shrimp of Thailand. The Siam Society Monograph Series, 3, 1 - 168.","Johnson, D. S. (1976) Prawns of the Malacca Straits and Singapore waters. Journal of the Marine Biological Association of India, 18, 1 - 54.","De Man, J. G. (1897) Bericht u ¨ ber die von Herrn Schiffscapitan Storm zu Atjeh, an den westlichen Ku ¨ sten von Malakka, Borneo und Celebes sowie in der Java-See gesammelten Decapoden und Stomatopoden. Fu ¨ nfter Theil. Z oologische Jahrbu ¨ cher. Abtheilung fu ¨ r Systematik, Geographie und Biologie der Thiere, 9, 725 - 790, pls. 34 - 37 (in volume 10).","Banner, D. M. & Banner, A. H. (1981) Annotated checklist of the alpheid shrimp of the Red Sea and Gulf of Aden. Zoologische Verhandelingen, Leiden, 190, 1 - 99.","Banner, D. M. & Banner, A. H. (1982) The alpheid shrimp of Australia. Part III: The remaining alpheids, principally the genus Alpheus and the family Ogyrididae. Records of the Australian Museum, 34, 1 - 357. https: // doi. org / 10.3853 / j. 0067 - 1975.34.1982.434","Banner, A. H. & Banner, D. M. (1983) An annotated checklist of the alpheid shrimp from the western Indian Ocean. Travaux et Documents de l'ORSTOM, 158, 1 - 164.","Banner, D. M. & Banner, A. H. (1985) The alpheid shrimp of Indonesia, based upon J. G. de Man's \" The Decapoda of the Siboga Expedition, Part II. Family Alpheidae \" (1911). Marine Research in Indonesia, 25, 1 - 79. https: // doi. org / 10.14203 / mri. v 25 i 0.431","Chace, F. A. Jr. (1988) The caridean shrimps (Crustacea: Decapoda) of the Albatross Philippine Expedition, 1907 - 1910, Part 5: Family Alpheidae. Smithsonian Contributions to Zoology, 466, 1 - 99. https: // doi. org / 10.5479 / si. 19436696.391.1","Marin, I. N. (2021) A new infaunal species of the alpheid shrimp genus Salmoneus Holthuis, 1955 (Crustacea: Decapoda: Alpheidae) and a new crustacean association from anoxic mangrove habitats in southern Vietnam. Arthropoda Selecta, 30, 369 - 385. https: // doi. org / 10.15298 / arthsel. 30.3.10","Johnson, D. S. (1962) A synopsis of the Decapoda Caridea and Stenopodidea of Singapore, with notes on their distribution and a key to the genera of Caridea occuring in Malayan waters. Bulletin of the National Museum of Singapore, 30, 44 - 79, pl. 2.","Coutiere, H. (1905) Les Alpheidae. In: Gardiner, J. S. (Ed.), The fauna and geography of the Maldive and Laccadive Archipelagoes. Being the account of the work carried on and of the collections made by an expedition during the years 1899 and 1900. University Press, Cambridge, pp. 852 - 921, pls. 70 - 87.","Sha, Z., Wang, Y. & Cui, D. (2019) The Alpheidae from China seas. Crustacea: Decapoda: Caridea. Springer, Singapore, 323 pp. https: // doi. org / 10.1007 / 978 - 981 - 15 - 0648 - 2 _ 2","Harikrishnan, M. & Kurup, B. M. (2008) Potential of snapping shrimps as ornamentals. In: Kurup, B. M., Boopendranth, M. R., Ravindran, K., Sairabanu & Nair, A. G. (Eds.), Ornamental fish, breeding, farming and trade. Department of Fisheries, Government of Kerala, Kerala, pp. 1 - 280.","Yeo, D. C. J. & Ng, P. K. L. (1996) A new species of freshwater snapping shrimp, Alpheus cyanoteles (Decapoda: Caridea: Alpheidae) from Peninsular Malaysia and a redescription of Alpheus paludicola Kemp, 1915. Raffles Bulletin of Zoology, 44, 37 - 63."]}

Alpheus songkla Banner & Banner, 1966, stat. nov. (Figs. 41, 52C) [see also Fig. 42 for A. cf. songkla] Alpheus malabaricus songkla Banner & Banner 1966: 147, fig. 56; Angsupanich & Kuwabara 1999: 6; Angsupanich et al. 2005: 376; Naiyanetr 2007: 173. (?) Alpheus malabaricus songkla.— Thomas 1976: 668. Type material. Holotype, female (cl 6.1 mm), USNM 120407, Thailand, Thale Sap (Songkhla Lake), BR44, commercial shrimp trawl, sandy bottom, depth: approximately 1 m, leg. A.H. Banner, 27.03.1963; paratype, female (cl 6.4 mm), USNM 120408, same collection data as for previous specimen. Tentative identification. Alpheus cf. songkla. Singapore: 2 males (cl 8.8, 10.0 mm), 1 female (9.6 mm), ZRC 1992.11117 – 11119, Pulau Ubin, mudflat, leg. P.K.L. Ng, 08.1987; 1 male (cl 6.8 mm, missing minor cheliped), 2 females (cl 5.6, 7.5 mm), ZRC 2014.0665, near Sarimbun Scouts Camp (Jalan Bahtera), seining on mud, leg. H.H. Ng, H. Wong & Y.L. Teo, 14.02.2012 [48193–48195]. Malaysia: 3 males (cl 7.4–8.0 mm), 1 female (cl 9.5 mm), OUMNH. ZC. 2019.06.50 [ex ZRC 2009.0306], Terengganu, Telok Tebrau, intertidal mudflats, with gobies, leg. Z. Jaafar, 05.11.2002. Thailand: 1 male (cl 9.6 mm), USNM 65561, “ Sin Gora inland sea [locality not found], leg. H.M. Smith, 20.06.1925; 1 male (cl 10.6 mm, missing both chelipeds), USNM 65558, Paknam, Menam Chao Phraya near Bangkok, Klang Krang, leg. H. Smith, 21.05.1925; 1 female (cl 9.0 mm), USNM 65477 /2, Bangpakong River, leg. H. Smith, 01.07.1923. Vietnam: 1 male (cl 8.1 mm), MNHN-IU-2018-5655, Duyen-Hai near Ho Chi Minh City, leg. H. Dung, 12.08.1995; 1 male (cl 10.4 mm, minor cheliped abnormal, see below), MNHN-IU-2019-2290, same collection data as for previous specimen. Australia: 1 male (cl 6.5 mm), NTM Cr.015100, Northern Territory, Arnhem Land, Milingimbi, east of Nilpaiwa Islands, 12º03.389’S 134º52.835’E, depth: 2.2 m, leg. S.K. Horner & G.M. Dally, 04.12.2004. Comparative material. Alpheus malabaricus (Fabricius, 1775) sensu lato (sensu Banner & Banner 1982; Chace 1988; but see discussion below). Taiwan: 1 male (cl 7.1 mm), OUMNH. ZC. 2019.06.51, Chang Hua, mudflat, leg. T. Y. Chan et al., 07.1996. Thailand: 1 female (cl 12.0 mm), OUMNH. ZC. 2019.06.52, Lake Thale Sap (= Songkhla), leg. S. Hajisamae, 17.08.2013; 1 male (cl 11.7 mm), OUMNH. ZC. 2019.06.53, Pattani Bay, Pattani, leg. S. Hajisamae, 01.2013; 1 ov. female (cl 9.5 mm), OUMNH. ZC. 2019.06.54, Phuket, E Chalong Bay, mudflat in front of mangrove, depth: Indonesia: 1 male (cl 11.7 mm), OUMNH. ZC. 2019.06.55, Papua, Ajkwa Island, mangrove, leg. A. Darmawan et al., 20.07.2012; 1 ov. female (cl 12.4 mm), QM W25803-1, Papua, Ajkwa River estuary, 4°50’S 136°50’E, Environmental Laboratory, PT Freeport, Sta. Ajk-36, estuarine habitat, 30.03.2000; 3 males (cl 10.2–14.6 mm), 1 female (cl 11.0 mm), QM W25802, Papua, West Ajkwa River, river mouth at Lanal Base, 4°50’S 136°50’E, Environmental Laboratory, PT Freeport, Sta. Ajk-36, estuarine habitat, 29.07.1999; 1 ov. female (cl 11.6 mm), 1 male major cheliped, OUMNH. ZC. 2019.06.56, Lombok, Lembar, prawn ponds, muddy banks of brackish stream, low tide, suction pump, leg. A. Anker, I.S. Pratama, M. Firdaus & D.L. Rahayu, 14.05.2014 [St5-14]; 1 male (cl 9.4 mm), 1 ov. female (cl 7.8 mm), OUMNH. ZC. 2019.06.57, Lombok, Teluk Sekotong, near mangrove, 0.1–0.3 m at low tide, suction pump, leg. A. Anker, I.S. Pratama, M. Firdaus & D.L. Rahayu, 14.05.2014 [St6-03]; 1 ov. female (cl indet.), ZRC 2014.0682, Anambas, sta. EAJL 03, Pulau Jemaja Teluk, deep sheltered sandy bay, northern and eastern sides with fringing Rhizophora and Bruguiera mangrove inlets, leg. J.C.Y. Lai et al., 13.03.2002. Singapore: 2 males (cl 15.4, 15.9 mm), 1 ov. female (cl 16.7 mm), ZRC 1992.11124 – 11126, Pulau Ubin, mudflat, leg. P.K.L. Ng, 08.1987; 2 males (cl 12.4, 12.5 mm), ZRC 1992.11122 – 11123, same collection data as for previous specimens; 1 male (cl 9.0 mm), ZRC 1994.4392, Kallang Basin, sta. 5, dredge, leg. Reef Ecology Study Team, 16.12.1994. Australia: 1 male (cl indet.), QM W25812, SE Queensland, Innes Park, creek, 300 m from mouth, mangroves, sand, leg. J. Johnson & A. Gill, 12.05.2000; 1 male (cl 13.4 mm), 1 ov. female (cl 14.9 mm), NTM Cr. 008307, Northern Territory, Darwin, Ludmilla Creek mouth, 12º24.8’S, 130º50.7’ E, undisturbed mangrove, low tide, leg. M. Burke, 04.06.1991. Description. See Banner & Banner (1966) for original description and illustrations, as A. malabaricus songkla; see also discussion below. Colour pattern. The colour pattern of the type specimens from Songkhla Lake is unknown. The colour pattern of the Indian specimens identified by Thomas (1976) as A. malabaricus songkla was described as following: general body colour cream with dark brown cross bands along posterior margins of pleonites and carapace; tips of uropods and anterior border of carapace between orbital hood and lateral angle dark brown; antennal flagella bluish violet, antennular flagella with brownish tinge; chelipeds grey with violet inner [mesial?] depression of large chela; remaining portion of chela and legs pinkish; exopods and endopods of pleopods [erroneously called chelipeds] bright red with paler bases; underside of pleon and chela white (adapted from Thomas 1976; however, see below). Type locality. Thale Sap (Songkhla Lake), Thailand. Distribution. Indo-West Pacific: with certainty known only from the type locality in Thailand, Thale Sap = Songkhla Lake (Banner & Banner 1966); record from India (Thomas 1976) requires confirmation; additional records of A. cf. songkla from Thailand, Vietnam, Malaysia, Singapore and Northern Territory, Australia (Fig. 52C) (see discussion below). Common name proposed. Songhkla snapping shrimp. Ecology and biology. Shallow subtidal species found on mud and sand bottoms usually off mangroves or inside brackish and saltwater lagoons; the South-East Asian and Australian specimens were collected in 1–3 m deep water (Banner & Banner 1966; present study), whereas the Indian specimens (which may or may not be A. songkla) came from a depth range of 3–10 m (Thomas 1976). Taxonomic remarks. Banner & Banner (1966) separated their new subspecies A. malabaricus songkla from A. malabaricus, including the forms known as A. malabaricus dolichodactylus Ortmann, 1890 and A. malabaricus leptopus De Man, 1910 (both now in the synonymy of A. malabaricus, see below), by the relative length of the fingers of the minor chela, which are 1.5 times as long as the palm (vs. at least three times as long as the palm in the other three forms). In addition, in A. malabaricus songkla, the distal parts of the fingers are strongly crossing, which is not the case of A. malabaricus dolichodactylus and A. malabaricus leptopus. In the general shape and proportions of the minor chela, A. malabaricus songkla approaches A. malabaricus mackayi Banner, 1959, which was elevated to species rank, as A. mackayi (Banner 1959; Banner & Banner 1974). However, Chace (1988) placed all the above-mentioned species and varieties, including A. malabaricus songkla and A. mackayi, as well as A. macrodactylus Ortmann, 1890, A. malabaricus trefzae Banner & Banner, 1982 and A. mazatlanicus Wicksten, 1983, in the synonymy of A. malabaricus. The clearly premature synonymisations of Chace (1988) resulted in the morphologically and ecologically highly variable A. malabaricus sensu lato, a “species” with a vast geographic range, intertidal and deep-water populations, specimens with relatively short to extremely elongated fingers of the minor cheliped, specimens with the dactylar plunger of the major cheliped ranging from very large and stout to greatly reduced, and a great deal of other “intraspecific variation”. Although the revision of the entire A. malabaricus complex (A. malabaricus sensu Chace 1988) is well beyond the scope of the present study, the author sees no reason for treating A. macrodactylus, A. mackayi, A. mazatlanicus and A. malabaricus songkla as junior synonyms of A. malabaricus (cf. Banner & Banner 1966, 1974, 1982, 1983; Kim & Abele 1988), whilst the taxonomic status of A. malabaricus dolichodactylus, A. malabaricus leptopus and A. malabaricus trefzae (cf. De Man 1911; Banner & Banner 1982) will need further clarification. Nevertheless, awaiting a long-needed revision of A. malabaricus, the comparative material examined in this study is listed under A. malabaricus sensu lato, even though it certainly contains more than one species. Banner & Banner (1966) noted that A. malabaricus songkla differs from A. mackayi by the longer rostrum; the lack of rostro-orbital furrows; and the fingers of the minor chela slenderer and with longer, distally stronger crossing tips. In A. malabaricus songkla, the orbital hoods seem to be less projecting than in A. mackayi, whereas the second article of the antennular peduncle is noticeably shorter than in A. mackayi (cf. Banner & Banner 1966: fig. 56A; Banner 1959: fig. 12a, b). Furthermore, the telson of A. malabaricus songkla has straight lateral margins; these are strongly convex in A. mackayi (cf. idem: fig. 56H; fig. 12m). The main differences between A. malabaricus songkla and A. macrodactylus consist in the less projecting orbital hoods (cf. Banner & Banner 1966: fig. 56A; Banner & Banner 1982: fig. 65a); the ratio of the fingers to the palm in the major chela being around 0.7 in A. malabaricus songkla vs. closer to 1.0 in A. macrodactylus (cf. idem: fig. 56B, C; fig. 65b, c); and the minor cheliped fingers somewhat shorter relative to the palm (1.5 times as long as the palm), without armature on the cutting edges and with the fingertips strongly crossing in A. malabaricus songkla vs. noticeably longer (1.8 times as long as the palm), with small teeth in the proximal portion of the cutting edges and with the fingertips not strongly crossing in A. macrodactylus (cf. idem: fig. 56D, E; fig. 65e, f). Finally, A. malabaricus songkla can be easily separated from the eastern Pacific A. mazatlanicus, for instance, by the shape of the rostro-orbital area; the length of the second article of the antennular peduncle; the shape of the major chela dactylus; and the relative proportions of the carpal subarticles in the second pereiopod (cf. Banner & Banner 1966: fig. 56; Kim & Abele 1988: fig. 36). Therefore, A. malabaricus songkla is herein elevated to full species rank, as A. songkla stat. nov. Despite the fact that A. songkla is morphologically different from all species and subspecies (or varieties) currently assigned to the A. malabaricus complex, it remains a problematic taxon. The description by Banner & Banner (1966) was based exclusively on females, i.e., the morphology of the male minor chela, of critical importance for taxonomy of Alpheus, currently remains unknown. Therefore, at this stage, a redescription or a new diagnosis for A. songkla would not be very useful. However, an opportunity is taken to correct a number of small errors and inaccuracies in the original description of A. songkla by Banner & Banner (1966), based on re-examination of the type material (Fig. 41). One of the most important features of A. songkla setting it clearly apart from the A. malabaricus complex is the relatively weak anterior projection of the orbital hoods (Banner & Banner 1966: fig. 56A; see also Figs. 41A, 42A; cf. Banner & Banner 1974: fig. 12a; Banner & Banner 1982: fig. 64a, 65a; Kim & Abele 1988: fig. 36a, b; Chace 1988: fig. 9a). The antennal basicerite of the holotype (USNM 120407) has a small, curved tooth in the middle of the distolateral margin (apparently broken on the right side), which seems to correspond to “minute lateral spine” in the description of Banner & Banner (1966). However, the antennal basicerite of the examined paratype (USNM 120408) has a similar curved tooth only on the right side, being unarmed on the left side. The ischium of the third pereiopod has a small spiniform seta, as correctly figured by Banner & Banner (1966: fig. 56G). The propodus of the third pereiopod has a row of tightly appressed spiniform setae between the long stiff setae; only the latter were shown by Banner & Banner (1966: fig. 56G), who stated that the propodus is “bearing strong setae but no spinules”, which is incorrect. Furthermore, the mesial face of the major chela is not perfectly smooth, but has a large field of fine granules distally, i.e., on the distal portion of the palm and most of the pollex, reminiscent of the granulation of A. eurydactylus (although weaker). The mesial subdistal ridge of the pollex is long and sharp, and departs from a conspicuous proximal bump, as in A. euphrosyne, A. eurydactylus, A. takla sp. nov., etc. The last two taxonomically important features of the major chela (granulation and mesial subdistal ridge) were not mentioned nor illustrated by Banner & Banner (1966). In addition, the minor chela fingers of both examined type specimens are quite setose, covered by tufts of numerous stiff setae; these setae were omitted in the original illustrations of the minor cheliped by Banner & Banner (1966: fig. 56D, E). The ventromesial carina of the first article of the antennular peduncle (not illustrated in the original description) is broadly rounded-triangular and without an acute point. The taxonomic identity of two males and four females from southern India (Korapuzha estuary north of Kozhikode on the Malabar Coast) reported as Alpheus malabaricus songkla by Thomas (1976) remains uncertain. According to Thomas (1976), both male and female specimens had “dense setae” on the lateral margins of the minor chela fingers, which were not referred to as “balaeniceps setae” (the same author reported a balaeniceps minor chela in A. euphrosyne). Banner & Banner (1966) did not mention nor illustrated dense setae on the female minor cheliped of A. songkla, but the re-examination of the type material confirmed that the minor chela is indeed quite setose (see above). Noteworhy is that Thomas’ (1976) descriptions of the colour patterns of A. m. songkla and A. euphrosyne are almost identical. The material from Singapore, Malaysia, Vietnam, Thailand and Australia herein tentatively identified as A. cf. songkla is morphologically somewhat heterogeneous and needs further study. The material from Singapore, Thailand and Vietnam is discussed in more detail below. (1) The specimens from Pulau Ubin and Sarimbun, Singapore (ZRC 1992.11117–11119, ZRC 2014.0665), are morphologically most similar to A. songkla, e.g., in the configuration of the rostro-orbital area and antennules; the presence of a small tooth on the antennal basicerite; the relatively slender major chela, with a moderately developed plunger on the dactylus; and the general proportions of the female chela. The general characteristics of the male minor cheliped place these specimens close to A. eurydactylus. However, they differ from A. eurydactylus in the longer distolateral tooth of the scaphocerite, exceeding the distal margin of the blade (vs. not exceeding it in A. eurydactylus), and in the dorsal surface of the telson without longitudinal depression (present in A. eurydactylus). In addition, the Singaporean specimens have slenderer chelipeds compared to those of similarly sized specimens of A. eurydactylus. (2) The male from “Sin Gora inland sea” in Thailand (USNM 65561) generally agrees with A. songkla, for instance, in the similar frontal area; the third pereiopod ischium armed with a spiniform seta; the third pereiopod propodus bearing stout appressed spines and long stiff setae; and the spatulate third pereiopod dactylus. However, the distolateral margin of the antennal basicerite of this specimen has no trace of tooth on both sides. The minor chela of this male specimen, which cannot be directly compared with that of the females of A. songkla, is very similar to that of A. eurydactylus. (3) The incomplete male from Menam Chao Phraya near Bangkok, Thailand (USNM 65558), matches A. songkla in most characters, including the very similar rostro-orbital area; the presence of a small, sharp tooth on the distolateral margin of the antennal basicerite (on both sides); the major chela with a gently sloping dorsal shoulder and granulated distomesially (with minute granules on the distal portion of the palm, pollex and dactylus); the major chela pollex with a distinct mesial subdistal ridge; the second pereiopod with the first carpal article much longer than the second; and the propodus of the third and fourth pereiopods furnished with stout appressed spines and long stiff setae. However, the ischium of the third and fourth pereiopods of this specimen is unarmed (vs. armed with a spiniform seta in A. songkla), and the third pereiopod merus appears to be somewhat broader than in A. songkla, as illustrated by Banner & Banner (1966: fig. 56G). The female from Bangpakong River, Thailand (USNM 65477/2), has essentially the same characterstics as the male from Menam Chao Phraya and seems to belong to the same species. (4) The two males from Duyen-Hai near Ho Chi Minh City, Vietnam (MNHN-IU-2018-5655, MNHN-IU-2019- 2290, see Fig. 42), were initially identified by the author as A. cf. eurydactylus. They differ from the typical A. eurydactylus, however, by the slenderer second pereiopod, with the first article much longer than the second; the third pereiopod ischium armed with a spiniform seta (as in A. songkla); and the slightly different proportions of the minor chela fingers to the palm in the male with the normally developed minor cheliped (MNHN-IU-2018-5655). In the male with a somewhat abnormal minor cheliped (MNHN-IU-2019-2290), the palm has a distinct dorsal notch, whilst the fingers possess a weak balaeniceps crest. On the other hand, they appear to be more similar to the afore-mentioned specimens of A. cf. songkla from Singapore. Thus, the above character combinations do not allow identifying the material from Singapore, Vietnam and Thailand reliably as A. songkla (or as A. eurydactylus) and the situation is further complicated by the variation in the armature of the antennal basicerite in the type material of A. songkla and of the ischium of the third pereiopod in A. eurydactylus. Since A. songkla is presentl, Published as part of Anker, Arthur, 2023, Revision of Alpheus euphrosyne De Man, 1897 and A. microrhynchus De Man, 1897, with description of three new species and taxonomic remarks on several other morphologically and ecologically similar snapping shrimps (Malacostraca: Decapoda: Alpheidae), pp. 1-115 in Zootaxa 5282 (1) on pages 75-81, DOI: 10.11646/zootaxa.5282.1.1, http://zenodo.org/record/7912292, {"references":["Banner, A. H. & Banner, D. M. (1966) The alpheid shrimp of Thailand. The Siam Society Monograph Series, 3, 1 - 168.","Angsupanich, S. & Kuwabara, R. (1999) Distribution of macrobenthic fauna in Phawong and U-Taphao canals flowing into a lagoonal lake, Songkhla, Thailand. Lakes & Reservoirs: Research and Management, 4, 1 - 13. https: // doi. org / 10.1046 / j. 1440 - 1770.1999.00080. x","Angsupanich, S., Siripech, A. & Charoenpornthip, M. (2005) Macrobenthic fauna community in the Middle Songkhla Lake, southern Thailand. Songklanakarin Journal of Science and Technology, 27 (Supplement 1), 365 - 390.","Naiyanetr, P. (2007) Checklist of crustacean fauna in Thailand (Decapoda, Stomatopoda, Anostraca, Myodocopa and Isopoda). Office of Natural Resources and Environmental Policy and Planning, Bangkok, 196 pp.","Thomas, M. M. (1976) New records of four alpheid shrimp from the Indian waters. Journal of the Marine Biological Association of India, 18, 666 - 669.","Fabricius, J. C. (1775) Systema Entomologiae, sistens Insectorum classes, ordines, genera, species, adiectis synonymis, locis, descriptionibus, observationibus. Officina Libraria Kortii, Flensbergi et Lipsiae, xxx + 832 pp. https: // doi. org / 10.5962 / bhl. title. 36510","Banner, D. M. & Banner, A. H. (1982) The alpheid shrimp of Australia. Part III: The remaining alpheids, principally the genus Alpheus and the family Ogyrididae. Records of the Australian Museum, 34, 1 - 357. https: // doi. org / 10.3853 / j. 0067 - 1975.34.1982.434","Chace, F. A. Jr. (1988) The caridean shrimps (Crustacea: Decapoda) of the Albatross Philippine Expedition, 1907 - 1910, Part 5: Family Alpheidae. Smithsonian Contributions to Zoology, 466, 1 - 99. https: // doi. org / 10.5479 / si. 19436696.391.1","Ortmann, A. (1890) Die Decapoden-Krebse des Strassburger Museums, mit besonderer Beru ¨ cksichtigung der von Herrn Dr. D ˆ derlein bei Japan und bei den Liu-Kiu-Inseln gesammelten und z. Z. im Strassburger Museum aufbewahrten Formen. I. Theil. Die Unterordnung Natantia (Boas) (Abtheilungen: Penaeidae und Eucyphidea = Caridae der Autoren). Zoologische Jahrbu ¨ cher. Abtheilung fu ¨ r Systematik, Geographie und Biologie der Thiere, 5, 437 - 542, pls. 36, 37.","De Man, J. G. (1910) Diagnoses of new species of macrurous decapod Crustacea from the \" Siboga-Expedition \". Tijdschrift der Nederlandsche Dierkundige Vereeniging, (2) 11, 287 - 319.","Banner, A. H. (1959) Contributions to the knowledge of the alpheid shrimp of the Pacific Ocean. Part IV. Various small collections from the Central Pacific area, including supplementary notes on alpheids from Hawaii. Pacific Science, 13, 130 - 155.","Banner, A. H. & Banner, D. M. (1974) Contributions to the knowledge of the alpheid shrimp of the Pacific Ocean. Part XVII. Additional notes on the Hawaiian alpheids: new species, subspecies and some nomenclatorial changes. Pacific Science, 28, 423 - 437.","Wicksten, M. K. (1983) A monograph on the shallow water caridean shrimps of the Gulf of California, Mexico. Allan Hancock Monographs in Marine Biology, 13, 1 - 59.","Banner, A. H. & Banner, D. M. (1983) An annotated checklist of the alpheid shrimp from the western Indian Ocean. Travaux et Documents de l'ORSTOM, 158, 1 - 164.","Kim, W. & Abele, L. G. (1988) The snapping shrimp genus Alpheus from the eastern Pacific (Decapoda: Caridea: Alpheidae). Smithsonian Contributions to Zoology, 454, 1 - 119. https: // doi. org / 10.5479 / si. 00810282.454","De Man, J. G. (1911) The Decapoda of the Siboga Expedition. Part II. Family Alpheidae. Siboga-Expeditie, 39 a 1 (2), 133 - 465; (1915) Supplement - Explanations of plates of Alpheidae, pls. 1 - 23. E. J. Brill, Leiden. https: // doi. org / 10.5962 / bhl. title. 10512"]}

Alpheus paludicola Kemp, 1915 (Figs. 3A–G, 43, 52D) Alpheus paludicola Kemp 1915: 303; Yeo & Ng 1996: 39, figs. 2, 3. (?) Not Alpheus paludicola. — Kemp 1918: 273 (= possibly A. microrhynchus De Man, 1897). Not Alpheus paludosus (lap. cal.).— Johnson 1965: 9 (part. = A. cyanoteles Yeo & Ng, 1996). Type material. Lectotype, male (cl 5.5 mm, tl 18.0 mm, chl 10.0 mm), NHM 1919.11.1.1, India, Orissa, Chilika (= Chilka) Lake, Outer Channel, leg. S. Kemp, 1914; paralecotype, male (cl 5.2 mm, tl 14.0 mm, chl 9.7 mm), NHM 1919.11.1.2, same collection data as for lectotype. Description. See Kemp (1915) for original description and illustrations (Fig. 3A–G), and Yeo & Ng (1996) for full redescription and additional figures; complementary illustrations of the type material are provided in Fig. 43. Colour pattern. Overall translucent with brownish red rostrum; antennular peduncle and lateral margin of scaphocerite tinged with reddish brown; posterior edge of carapace and each pleonite with transverse brownish to bluish green band; telson and uropods dusky (adapted from Kemp 1915). Type locality. Chilika Lake (formerly Chilka Lake), Orissa, India. Distribution. Northern Indian Ocean: currently known only from the type locality in Orissa, northeastern India (Fig. 52D). The material from Lake Thale Sap Thailand reported by Kemp (1918) as A. paludicola most likely represents another, perhaps undescribed species (see also comments in Yeo & Ng 1996). Common name proposed. Chilika snapping shrimp. Ecology and biology. Alpheus paludicola is confined to brackish lagoons, where it lives in burrows on soft mud, at a depth range of 1– 4 m. The species can be classified as euryhaline, being able to tolerate fluctuations in salinity (Kemp 1915). The small number of eggs and their large size (diameter about 1.4 mm) suggest an abbreviated larval development, as seen in A. microrhynchus and A. cyanoteles (see above). Taxonomic remarks. In their very detailed redescription of A. paludicola, Yeo & Ng (1996) noted that the stylocerite is missing its distal point on both sides in the lectotype (see Yeo & Ng 1996: fig. 2c) vs. terminating in a small acute point in the paralectotype. The major chela was illustrated by Yeo & Ng (1996: fig. 2e) without a mesial subdistal ridge on the pollex, whereas the fine granulation of the major chela observed by Kemp (1915) was “not visible” according to these authors. However, the reexamination of the lectotype and paralectotype of A. paludicola by the present author revealed that the major chela pollex of the lectotype has a small mesial subdistal ridge, whereas the distal portion of the mesial side of the major chela palm and the adjacent area of the pollex are indeed covered with minute granules, as correctly observed by Kemp (1915). Another possible small inaccuracy in Yeo & Ng’s (1996) illustrations of the major cheliped of A. paludicola is the absence of a mesial longitudinal groove connecting with the dorsal transverse groove, which is present, albeit very shallow, in the lectotype. Alpheus paludicola (Figs. 3A–G, 43) can be separated from A. euphrosyne, A. eurydactylus, A. richardsoni, A. microrhynchus, A. cyanoteles, A. nomurai sp. nov., A. takla sp. nov., A. mangalis sp. nov. and A. songkla by the extremely short rostrum (which is noticeably longer in most other species, except for A. microrhynchus); the development of the rostro-orbital furrows (absent in A. paludicola vs. present in most other species, although very shallow in A. eurphrosyne and not distinct in A. microrhynchus); the distally unarmed antennal basicerite (vs. armed with a more or less strong tooth in A. richardsoni, A. cyanoteles, A. nomurai sp. nov., A. mangalis sp. nov. and A. takla sp. nov., occasionally with a minute tooth also in A. microrhynchus); the shape of the dorsal shoulder of the major chela (gently sloping in A. paludicola vs. with an overhanging tooth in A. euphrosyne); the proportions of the fingers to the palm in the major chela (0.4 in A. paludicola vs. 0.6–1.0 in all the other species); the development of the mesial subdistal ridge on the major chela pollex (feebly developed in A. paludicola vs. very strong in A. euphrosyne, A. eurydactylus, A. microrhynchus, A. cyanoteles and A. takla sp. nov.); the degree of granulation on the mesial surface of the major chela (weak and restricted to the distal portion of the palm and pollex in A. paludicola vs. much stronger and extensive in A. euphrosyne and A. eurydactylus, or absent in A. microrhynchus, A. cyanoteles and A. mangalis sp. nov.); the sculpture of the palm of the male minor chela (with weak dorsal and ventral transverse grooves in A. paludicola vs. with much stronger ones in A. euphrosyne and A. eurydactylus, or without a distinct dorsal notch in A. richardsoni and A. takla sp. nov.); the shape of the dactylus of the third to fifth pereiopods (spatulate in A. paludicola vs. trigonal-subspatulate in A. microrhynchus and A. cyanoteles); the armature of the ischium of the third pereiopod (unarmed in A. paludicola vs. armed with a spiniform seta in A. euphrosyne, A. eurydactylus, A. richardsoni, A. nomurai sp. nov. and A. mangalis sp. nov., although variable in some taxa); and the number and size of eggs in females (few and very large in A. paludicola vs. numerous and small in the other species, except for A. microrhynchus and A. cyanoteles). In addition, A. paludicola differs from A. mangalis sp. nov. by the distally unarmed ventromesial margin of the chelipeds. The type specimens, with maximal cl 5.5 mm / tl 22.0 mm, are also much smaller than all the above-listed species of the A. euphrosyne — A. microrhynchus complex. In life, A. paludicola may be easily separated from A. euphrosyne, A. richardsoni, A. cyanoteles and A. nomurai sp. nov., by its duller and more uniform colour pattern (Kemp 1915), although young or paler individuals of A. microrhynchus, A. eurydactylus, A. mangalis sp. nov. and A. takla sp. nov. may bear some resemblance with A. paludicola., Published as part of Anker, Arthur, 2023, Revision of Alpheus euphrosyne De Man, 1897 and A. microrhynchus De Man, 1897, with description of three new species and taxonomic remarks on several other morphologically and ecologically similar snapping shrimps (Malacostraca: Decapoda: Alpheidae), pp. 1-115 in Zootaxa 5282 (1) on page 81, DOI: 10.11646/zootaxa.5282.1.1, http://zenodo.org/record/7912292, {"references":["Kemp, S. (1915) Fauna of the Chilka Lake. No. 3. Crustacea Decapoda. Memoirs of the Indian Museum, 5, 201 - 325, pls. 12 - 13. https: // doi. org / 10.5962 / bhl. title. 10414","Yeo, D. C. J. & Ng, P. K. L. (1996) A new species of freshwater snapping shrimp, Alpheus cyanoteles (Decapoda: Caridea: Alpheidae) from Peninsular Malaysia and a redescription of Alpheus paludicola Kemp, 1915. Raffles Bulletin of Zoology, 44, 37 - 63.","Kemp, S. (1918) Crustacea Decapoda of the Inle Lake Basin. Records of the Indian Museum, 14, 81 - 102, pls. 24 + 25. https: // doi. org / 10.26515 / rzsi / v 14 / i 4 / 1918 / 163580","De Man, J. G. (1897) Bericht u ¨ ber die von Herrn Schiffscapitan Storm zu Atjeh, an den westlichen Ku ¨ sten von Malakka, Borneo und Celebes sowie in der Java-See gesammelten Decapoden und Stomatopoden. Fu ¨ nfter Theil. Z oologische Jahrbu ¨ cher. Abtheilung fu ¨ r Systematik, Geographie und Biologie der Thiere, 9, 725 - 790, pls. 34 - 37 (in volume 10).","Johnson, D. S. (1965) A review of brackish water prawns of Malaya. Bulletin of the National Museum of Singapore, 33, 7 - 11."]}

Alpheus bunburius Banner & Banner, 1982 (Fig. 46A–D, Fig. 52D) Alpheus bunburius Banner & Banner 1982: 213, fig. 66. Type material. Holotype: ov. female (cl 13.7 mm, tl 38.0 mm, chl 13.5 mm), WAM 271-65, Australia, Western Australia, Bunbury, leg. W.H. Butler, 03.1962. Description. See Banner & Banner (1982) for detailed description and illustrations; see also Fig. 46A–D and remarks below. Colour pattern. Unknown. Type locality. Bunbury, Western Australia. Distribution. Southeastern Indian Ocean: presently known only from the type locality in Western Australia (Fig. 52D). Common name proposed. Bunbury snapping shrimp. Ecology and biology. The general ecology and biology of A. bunburius remain largely unknown (but see below). The presence of numerous small eggs (diameter ~1.0 mm) in the female holotype indicates an extended larval development. Taxonomic remarks. Alpheus bunburius was described on the basis of a single female specimen from Bunbury, Western Australia, without any biological or ecological information.According to Banner & Banner (1982), the form of this species [A. bunburius] is very similar to A. euphrosyne euphrosyne De Man and A. euphrosyne richardsoni Yaldwyn, but in addition to the differences in the palmar/finger length this species differs in the low dactylar plunger of the dactylus of the large chela and the trigonal dactylus of the third pereiopod. However, the rather unusual proportions and small size (relative to the body) of the major cheliped of the holotype of A. bunburius suggest that this appendage may be a former minor cheliped in the process of transformation into a major cheliped. This type of regeneration, i.e., transformation of a major cheliped into a minor cheliped, is not uncommon in Alpheus, often after a loss (autotomy or injury) of the original major cheliped (A. Anker, pers. obs.; see also Shin-Ike 1956; Read & Govind 1998 and references therein). In most morphological characters, except for the afore-mentioned trigonal dactylus, A. bunburius is identical to the female specimens of A. richardsoni. Bunbury, the type locality of A. bunburius, is a small town with two large rivers and an extensive saltwater lagoon connected to the open ocean by a large breach; mangroves are present in the estuary of at least one of the rivers, in Vittoria Bay. According to Banner & Banner (1982), A. richardsoni occurs in “large brackish rivers systems in Bunbury and Perth in Western Australia ”. On the other hand, A. bunburius has never been reported since its original description, which is somewhat surprising for a relatively well-sampled area. This fact, combined with the aberrant condition of the major cheliped in the holotype of A. bunburius and the great general similarity between A. bunburius and A. richardsoni, suggest that A. bunburius could also be a junior synonym of A. richardsoni. However, a side-by-side comparison of the third pereiopods of the holotype of A. bunburius and a female specimen of A. richardsoni from New South Wales revealed some differences in the shape and relative length of the dactylus (Fig. 46). In A. bunburius, the third pereiopod dactylus is feebly expanded, trigonal-conical in cross-section, and about 0.4 length of the propodus (Fig. 46A–D; see also Banner & Banner 1982: fig. 66i). In contrast, in A. richardsoni, the third pereiopod dactylus is more expanded, spatulate, and almost 0.5 length of the propodus (Fig. 46E–G; see also Banner & Banner 1982: fig. 74n). Based on this difference, A. bunburius is herein tentatively maintained as a valid species, i.e., distinct from A. richardsoni. Nevertheless, it remains a problematic taxon, being based on a single female specimen, possibly with a regenerated major cheliped, and without any information on the species’ habitat and colour pattern. It is possible that A. bunburius represents a unique western Australian transitional freshwater—brackish water form derived from A. richardsoni, like A. microrhynchus is in South-East Asia, deriving from mangrove forms, such as A. eurydactylus or A. takla sp. nov. A more intensive sampling of estuarine snapping shrimps in Bunbury and Perth areas will hopefully result in the collection of additional male and female specimens of A. bunburius, which would then allow a full redescription of the species and clarify its ecology and phylogenetic affinities., Published as part of Anker, Arthur, 2023, Revision of Alpheus euphrosyne De Man, 1897 and A. microrhynchus De Man, 1897, with description of three new species and taxonomic remarks on several other morphologically and ecologically similar snapping shrimps (Malacostraca: Decapoda: Alpheidae), pp. 1-115 in Zootaxa 5282 (1) on pages 85-86, DOI: 10.11646/zootaxa.5282.1.1, http://zenodo.org/record/7912292, {"references":["Banner, D. M. & Banner, A. H. (1982) The alpheid shrimp of Australia. Part III: The remaining alpheids, principally the genus Alpheus and the family Ogyrididae. Records of the Australian Museum, 34, 1 - 357. https: // doi. org / 10.3853 / j. 0067 - 1975.34.1982.434","Shin-Ike, T. (1956) Asymmetry in the chela of Alpheus. Annotationes Zoologicae Japonenses, 29, 79 - 85.","Read, A. T. & Govind, C. K. (1998) Cell types in regenerating claws of the snapping shrimps, Alpheus heterochaelis. Canadian Journal of Zoology, 76, 1080 - 1090. https: // doi. org / 10.1139 / z 98 - 048","Yaldwyn, J. C. (1971) Preliminary descriptions of a new genus and twelve new species of natant decapod Crustacea from New Zealand. Records of the Dominion Museum, 7, 85 - 94."]}

Alpheus nipa Banner & Banner, 1985 (Figs. 44, 45, 52D) Alpheus nipa Banner & Banner 1985: 21, fig. 2. Type material. Paratypes, 5 males (cl 6.2–7.5 mm, tl 16.0 –20.0 mm, all missing chelipeds, but detached major and minor chelipeds present in the vial), 1 ov. female (cl 9.5 mm, tl 23.5 mm, chl 12.0 mm), ZMUC CRU-7526, Indonesia, Strait of Malacca off Sumatra, vicinity of Medan, Galathea Expedition sta. 325, 4°20’N 98°53’E, palm fronds floating on water surface (depth at collection site: 40 m), 10.05.1951. Description. See Banner & Banner (1985) for detailed description and illustrations; complementary illustrations of one of the paratypes are provided in Figs. 44, 45. Colour pattern. Unknown. Type locality. Off Sumatra, Indonesia. Distribution. Indo-West Pacific: presently known only from the type locality in the Straits of Malacca off Sumatra, western Indonesia (Fig. 52D). Common name proposed. Nipa palm snapping shrimp. Ecology and biology. All known specimens of A. nipa were found on floating fronds of nipa palm (Nypa fructicans Wurmb, Arecaceae), suggesting that they may inhabit estuarine muddy banks fringed by nipa palms, a habitat that is both difficult to access and dangerous to sample due to the presence of saltwater crocodiles [Crocodylus porosus (Schneider)]. Taxonomic remarks. Banner & Banner (1985) provided a reasonably detailed description of A. nipa, accompanied by illustrations of all major diagnostic features. Two important features of A. nipa not mentioned nor illustrated by Banner & Banner (1985) are the unarmed antennal basicerite (Fig. 45B) and the presence of a faint mesial subdistal ridge on the major cheliped pollex (Figs. 44B, 45G), which may link this species to the A. euphrosyne — A. microrhynchus complex. Another previously unknown feature of A. nipa is the presence of three small, gill-like leaflets at the base of the third maxilliped exopod; this feature was observed on two dissected third maxillipeds and one maxilliped still in situ. This gill-like structure does not appear to be homologous to the “rudimentary pleurobranch at the base of the third maxilliped” in A. pontederiae (Christoffersen 1984) and “une petite pleurobranchie de forme très spéciale sur le 3 e mxp.” in several other species of Alpheus, including A. euphrosyne, A. microrhynchus and A. macrodactylus (see discussion in Coutière 1899: 279–281). As noted by Banner & Banner (1985), A. nipa can be separated from A. euphrosyne, A. richardsoni, A. paludicola and A. microrhynchus by the dactyli of the third and fourth pereiopods being short and conical rather than spatulate (in the first three species) or trigonal-subspatulate (in the latter species). This distinguishing character is also valid for A. eurydactylus, A. nomurai sp. nov., A. mangalis sp. nov., A. takla sp. nov. and A. songkla, all of which with spatulate dactyli on the ambulatory pereiopods. In addition, A. nipa differs from A. microrhynchus by the markedly different ratio of the first two carpal subarticles of the second pereiopod (1: 0.8 in A. nipa vs. 1: 0.3 in A. microrhynchus); from both A. microrhynchus and A. cyanoteles by the much stronger sculpture of the male minor chela, with both dorsal and ventral transverse grooves well pronounced and bordered by shoulders (vs. feebly delineated, if it all, and without shoulders, in A. microrhynchus and A. cyanoteles); and the scaphocerite blade much shorter than the distolateral tooth (vs. reaching slightly or far beyond the distolateral tooth in A. microrhynchus and A. cyanoteles). Another diagnostic character of A. nipa is the presence of several long erect setae on the rostrum (Fig. 45A, B; see also Banner & Banner 1985: fig. 2a, b); these setae are absent in the other species being dealt with in this study (or if present, not nearly as conspicuous as in A. nipa). Banner & Banner (1985) suggested that A. nipa “is derived from one of these euryhaline mud-or soft sedimentdwelling species, possibly from the most common and widespread A. euphrosyne * and has left the mud and silt where the other three species [A. euphrosyne *, A. richardsoni, A. paludicola] dwell and has invaded a unique habitat, the nipa palms. In so doing, it has lost the spatulate dactyli on the walking legs, so necessary for digging in soft sediments” [*note: A. euphrosyne not sensu De Man (1897)]. This is indeed an interesting hypothesis to be tested in the future, and to be contrasted to an alternative hypothesis of A. nipa not being closely related to the A. euphrosyne — A. microrhynchus complex and being derived, for instance, from the morphologically similar A. lobidens complex, in which the walking leg dactyli are also conical. Whatever the case may be, the origin and phylogenetic affinites of A. nipa are difficult to ascertain without a comprehensive phylogenetic analysis of the entire A. edwardsii group, with inclusion of as many members as possible of the A. euphrosyne — A. microrhynchus and A. lobidens complexes.

Alpheus pontederiae de Rochebrune, 1883 (Figs. 3J–M, 47, 48A–D, 52E) Alpheus Pontederiae de Rochebrune 1883: 174. Alpheus pontederiae.— Coutière 1899: 37; Holthuis 1951: 85, fig. 17; Longhurst 1958: 31, 46, 47, 49, 91; Rossignol 1962: 131; Crosnier & Forest 1965: 607; Crosnier & Forest 1966: 278, fig. 23a–j; Powell 1979: 127; Christoffersen 1980: 105, figs. 24, 25; Christoffersen 1984: 197, figs. 3, 4; Christoffersen 1998: 360; Almeida et al. 2006: 9; Almeida et al. 2012: 12; Soledade & Almeida 2013: 104, fig. 6D; Pachelle et al. 2016: 9; Soledade et al. 2017: 182, fig. 1; Almeida et al. 2018: 341. Alpheus Edwardsi. — Aurivillius 1898: 30 [not Alpheus edwardsii (Audouin, 1826)]. Alpheus megacheles.— Coutière 1899: 37 [not Alpheus megacheles Norman, 1868]. Alpheus macrocheles.— Rathbun 1900: 312; Balss 1916: 20 [not Alpheus macrocheles (Hailstone, 1835)]. Crangon langi Schmitt 1926: 20, fig. 63. Alpheus Bouvieri. — Monod 1927: 594 [not Alpheus bouvieri A. Milne Edwards, 1878]. Alpheus Langi. — Monod 1928: 252. Alpheus heterochaelis var. orientalis Vilela 1949: 55, fig. 3. Alpheus euphrosyne langi.— Banner & Banner 1982: 238. Material examined. Brazil: 1 female (cl 10.0 mm), MZUSP 21489, Pará, Bragança, Praia de Ajuruteua, Furo do Meio, sta. 6, 0052,440’S 4939,000’W, mangrove, leg. M. Tavares et al., 03.02.2010; 2 ovig. females (cl 11.3, 16.4 mm), MCZ 96843, Pará, Thayer Expedition 1865, leg. Agassiz & Bouget, 1865 (det. as Crangon lutarius); 1 male, (cl indet.), MPEG, Pará, near Bragança, leg. R.R.R. Vieira, 2016. Venezuela: 1 female (cl 19.3 mm), OUMNH. ZC. 2011.06.4, Orinoco Delta, sta. XI-52-24, leg. G. Pereira et al., date unknown; 1 male (cl 11.7 mm, missing minor cheliped), OUMNH. ZC. 2011.06.2, Orinoco Delta, sta. XI-50-37, leg. G. Pereira et al., date unknown; 1 male (cl 7.2 mm), OUMNH. ZC. 2011.06.010, north of Maracaibo, El Nazareth, leg. A. Godoy, date unknown; 1 male (cl indet.), 1 female (cl indet.), MZUSP, Orinoco Delta, further details not recorded. São Tomé and Príncipe: 1 male (cl ~6.0 mm), OUMNH. ZC. 2011.06.8, S„o Tomé Island, near Porto Alegre, mangrove, in mud under rocks, leg. A. Anker, 05.02.2006 [06-154]. Description. See de Rochebrune (1883) for original description (without illustrations), Schmitt (1926, as Crangon langi), Holthuis (1951), Crosnier & Forest (1966), Christoffersen (1984) and Soledade et al. (2017) for additional accouts and taxonomic remarks, all with illustrations; see also Fig. 47 for complementary illustrations. Colour pattern. Body uniform brownish to brown-green or green-blue; pleon with weak pale yellow markings; posterior-most part of each pleonite somewhat darker, resulting in impression of dark transverse banding; antennular and antennal flagella pale greenish or yellowish; mesial face of major greenish or bluish brown with some blue-grey, purplish and paler yellow areas; fingers darker, more olive-green, with contrasting pink fingertips; mesial subdistal ridge on pollex conspicuously blue; dactylar plunger whitish with green-yellow tinge and blue spot; minor chela greenish or bluish, with darker fingers; walking legs pale reddish; telson marbled with whitish; uropods whitish proximally, mottled with brown or grey distally, exopod darker brown with blue tinge posterior to transverse suture (Fig. 48A–D); see also Soledade & Almeida (2013: fig. 6D). Type locality. Mouth of the rivers Leybar, Thiank and Dakar-Bango, Senegal. Distribution. Eastern and western Atlantic, from Senegal to Congo, and from Venezuela to southern Brazil (Fig. 52E); specific localities include, in the eastern Atlantic: Senegal (Leybar, Thiank, Dakar-Bango), Guinea (Conakry), Nigeria (Niger delta), Cameroon (Bibundi, Souelaba, Malimba Bay), Pop. Rep. Congo (Pointe Noire), Dem. Rep. Congo (Banana), Equatorial Guinea (Passo, Canchungo, Rubane Island), S„o Tomé (Schmitt 1926; Vilela 1949; present study); in the western Atlantic: Venezuela (Orinoco Delta), Suriname, Brazil (Pará, Maranh„o, Ceará, Paraíba, Alagoas, Bahia, S„o Paulo, Paraná) (Christoffersen 1984, 1998; Almeida et al. 2006, 2012; Soledade & Almeida 2013; Pachelle et al. 2016; Soledade et al. 2017; present study). Common name proposed. Rochebrune’s snapping shrimp. Ecology and biology. Alpheus pontederiae is mainly an estuarine snapping shrimp, inhabiting mudflats and mangrove channels close to river mouths and deltas, often in brackish water (salinity range: 30 / 00 – 200 / 00) (see also Fig. 48E). The species lives in deep galleries built in mud or muddy sand, sometimes under rocks, decaying wood or other debris, usually in the intertidal zone, but also in deeper water down to 30 m (Schmitt 1926; Monod 1927; Holthuis 1951; Christoffersen 1984; Almeida et al. 2012; Soledade & Almeida 2013; Pachelle et al. 2016). In West Africa, A. pontederiae is often found close to burrows of mangrove-dwelling crabs, viz. Panopeus africanus A. Milne-Edwards and Sarmatium curvatum H. Milne Edwards, whilst young shrimps occasionally dwell in decomposing mangrove wood, together with the mud shrimps Upogebia furcata (Aurivillius) (Aurivillius 1898; Schmitt 1926). According to de Rochebrune (1883), the type specimen was found “between floating clusters of the marine herb Eichhornia natans (P. Beauv.) [= Pontederia natans] in rivers of Senegal at the time when their water is saltish” (Holthuis 1951). In some areas of eastern Brazil, A. pontederiae also occurs in mangrove oyster [Crassostrea rhizophorae (Guilding)] beds (Almeida et al. 2012). The large number of eggs and their small size suggest that this species has an extended larval development. Taxonomic remarks.Several morphological features link A.pontederiae to the A.euphrosyne — A.microrhynchus complex. These are the general shape of the chelipeds, including the presence of a well-developed mesial subdistal ridge on the major chela pollex (Fig. 47B, D; Christoffersen 1984: fig. 4a) and the male minor chela with strongly balaeniceps fingers (Fig. 47G, H; Christoffersen 1984: fig. 4c, d); and the third and fourth pereiopods with broadly spatulate dactyli (Christoffersen 1984: fig. 4g). The Indo-West Pacific species, which are morphologically closest to A. pontederiae, are A. eurydactylus and A. takla sp. nov. The Atlantic species differs from A. eurydactylus by the mesial face of the major chela non-granulated (vs. granulated on the distal portion of palm and pollex in A. eurydactylus) and the antennal basicerite distolaterally armed with a strong tooth (vs. unarmed in A. eurydactylus); and from A. takla sp. nov. by the slenderer third pereiopod, with its merus almost seven times as long as wide (vs. five times in A. takla sp. nov.) and ischium armed with a stout spiniform seta (vs. usually unarmed in A. takla sp. nov.). In addition, A. pontederiae, A. eurydactylus and A. takla sp. nov. have differently coloured plungers of the major chela dactylus: whitish, tinged with green-yellow and a blue spot in A. pontederiae; white and blue proximally and orange-yellow distally in A. eurydactylus; and bright red-orange in A. takla sp. nov. (cf. Figs. 16C, 32A, 35A, 48C). Alpheus pontederiae has a disjunct amphi-Atlantic distribution (Fig. 52E), being absent from the islands of the Central Atlantic, probably due to the lack of suitable habitats there. Interestingly though, A. pontederiae also seems to be absent from the Caribbean Sea and Gulf of Mexico / Florida region, where estuarine and mangrove habitats are plentiful; the western-most record of this species is the Orinoco Delta in Venezuela. A genetic comparison of the western and eastern Atlantic populations of A. pontederiae, similar to the one recently performed for A. intrinsecus Spence Bate, 1888 (Cunha et al. 2017), is highly desirable. In this context, it is noteworthy that A. pontederiae was reported to be variable in the relative length of the antennal scaphocerite blade, which ranges from being much shorter than the adjacent distolateral tooth to reaching distinctly beyond it (Crosnier & Forest 1966; Christoffersen 1984)., Published as part of Anker, Arthur, 2023, Revision of Alpheus euphrosyne De Man, 1897 and A. microrhynchus De Man, 1897, with description of three new species and taxonomic remarks on several other morphologically and ecologically similar snapping shrimps (Malacostraca: Decapoda: Alpheidae), pp. 1-115 in Zootaxa 5282 (1) on pages 87-90, DOI: 10.11646/zootaxa.5282.1.1, http://zenodo.org/record/7912292, {"references":["Rochebrune, A. T. de (1883) Diagnoses d'arthropodes nouveaux propres la Senegambie. Bulletin de la Societe Philomathique de Paris, Serie 7, 7, 167 - 182.","Coutiere, H. (1899) Les \" Alpheidae \". Morphologie externe et interne, formes larvaires, bionomie. Annales des Sciences Naturelles, Zoologie et Paleontologie, Serie 8, 9, 1 - 559, pls. 1 - 6. Masson, Paris. https: // doi. org / 10.5962 / bhl. title. 13143","Holthuis L. B. (1951) The caridean Crustacea of tropical West Africa. Atlantide Report, 2, 1 - 187.","Longhurst, A. R. (1958) An ecological survey of the West African marine benthos. Colonial Office, Fisheries Publication, 11, 1 - 102.","Rossignol, M. 1962. Catalogue des crustaces decapodes brachyoures, anomoures et macroures littoraux en collection au Centre d'Oceanographie de Pointe-Noire. Cahiers de l'ORSTOM, Oceanographie, 2, 111 - 138.","Crosnier, A. & Forest, J. (1965) Note preliminaire sur les Alpheidae recuellis par la Calypso dans l'Atlantique oriental tropical (Crustacea Decapoda Natantia). Bulletin du Museum National d'Histoire Naturelle, Paris, Serie 2, 36, 602 - 610.","Crosnier, A. & Forest, J. (1966) Crustaces Decapodes: Alpheidae. In: Campagnes de la Calypso dans le Golfe de Guinee et aux Iles Principe, S \" o Tom et Annobon (1956), et Campagne aux Iles du Cap Vert (1959). Part 19. Resultats Scientifiques des Campagnes de la Calypso. 7 (27). Annales de l'Institut Oceanographique de Monaco, 44, 199 - 314. https: // doi. org / 10.5962 / bhl. title. 13143","Powell, C. B. (1979) Three alpheid shrimps of a new genus from West African fresh and brackish waters: taxonomy and ecologial zonation (Crustacea Decapoda Natantia). Revue de Zoologie Africaine, 93, 116 - 150.","Christoffersen, M. L. (1980) Taxonomia e distribuicao geografica dos Alpheoidea (Crustacea, Decapoda, Natantia) do Brasil, Uruguay e norte da Argentina, incluindo consideracles sobre a divisao do sul do continente em provincias biogeograficas marinhas. PhD Thesis, University of S \" o Paulo, S \" o Paulo, 467 pp.","Christoffersen, M. L. (1984) The western Atlantic snapping shrimps related to Alpheus heterochaelis Say (Crustacea, Caridea) with the description of a new species. Papeis Avulsos de Zoologia, 35, 189 - 208.","Christoffersen M. L. (1998) Malacostraca. Eucarida. Caridea. Crangonoidea and Alpheoidea (Except Glyphocrangonidae and Crangonidae). In: Young, P. S. (Ed.), Catalogue of Crustacea of Brazil. Serie Livros N. 6. Museu Nacional, Rio de Janeiro, pp. 351 - 372.","Almeida, A. O. de, Coelho, P. A., Santos, J. T. A. dos & Ferraz, N. R. (2006) Crustaceos decapodos estuarinos de Ilheus, Bahia, Brasil. Biota Neotropica, 6 (2), bn 03406022006, 1 - 24. https: // doi. org / 10.1590 / S 1676 - 06032006000200024","Almeida, A. O. de, Boehs, G., Araujo-Silva, C. L. & Bezerra, L. E. A. (2012) Shallow-water caridean shrimps from southern Bahia, Brazil, including the first record of Synalpheus ul (Rios & Duffy, 2007) (Alpheidae) in the southwestern Atlantic Ocean. Zootaxa, 3347 (1), 1 - 35. https: // doi. org / 10.11646 / zootaxa. 3347.1.1","Soledade, G. O. & Almeida, A. O. de (2013) Snapping shrimps of the genus Alpheus Fabricius, 1798 from Brazil (Caridea: Alpheidae): updated checklist and key for identification. Nauplius, 21, 89 - 122. https: // doi. org / 10.1590 / S 0104 - 64972013000100010","Pachelle, P. P. G., Anker, A., Mendes, C. B. & Bezerra, L. E. A. (2016) Decapod crustaceans from the state of Ceara, northeastern Brazil: an updated checklist of marine and estiarine species, with 23 new records. Zootaxa, 4131 (1), 1 - 63. https: // doi. org / 10.11646 / zootaxa. 4131.1.1","Soledade, G. O., Oliveira, M. V. & Almeida, A. O. de (2017) A specimen of the snapping shrimp Alpheus pontederiae de Rochebrune, 1883 with symmetric chelipeds. Spixiana, 40, 181 - 184.","Aurivillius, C. W. S. (1898) Krustaceen aus dem Kamerun-Gebiete. Bihang till Kungliga Svenska Vetenskapsakademiens Handlingar, 24, 1 - 31, pls. 1 - 4.","Audouin, V. (1826) Explication sommaire des planches de Crustaces de l'Egypte et de la Syrie, publiees par Jules-Cesar Savigny, Membre de l'Institut; offrant un expose des characteres naturels des genres avec la distinction des especes. Animaux invertebres. In: Savigny, J. C. (Ed.), Description de l'Egypte ou receuil des observations et des recherches qui ont et faites en Egypte pendant l'expedition de l'armee francaise, publie par les orders de sa Majeste l'Empereur Napoleon le Grand. Imprimerie Imperiale, Paris, pp. 77 - 98.","Norman, A. M. (1868) On the British species of Alpheus, Typton, and Axius, and on Alpheus Edwardsii of Audouin. Annals and Magazine of Natural History, Series 4, 2, 173 - 178. https: // doi. org / 10.1080 / 00222936808695776","Rathbun, M. J. (1900) The decapod crustaceans of West Africa. Proceedings of the United States National Museum, 22, 271 - 316. https: // doi. org / 10.5479 / si. 00963801.22 - 1199.271","Balss, H. (1916) Crustacea II: Decapoda Macrura und Anomura (ausser Fam. Paguridae). In: Michaelsen, W. (Ed.), Beitrage der Meeresfauna Westafrikas. L. Friederichsen & Co., Hamburg, pp. 13 - 46.","Hailstone, S. Jr. (1835) Notices of another species of Pontophilus, and of a crustacean allied to the genus Hippolyte. The Magazine of Natural History and Journal of Zoology, Botany, Mineralogy, Geology, and Meteorology, 8, 270 - 273.","Schmitt, W. L. (1926) The macruran, anomuran and stomatopod crustaceans collected by the American Museum Congo Expedition, 1909 - 1915. Bulletin of the American Museum of Natural History, 53, 1 - 67, pls. 1 - 9.","Monod, T. (1927) Crustacea IV. Decapoda (excl. Palaemonidae, Atyidae et Potamomdae). In: Monod, T. (Ed.), Contribution a l'etude de la faune du Cameroun. Faune des Colonies Francaises, 1, 593 - 624.","Monod, T. (1928) Additions a ma liste des decapodes marins du Cameroun. Bulletin du Museum d'Histoire Naturelle, 34, 252.","Vilela, H. (1949) Crustaceos decapodes e estomatopodes da Guine Portuguesa. Anais da Junta de Investigacles Coloniais, Lisboa, 4, 47 - 70.","Banner, D. M. & Banner, A. H. (1982) The alpheid shrimp of Australia. Part III: The remaining alpheids, principally the genus Alpheus and the family Ogyrididae. Records of the Australian Museum, 34, 1 - 357. https: // doi. org / 10.3853 / j. 0067 - 1975.34.1982.434","Spence Bate, C. (1888) Report on the Crustacea Macrura collected by the Challenger during the years 1873 - 76. In: Report on the Scientific Results of the Voyage of H. M. S. \" Challenger during the years 1873 - 76. Vol. 24. Eyre & Spottiswoode, London, pp. i - xc + 1 - 942, pls. 1 - 157.","Cunha, A. M., Terossi, M., Mantelatto, F. L. & Almeida, A. O. de (2017) Morphological and molecular analyses support the amphi-Atlantic distribution and taxonomic status of the snapping shrimp Alpheus intrinsecus Spence Bate, 1888 (Crustacea: Decapoda: Alpheidae). Zootaxa, 4303 (4), 573 - 589. https: // doi. org / 10.11646 / zootaxa. 4303.4.8"]}

Alpheus mangalis sp. nov. (Figs. 36–40, 52B) (?) Alpheus sp. cf. audouini .— Johnson 1962: 53 (part.). (?) Alpheus microrhynchus. — Johnson 1965: 9 (part.) [not A. microrhynchus De Man, 1897]. (?) Alpheus euphrosyne. — Banner & Banner 1966: 130 (part.?) [not A. euphrosyne De Man, 1897]. (?) Alpheus euphrosyne euphrosyne. — Banner & Banner 1982: 232 (part.?, not fig. 73); Chace 1988: 27 (part.?) [not A. euphrosyne De Man, 1897]. Type material. Holotype, male (cl 11.4 mm, tl 33.5 mm, chl 17.5 mm), ZRC 2014.0667, Singapore, Pulau Ubin, Kampung Melayu, mudflat near mangrove, leg. B.Y. Lee & B.W. Lee, 26.09.2011. Paratypes: 1 ov. female (cl 12.0 mm), ZRC 2014.0683, same collection data as for holotype; 2 males (cl 7.5, 7.7 mm), 2 females (cl 7.0, 8.3 mm), 1 ov. female (cl 7.1 mm), 1 juv. male (cl indet.), ZRC 2014.0685, Singapore, Pulau Ubin, OBS Camp 2, mouth of Sungei Teris, sta. SW77, mud, gill net, leg. H.H. Ng & M. R. bin Duria, 24.10.2012; 5 males (cl 7.3–9.4 mm), 3 ov. females (cl 8.2–11.0 mm), OUMNH. ZC. 2019.06.60, Singapore, Pulau Ubin, Chek Jawa, mangrove, yabby pump, leg. K.L. Yeo, 21.08.2001; 1 male (cl 10.4 mm), 1 ov. female (cl 10.5 mm), ZRC 2014.0666, same collection data as for previous paratypes; 5 males (cl 7.6–11.3 mm), 1 female (cl 9.2 mm), 4 ov. females (cl 9.4–11.5 mm), MNHN-IU-2018-5732, Singapore, Pulau Ubin, between OBS camps 1 and 2, sta. M04, mudflat, edge of mangrove, leg. J.C. Mendoza, Y.L. Lee & R. Tan, 06.03.2012; 1 male (cl 10.5 mm), 1 ov. female (cl 12.8 mm), ZRC 2014.0684, same collection data as for previous specimens [M04-141]; 2 males (cl 8.0, 8.1 mm), MNHN-IU-2018-5750, Singapore, Pulau Ubin, west of Celestial Resort, sta. M01, mangrove, mounds of Thalassina sp., mud combing, leg. D.L. Rahayu & K.S. Tan, 06.03.2012; 1 male (cl 7.7 mm), 1 female (cl 7.1 mm), OUMNH. ZC. 2019.06.61, Singapore, mouth of Sungei Besar, sta. SW95, sandy shore with mudflat and mangrove, leg. K.S. Koh et al., 26.10.2012; 1 male (cl 9.3 mm), MNHN-IU-2018-5743, same collection data as for previous specimens [JS-2430]; 1 male (cl 10.2 mm), MNHN-IU-2018-5592, same collection data as for previous specimens [JS-2429];> 40 specimens of both sexes (cl indet.), ZRC 2014.0678, Singapore, Sungei Buloh, sta. KR3, mangrove, leg. Y.L. Teo & N. Sivasothi, 12.03.2004 [2011-SIN-082]; 2 males (cl 8.9, 9.3 mm), 2 ov. females (cl 9.8, 10.0 mm), MNHN-IU-2018-5600, same collection data as for previous specimens; 1 male (cl 9.8 mm), MNHN-IU-2018-5749, Singapore, Lim Chu Kang, mangrove, sta. SW106, mud, hand and tangle nets, leg. A. Anker, B.Y. Lee & D. Uyeno, 28.10.2012 [JS-2414]. Additional material. Singapore: 1 male (cl 6.0 mm), 2 females (cl 5.5, 6.5 mm), ZRC 2014.0679, Pulau Ubin, 01°24.340’N 103°58.253’, leg. K.S. Tan et al., 12.02.2011 [4017-4019]; 2 males (cl 6.6, 9.3 mm), ZRC 2014.0721, Sungei Simpang, sta. MF49, leg. H.H. Ng, 17.02.2012 [49011-49012]; 1 ov. female (cl indet.), OUMNH. ZC. 2019.06.62, Sungei Buloh, Left Main Stream, sta. KR4, mangrove, leg. Y.L. Teo, 05.03.2004 [2011-SIN-031]; 1 female (cl 10.3 mm, without rostrum), MNHN-IU-2018-5747, same collection data as for pervious specimen [2011-SIN-025]; 1 female (cl 11.2 mm), OUMNH. ZC. 2019.06.62, Sungei Buloh, sta. KR3, mangrove, leg. Y.L. Teo, 13.02.2004 [2011-SIN-036]; 1 ov. female (cl 9.5 mm, without rostrum), OUMNH. ZC. 2019.06.63, Sungei Buloh, sta. KR3, mangrove, leg. Y.L. Teo, 12.03.2004 [2011-SIN-047]; 2 males (cl 7.3, 7.7 mm), OUMNH. ZC. 2019.06.64, Sungei Buloh, mangrove, leg. Y. Cai, 19.04.1997 [2011-SIN-061];> 20 specimens of both sexes (cl indet.), MNHN-IU-2018-5753, Sungei Buloh, East Channel, mangrove, leg. Y.L. Teo, 13.02.2004 [2011-SIN-078];> 10 specimens of both sexes (cl indet.), FLMNH UF 68863, Sungei Buloh, East Channel, mangrove, leg. Y.L. Teo & K.M. Sheng, 21.02.2004; 2 males (cl 6.5, 8.9 mm), 1 female (cl 8.7 mm), 1 ov. female (cl 7.9 mm), ZRC 1996.44, Sungei Buloh, mangrove, leg. P.K.L. Ng et al., 05.08.1995; 1 male (cl 8.0 mm), 1 ov. female (cl 8.4 mm), ZRC 2014.0720, Sungei Buloh, mangrove, leg. C. Schubart, 14.01.2002; 2 males (cl 7.4, 8.0 mm), 1 female (cl 7.4 mm), 4 ov. females (cl 8.7–9.2 mm), ZRC 2004.0500, Sarimbun-Poyan mangrove, leg. H.H. Tan et al., 13– 20.01.2004; 1 female (cl 8.0 mm), 3 ov. females (cl 8.7–9.0 mm), ZRC 2014.0688, Sarimbun Beach, near Jalan Bahtera Scouts Camp, mudflat, H.H. Ng, Y.L. Teo & H. Wong, 14.02.2012 [48198-48201]; 2 males (cl 7.3, 7.4 mm), ZRC 1996.43, Lim Chu Kang, mangrove, leg. P.K.L. Ng, 18.08.1995; 1 ov. female (cl 9.2 mm), ZRC 2014.0691, Singapore, Lim Chu Kang, mangrove, sta. Y595, 04.04.1989; 1 male (cl 7.5 mm), ZRC 1989.3661, Lim Chu Kang, mangrove, in mound of Thalassina sp., leg. P.K.L. Ng, 17.03.1987; 1 male (cl 6.8 mm), 1 ov. female (cl 7.9 mm), ZRC 2014.0689, Lim Chu Kang—Sarimbun Scouts Camp, mangrove, leg. H. Wong et al., 21.07.2011 [27136–27137]; 1 male (cl 6.7 mm), 1 female (cl 5.5 mm), ZRC 2014.0692, same collection data as for previous specimens [27138–27139]; 1 female (cl 8.0 mm), ZRC 2014.0687, Lim Chu Kang, sta. SW110, mangrove and adjacent mud flat, B.Y. Lee et al., 28.X.2012 [JS-2395]; 1 male (cl 6.3 mm), ZRC 2014.0693, Lim Chu Kang, sta. SW106, mangrove and adjacent mudflat, leg. A. Anker et al., 27.10.2012; 3 males, 2 females, 1 ov. female (cl. indet.), MNHN-IU-2013-14787, Lim Chu Kang mangrove, leg. A. Anker et al., 01.2002; 3 males (cl 6.1–7.3 mm), ZRC 2014.0690, Loyang, East coast of river mouth, mud, leg. TMSI team, 11.07.2012 [67156–67158]; 1 male (cl 11.3 mm), 1 ov. female (cl 12.6 mm), ZRC 1996.45, Mandai, mangrove, leg. P.K.L. Ng, 21.06.1990; 1 male (cl 9.2 mm), 1 ov. female (cl 8.5 mm), MNHN-IU-2013-14785, Pandan mangrove, low tide, in pairs under logs in mud, leg. A. Anker, 13.02.2002; 1 male (cl 8.3 mm), 1 female (cl 7.9 mm), ZRC 1979.4.3.1–2, Jurong, prawn ponds, sta. B17, leg. S. R.F. R.S., 01.06.1954 [J8098-8098]; 2 males (cl 7.8, 10.2 mm), 2 ov. females (cl 9.5, 10.4 mm), ZRC 1979.4.3.3-6, Jurong, prawn ponds, sta. B29, leg. S. R.F. R.S., 05.07.1954 [J8086-8089]. Malaysia: 1 ov. female (cl 11.3 mm), ZRC 1979.4. 3.8, Selangor, Port Klang (Port Swettenham), mudflats, leg. S. Kumar, 27.08.1968; 2 ov. females (cl 11.2, 12.0 mm), MNHN-IU-2018-5745, Johor, Muar, sta. 2, mudflats, leg. N.K. Ng. & J.C.Y. Lai, 10.02.2009; 1 male (cl 7.7 mm), MNHN-IU-2013-14786, Tanjong Sendili, leg. Y. Cai & N.K. Ng, 20.09.1998; 1 male (cl 6.2 mm), ZRC 1999.0571, Sarawak, Bako, Telok Delima, leg. K.L. Yeo, 29.06.1994; 1 male (cl 9.1 mm), 1 ov. female (cl 8.3 mm), ZRC 2014.0669, Sarawak, Kuching prawn farm, no further data. Indonesia: 2 males (cl 10.2, 12.6 mm), 2 ov. females (cl 9.3, 14.4 mm, latter tl ~43.0 mm), MNHN-IU-2018- 5733, Java, Jepara, leg. D.L. Rahayu, 04.2010; 1 male (cl 9.2 mm), OUMNH. ZC. 2019.06.65, Lombok, Lembar, prawn ponds, fine mud, burrows in banks of brackish creek, low tide, leg. A. Anker & D.L. Rahayu, 14.05.2014; 1 male (cl 12.3 mm), 1 female (cl 13.0 mm), QM W25803-2, Papua, Ajkwa River estuary, 4°50’S 136°50’E, Environmental Laboratory, PT Freeport, Sta. Ajk-36, estuarine habitat, 30.03.2000; 5 males (cl 8.0– 13.6 mm), 1 ov. female (cl 9.7 mm), MNHN-IU-2018-5740, Papua, Ajkwa Island, mangrove, leg. A. Darmawan et al., 20.07.2012; 1 male (cl 7.4 mm, missing minor cheliped), ZRC 2014.0722, same collection data as for previous specimens; 3 males (cl 8.8–9.5 mm), 1 female (cl 11.4 mm), MNHN-IU-2018-5736, Papua, Kamora, mangrove, leg. A. Darmawan et al., 07.08.2012. Philippines: 1 juv. female (cl 5.7 mm, both chelipeds detached, other pereiopods missing, identification somewhat tentative), ZRC 2014.0681, Palawan, Sabang, Underground River National Park, brackish river near cave entrance, leg. H. Freitag, 16.02.2001. Taiwan: 1 male (cl 8.2 mm, missing minor cheliped), ZRC 2000.2132, Shinzhu, mangrove, leg. Y. Cai & C.W. Lin, 28.11.1997. Australia: 2 males (cl 10.5, 12.1 mm), 1 ov. female (cl 11.7 mm), NTM Cr. 008307, Northern Territory, Darwin, Ludmilla Creek mouth, 1224.8’S 13051.7 E, undisturbed mangrove, low tide, leg. M. Burke, 04.06.1991; 1 male (cl 11.2 mm), NTM Cr. 008317, Northern Territory, Darwin, Ludmilla Creek mouth, 12°24.8’S 130°51.7’ E, disturbed mangrove, low tide, leg. M. Burke, 05.06.1991; 4 males (cl 13.0– 13.4 mm), 1 female (cl 14.0 mm), 1 ov. female (cl 14.4 mm), NTM Cr. 008305, Northern Territory, Darwin, mouth of Blackmore River, Channel Island, 12°37.3’S 130°56.6’ E, undisturbed mangrove, low tide, leg. M. Burke, 13.06.1991; 1 ov. female, NTM Cr. 013093, Northern Territory, Darwin, mouth of Blackmore River, Channel Island, 12°37.3’S 130°56.6’ E, undisturbed mangrove, low tide, leg. M. Burke, 12.06.1991; 1 male (cl indet., missing both chelipeds), NTM Cr. 013094, Northern Territory, Darwin, mouth of Blackmore River, Channel Island, 12°37.3’S 130°56.6’ E, undisturbed mangrove, low tide, leg. M. Burke, 17.06.1991; 1 male, 1 female (cl indet.), NTM Cr. 008305, Northern Territory, Darwin, Channel Island, near bridge, 12°34.9’S 130°55.4’ E, undisturbed mangrove, low tide, leg. M. Burke, 02.07.1991. Tentative identification. Alpheus cf. mangalis sp. nov. New Caledonia: 2 males (cl 10.2, 11.6 mm), 1 ov. female (cl 11.6 mm), MNHN-IU-2019-3217, Dumbéa River estuary, mangrove, leg. C. Vadon, 22.09.1978. Description. See Figs. 36–38. Medium-sized species of Alpheus (maximal cl 14.4 mm, tl ~43.0 mm). Carapace glabrous, without pubescence, with slight grooves. Rostrum short, triangular, distally acute, reaching or slightly overreaching mid-length of first article of antennular peduncle, occasionally minute or (rarely) absent; rostral carina present as low elevation, fading at eye level; rostro-orbital furrows poorly demarcated. Pterygostomial angle broadly rounded; cardiac notch deep. Pleon smooth, without pubescence. Telson moderately broad, ovate-rectangular, gently tapering towards posterior margin, about 1.5 times as long as maximal width near proximal margin; lateral margins slightly convex; dorsal surface without trace of median depression, with two pairs of short spiniform setae inserted in deep pits at some distance from lateral margins, first pair at or just before telson mid-length, second pair at about 0.7 of telson length; posterior margin broadly rounded, with two pairs of slender spiniform setae at posterolateral angles, lateral much shorter than mesial. Antennular peduncle with stylocerite broad, markedly convex laterally, with acute tip, latter reaching, but not overreaching distal margin of first article; ventromesial carina with broad blunt tooth; second article about 2.3 times as long as wide. Antennal peduncle with basicerite armed with strong sharp tooth; carpocerite reaching slightly beyond scaphocerite; scaphocerite with lateral margin nearly straight; blade broad, separated from broad distolateral tooth by deep cleft; anterior margin of blade broadly rounded, exceeded by sharp distolateral tooth. Third maxilliped with antepenultimate article not particularly broadened; penultimate article moderately long, widening distally, about three times as long as maximal width; ultimate article with dense transverse rows of short serrulate setae on mesial surface and some longer setae near apex; coxal lateral plate bluntly protruding dorsally; exopod distinctly overreaching end of antepenultimate article. Major cheliped of A. edwardsii - type. Merus stout, about twice as long as wide; ventromesial margin distally armed with strong sharp tooth. Carpus very short, cup-shaped. Chela large, heavy, with fingers about 0.6 length of palm; all surfaces smooth, without granulation. Mesial face with weak mesial transverse ridge extending obliquely from proximal edge of mesial longitudinal groove to ventral surface of palm, although far from reaching it. Lateral face of palm with lateral longitudinal groove deep, subrectangular; proximal half without distinct transverse ridge. Dorsal shoulder gently sloping into broad transverse groove with angle of about 45°. Ventral shoulder pronounced, slightly protruding in lateral view, broadly rounded. Pollex with mesial face lacking subdistal mesial ridge and adjacent proximal protuberance; distomesial angle distinctly superior to 90°, blunt; lateral face with distolateral angle wide, poorly marked. Dactylus without dorsal ridge; dactylar plunger stout, elongate, with anterior margin clearly demarcated from ventral margin of dactylus; distal surface of plunger rounded anteriorly, otherwise flattened. Adhesive discs small. Male minor cheliped with chela strongly balaeniceps. Merus distinctly slenderer than that of major cheliped, almost three times as long as wide; ventral margin with distomesial angle armed with stout, sharp tooth, as in major cheliped. Chela feebly swollen, slender in lateral view, with fingers about 0.8 length of palm; all surfaces smooth, without granulation. Palm subcylindrical, compressed, feebly swollen; mesial and lateral longitudinal grooves poorly marked; dorsal shoulder poorly developed, present only as low elevation gently sloping into broad dorsal transverse groove; ventral transverse groove and adjacent shoulder similarly poorly developed, recognisable as shallow sinus and low bump, respectively. Pollex with mesial surface bearing row of balaeniceps setae on low crest extending from base to about mid-length of pollex; lateral surface with similar row of balaeniceps setae as on mesial surface. Dactylus with marked lateral expansion (balaeniceps expansion) in proximal half, about twice as long as maximal width; lateral and mesial dactylar ridges each with dense rows of balaeniceps setae. Cutting edges of both fingers shallowly excavated on each side of sharp, blade-like ridge. Female minor cheliped simple, not balaeniceps. Chela slender, not particularly swollen, with fingers subequal to palm; all surfaces smooth, not granulated. Palm with mesial longitudinal groove not distinct; lateral longitudinal groove distinct, but very shallow; dorsal and ventral transverse grooves and shoulders absent. Fingers simple, with sharp cutting edges, not gaping when closed; fingertips moderately curved and crossing. Second pereiopod with ratio of carpal subarticles approximately equal to 4: 2.1: 1: 1: 1.5. Third pereiopod relatively slender; ischium armed with stout spiniform seta on ventrolateral surface; merus about 5.2 times as long as maximal width, unarmed; propodus with five or six short spiniform setae on ventral margin and one pair of spiniform setae on distoventral margin adjacent to dactylus; dactylus almost half-length of propodus, spatulate, moderately expanded, slightly wider than propodus in dorsal view, with row of setae along mesial margin. Fourth pereiopod generally similar to third pereiopod, shorter and slenderer. Fifth pereiopod much slenderer than third and fourth pereiopods; ischium unarmed; propodus with single spiniform seta at mid-length, one distal spiniform seta adjacent to dactylus, and well-developed setal brush; dactylus subspatulate, about 0.4 length of propodus. Male second pleopod with appendix masculina 0.8 length of appendix interna. Uropod with each protopodal lobe ending in subacute tooth distally; exopod and endopod broad, ovate; diaeresis of exopod complete, almost straight, with broadly subtriangular lobe near small distolateral spiniform seta and adjacent distolateral tooth; lateral margin of exopod broadly convex. Eggs numerous (> 100 in larger females), small; egg diameter about 0.51 0.55 mm to 0.55 0.60 mm. Colour pattern. Body (carapace and pleon) uniform dark olive-green, brown-green or green-blue; smaller individuals with broad transverse bands on pleon, one per pleonite; antennules and antennae greenish blue; flagella dull greenish; mesial face of major chela dark olive-green to blue-green; lateral face of major chela whitish or bluish; distal portion of both fingers pink; dactylar plunger greenish or pale green-yellow; minor chela dark bluish green; second to fifth pereiopods reddish; pleopods dull greenish or bluish; telson and uropods dark blue-green, uropodal exopod with deep blue area posterior to transverse suture (Figs. 39, 40). Type locality. Pulau Ubin, Singapore (collection locality of the holotype). Distribution. Tropical Indo-West Pacific from Taiwan to northern Australia (Fig. 52B); specific localities include: Taiwan (Shinzhu), Philippies (Palawan), Singapore (Pulau Ubin, Lim Chu Kang, Sungei Buloh, Jurong, etc.), Indonesia (Lombok, Papua), Malaysia (Selangor, Johor, Sarawak), and Australia (Northern Territory); possibly extending to New Caledonia (Dumbéa), but see below (closely related species?). Etymology. The new species’s name refers to its typical habitat, mudflats within or near mangrove forests; used as an adjective. Common name proposed. Lesser mangrove snapping shrimp. Ecology and biology. Alpheus mangalis sp. nov. is one of the most common snapping shrimps on mangrove mudflats of Singapore, Malaysia and Indonesia. For instance, in Sungei Buloh and Pulau Ubin, Singapore, A. mangalis sp. nov. may be easily collected under large rocks embedded in mud, among mangrove roots, under wood logs, debris, etc. Like many other species of Alpheus associated with mangroves, this species appears to be euryhaline, living in marine water, but also tolerating brackish conditions. The large number of eggs and their small size suggest an extended larval development in A. mangalis sp. nov. Taxonomic remarks. Alpheus mangalis sp. nov. can be easily separated from A. euphrosyne, A. eurydactylus, A. richardsoni, A. microrhynchus, A. cyanoteles, A. nomurai sp. nov. and A. takla sp. nov. by the presence of a stout, sharp tooth on the distomesial margin of the cheliped meri (Figs. 36E, G, 38B, G; this tooth is absent in all the afore-mentioned species, cf. Figs. 2E, 7C, G, 15B, G, 27B, etc.), and the longer and/or differently shaped dactylar plunger of the major chela (Figs. 38D, 40A, E; cf. Figs. 7D, 15D, 24D, 27D, 31C; Banner & Banner 1982: fig. 74e). In addition, A. mangalis sp. nov. differs from A. euphrosyne and A. eurydactylus by the smooth, non-granulated major chela surface (granulated in the other two species); from A. euphrosyne, A. eurydactylus, A. microrhynchus, A. cyanoteles, A. nomurai sp. nov. and A. takla sp. nov. by the absence of a subdistal mesial ridge on the major chela pollex and the absence of dorsal and ventral shoulder on the minor chela (present in all the other species, although less pronounced in A. nomurai sp. nov.); from A. euphrosyne by the non-overhanging dorsal shoulder of the major chela (overhanging in A. euphrosyne); from A. richardsoni by the absence of ventral sinus on the male minor chela (present in A. richardsoni); from A. microrhynchus and A. cyanoteles by the spatulate dactyli of the third to fi, Published as part of Anker, Arthur, 2023, Revision of Alpheus euphrosyne De Man, 1897 and A. microrhynchus De Man, 1897, with description of three new species and taxonomic remarks on several other morphologically and ecologically similar snapping shrimps (Malacostraca: Decapoda: Alpheidae), pp. 1-115 in Zootaxa 5282 (1) on pages 67-75, DOI: 10.11646/zootaxa.5282.1.1, http://zenodo.org/record/7912292, {"references":["Johnson, D. S. (1962) A synopsis of the Decapoda Caridea and Stenopodidea of Singapore, with notes on their distribution and a key to the genera of Caridea occuring in Malayan waters. Bulletin of the National Museum of Singapore, 30, 44 - 79, pl. 2.","Johnson, D. S. (1965) A review of brackish water prawns of Malaya. Bulletin of the National Museum of Singapore, 33, 7 - 11.","De Man, J. G. (1897) Bericht u ¨ ber die von Herrn Schiffscapitan Storm zu Atjeh, an den westlichen Ku ¨ sten von Malakka, Borneo und Celebes sowie in der Java-See gesammelten Decapoden und Stomatopoden. Fu ¨ nfter Theil. Z oologische Jahrbu ¨ cher. Abtheilung fu ¨ r Systematik, Geographie und Biologie der Thiere, 9, 725 - 790, pls. 34 - 37 (in volume 10).","Banner, A. H. & Banner, D. M. (1966) The alpheid shrimp of Thailand. The Siam Society Monograph Series, 3, 1 - 168.","Banner, D. M. & Banner, A. H. (1982) The alpheid shrimp of Australia. Part III: The remaining alpheids, principally the genus Alpheus and the family Ogyrididae. Records of the Australian Museum, 34, 1 - 357. https: // doi. org / 10.3853 / j. 0067 - 1975.34.1982.434","Chace, F. A. Jr. (1988) The caridean shrimps (Crustacea: Decapoda) of the Albatross Philippine Expedition, 1907 - 1910, Part 5: Family Alpheidae. Smithsonian Contributions to Zoology, 466, 1 - 99. https: // doi. org / 10.5479 / si. 19436696.391.1"]}

Alpheus richardsoni Yaldwyn, 1971 (Figs. 17, 18, 46F–G, 51C) Alpheus richardsoni Yaldwyn 1971: 88; Fransen et al. 1997: 36; Poore 2004: 106, fig. 27c, d, 28l, 29s, t, pl. 9b; Gowlett-Holmes 2008: 215; De Grave & Fransen 2011: 389. Alpheus euphrosyne.— Hutchings & Recher 1974: 106 [not A. euphrosyne De Man, 1897]. Alpheus euphrosyne richardsoni.— Banner & Banner 1982: 235, fig. 74 (part.?); Morgan & Jones 1991: 485; Morrison et al. 2014: 88, fig. 37c. Alpheus sp. — Richardson & Yaldwyn 1958: 37, fig. 35. Not Alpheus euphrosyne richardsoni.— Miya 1995: 275, fig. 2; Yang & Kim 1996: 106, figs. 1–3; Hayashi 1998: 292, figs. 353c, g, 354c, 355c, e [= A. nomurai sp. nov.]; Binh & Van 2021: 132 [= Alpheus sp., possibly A. euphrosyne De Man, 1897 or A. eurydactylus De Man, 1920]. Not Alpheus richardsoni. — Kunishima et al. 2022: 134, fig. 5 [= Alpheus sp. from A. malabaricus complex, possibly A. dolichodactylus Ortmann, 1890]. Material examined. Australia: 1 male (cl 9.5 mm), OUMNH. ZC. 2009.09.16, Queensland, Moreton Bay, Gold Coast, Loders Creek, 27°57.232’S 153°24.617’E, under rocks on muddy sand at high tide mark, leg. A. Anker & S. De Grave, 11.03.2009; 1 male (cl 8.3 mm), FLMNH UF 18694, same collection data as for prevous specimen [AUS-009]; 1 ov. female (cl 8.3 mm), FLMNH UF 18693, same collection data as for previous specimen [AUS-008]; 1 male (cl 9.3 mm), OUMNH. ZC. 2009.09.15, Queensland, Cleveland, Raby Bay Marina, 27°30.245’S 153°24.341’E, muddy lower intertidal, leg. A. Anker & S. De Grave, 12.03.2009 [AUS-055]; 1 male (cl 13.8 mm), USNM 106164, Queensland, North Stradbroke Island, Dunwich, leg. J.M. Moulton, 03.09.1960; 1 male (cl 10.0 mm), OUMNH. ZC. 2003.20.27, New South Wales, Clarence River, Thorny Island, ungated drainage system, leg. “H.K”, 07.2000; 1 male (cl 11.7 mm), OUMNH. ZC. 2003.20.24, same collection data as for previous specimen; 1 ov. female (cl 15.5 mm), RMNH Crus.D. 49642, New South Wales, north of Sydney, Broken Bay, Pittwater, Bayview, 17.04.1955, mud beach at low tide, near low tide mark, leg. L.B. Holthuis; 1 female (cl 10.0 mm), NMV J41038, Victoria, Port Phillip Bay, central muddy region, 3759.77’S, 14457.43’E, depth: 17 m, Smith MacIntyre grab, leg. R. S. Wilson, G. Walker-Smith & S. Heislers, 03.04.1995; 2 males (cl 12.4, 13.0 mm), 1 ov. female (cl 13.8 mm), NMV J40239, Victoria, Mallacoota Inlet, 37°32.7’S, 149°45.5’E, under wharf, depth: 1 m, oyster bag, leg. M.F. Gommon, 06.04.1989; 1 male (cl 12.7 mm), 1 ov. female (cl 14.6 mm), NMV J43683, Victoria, Port Phillip Bay, Corio muddy region, 38°07.0’S, 144°23.1’E, depth: 9 m, Smith MacIntyre grab, leg. R. S. Wilson & S. Heislers, 17.10.1995; 2 ov. females (cl 8.7, 10.8 mm), NMV J37327, Victoria, reef north of San Remo, intertidal, under rocks, leg. G. Poore et al., 28.04.1994; 1 male, 2 ov. females (cl 11.5–13.1 mm), NMV J21603, Victoria, Chesapeake Island, French Island, Western Port, collector unknown, 21.01.1972; 2 males (cl 11.5, 12.4 mm), NMV J21643-1, Victoria, Crib Point, Western Port, leg. G. Westcott, 22.06.1973; 2 ov. females (cl 11.5, 15.2 mm), NMV J21601, Victoria, Toora Beach, leg. S.W. Gunn, 01.09.1983; 1 male (cl 15.5 mm), 1 ov. female (cl 17.5 mm), NMV J21606, Victoria, Miller’s landing, Western Port, leg. Marine Research Group, 17.10.1981; 1 male (cl 13.2 mm), 1 ov. female (cl 13.8 mm), NMV J21642, South Australia, Pine Point, 19.11.1983; 1 ov. female (cl 10.2 mm), NMV J22006-2, Investigator Strait, leg. J.E. Watson, 17.01.1971. Description. See Yaldwyn (1971) for original description and Banner & Banner (1982) for detailed redescription and illustrations (as A. euphrosyne richardsoni); a complementary, fully revised diagnosis is provided below. Diagnosis. Large-sized species of Alpheus (maximal cl 20.0 mm, tl 65.0 mm). Carapace and pleon smooth, without pubescence. Rostrum subtriangular, acute, reaching or slightly overreaching mid-length of first article of antennular peduncle; rostral carina present, blunt, gently sloping into moderately deep rostro-orbital furrows. Orbital hoods broadly rounded in dorsal view. Pterygostomial angle broadly rounded. Telson ovate-rectangular, gradually tapering towards posterior margin, about 1.3–1.4 times as long as maximal width; dorsal surface with shallow, longitudinal, median depression, latter without setae, and two pairs of spiniform setae inserted at some distance from margin; posterior margin broadly rounded; posterolateral angle with two small spiniform setae. Antennular peduncle with stylocerite convex laterally, with tip subacute, not overreaching distal margin of first article; ventromesial carina with subtriangular tooth bearing small acute point; second article about 2.0–2.2 times as long as wide. Antennal basicerite with small distoventral tooth; scaphocerite with lateral margin shallowly concave; blade not broadened, separated from strong distolateral tooth by deep cleft; anterior margin of blade rounded, not reaching beyond distolateral tooth. Third maxilliped with antepenultimate article not broadened; penultimate article 2.5 times as long as wide. Chelipeds with ventromesial margin of meri unarmed distally. Major chela of A. edwardsii - type, with fingers about 0.4–0.5 length of palm; mesial face of palm smooth, with narrow mesial longitudinal groove, without mesial transverse ridge; lateral face of palm largely smooth, except for small field of minute granules above ventral shoulder (sometimes absent), with subrectangular lateral longitudinal groove; dorsal shoulder pronounced, rounded, sloping with angle superior to 45° into dorsal transverse groove, not overhanging; ventral shoulder pronounced, slightly protruding in lateral view, broadly rounded; mesial face of pollex faintly granulated, with poorly developed mesial subdistal ridge, without proximal protuberance and groove; lateral surface of pollex, as well as mesial and lateral surfaces of fingers smooth; dactylus with dorsal ridge slightly curved mesially, not protruding; dactylar plunger stout, distinct from dactylar ventral margin; distal portion of plunger with obliquely flattened ventromesial area; adhesive discs small. Male minor chela stout, strongly balaeniceps, with fingers about 0.8 length of palm; all surfaces smooth, without granulation; mesial face of palm with shallow mesial longitudinal groove; lateral face of palm with well-marked lateral longitudinal groove; dorsal shoulder not distinct, at most present as slight sinus; ventral transverse groove constriction-like; ventral shoulder poorly developed, rounded; dactylus with proximal balaeniceps expansion about 1.5 times as long as maximal dactylar width, with rows of balaeniceps setae. Female minor chela not balaeniceps; palm smooth, without sculpture; fingers slightly longer than palm, simple, with sharp cutting edges, not gaping. Second pereiopod with ratio of carpal subarticles approximately equal to 3.8: 2.2: 1: 1: 1.5. Third pereiopod with ischium typically armed with spiniform seta; merus about 4.0 times as long as maximal width, unarmed; propodus with six to eight stout spiniform setae on ventral margin and one pair of spiniform setae on distoventral margin adjacent to dactylus; dactylus about 0.4 length of propodus, spatulate. Fifth pereiopod with ischium unarmed. Male second pleopod with appendix masculina ~1.2 times length of appendix interna. Uropod with both protopod lobes ending in sharp tooth distally. Eggs numerous (> 100 in large females), small; egg diameter: 0.7 mm. Colour pattern. Carapace brownish to olive-green with more or less conspicuous, whitish, transverse, crescentshaped band and some white spots or whitish mottling; each pleonite with two transverse, greenish or brown-green bands, sometimes with some additional bands on first two or three somites; antennular peduncles and scaphocerite brownish or pale brown-green; antennular and antennal flagella bluish or bluish green; mesial face of major chela olive-green to bluish green, with some greyish and paler beige areas; fingers pinkish distally; minor chela dark olive-grey or brown-green; walking legs pinkish or yellowish; telson greyish blue; uropods greenish or brownish with large bluish areas, blue colour being more intense posterior to transverse suture (Figs. 17, 18; see also colour photographs in Poore 2004: pl. 9b; Gowlett-Holmes 2008: 215). Type locality. Bay of Islands, North Island, New Zealand. Distribution. Southern temperate and subtropical Indo-West Pacific (Fig. 51C): Australia (Western Australia, South Australia, Victoria, Tasmania, New South Wales, southern Queensland) and New Zealand (North Island). Common name proposed. Richardson’s snapping shrimp. Ecology and biology. Alpheus richardsoni is an intertidal and shallow subtidal snapping shrimp, which is frequently encountered on mudflats and sandflats with abundant rocks and other types of shelters, in mangroves, as well as estuarine habitats, typically between the lower intertidal zone and about 5 m, sometimes down to 24 m. Being both an eurythermic and an euryhaline species, A. richardsoni is capable of tolerating marked seasonal variations, with surface water temperature fluctuating from 6°C to 28°C, and salinities ranging from 140 / 00 to 350 / 00, for instance, in Tasmania and New Zealand. It lives in complex burrow systems, often dug under large rocks, with several chambers and galleries descending 30 cm or more below the surface (Banner & Banner 1982; present study). As for A. euphrosyne and A. eurydactylus, the large number of eggs and their small size suggest an extended larval development in A. richardsoni. Banner & Banner (1982: 239, 240) provided a detailed summary of the biology of the Tasmanian populations of A. richardsoni using data from an unpublished PhD thesis (U [U Khin Khin] 1977). Alpheus richardsoni is one of the largest snapping shrimps in southern Australia, with adult individuals reaching cl 20 mm and tl 65 mm (Banner & Banner 1982). Noteworthy is that the size of A. richardsoni gradually diminishes along the south-north gradient on the eastern coast of Australia. For instance, in the present material, the largest specimen from southern Queensland, near the northern limit of the species’ geographic range, is a female from Gold Coast (OUMNH. ZC. 2009.09.16) at cl 9.5 mm and tl 26 mm. The largest specimen from New South Wales in the present material is a male from Clarence River (OUMNH. ZC. 2003.20.24) at cl 12.5 mm and tl 30 mm. In contrast, several specimens from Victoria are significantly larger in size, often exceeding cl 13 mm, with one ovigerous female from Western Port (NMV J21606) reaching cl 17.5 mm and tl 56 mm. Taxonomic remarks. Alpheus richardsoni is easily distinguishable from both A. euphrosyne and A. eurydactylus by the feebly developed distomesial ridge on the major chela pollex (vs. a very prominent one in A. euphrosyne and A. eurydactylus); the granulation of the mesial palmar face of the major chela reduced to a few granules above the ventral shoulder (vs. with a much more extensive granulation in A. euphrosyne and A. eurydactylus); the greatly reduced sculpture of the male major chela, for instance, the absence of a clear dorsal groove and shoulder (vs. with longitudinal grooves and a distinct dorsal shoulder in A. euphrosyne and A. eurydactylus); and the antennal basicerite armed with a small distolateral tooth (vs. usually unarmed in A. euphrosyne and A. eurydactylus) (Banner & Banner 1982: fig. 74; cf. Figs. 6, 7, 14, 15). Alpheus richardsoni differs specifically from A. euphrosyne by the non-overhanging dorsal shoulder of the major chela palm (vs. overhanging the transverse groove in A. euphrosyne); the presence of distinct rostro-orbital furrows (which are barely distinct or absent in A. euphrosyne); and the dorsal surface of the telson bearing only a very shallow, non-setose median longitudinal depression (vs. with a much deeper median depression, fringed laterally by numerous long setae, in A. euphrosyne) (Banner & Banner 1982: fig. 74; cf. Figs. 6, 7). The unique and diagnostic colour pattern of A. richardsoni (Figs. 17, 18) separates this species at once from A. euphrosyne (Fig. 8–10) and A. eurydactylus (Fig. 16), but also from all other species of Alpheus, including all species treated below. In addition, A. richardsoni is confined to subtropical and temperate waters of Australia and New Zealand and is thus geographically separated from A. euphrosyne (not known from Australia) and A. eurydactylus (known only from tropical northern Australia) (Fig. 51A–C). For separation of A. richardsoni from A. euphrosyne, A. eurydactylus and other species included in this study see below or refer to Table 1. The specimens from two localities near Cairns, Queensland (Arlington Reef and an intertidal reef flat at Green Island), which were tentatively identified as A. euphrosyne richardsoni by Banner & Banner (1982), do not appear to belong to A. richardsoni. As noted by Banner & Banner (1982), these specimens “were from neither silty nor brackish conditions, yet morphologically they could not be distinguished from A. e. richardsoni.... However, they were markedly smaller than A. e. richardsoni, for none, including 3 ovigerous females, were over 25 mm in length, while A. e. euphrosyne and A. e. richardsoni both reach 65 mm in length at maturity”. According to these authors, these specimens may represent “yet another subspecies of the nominate species that has adapted to living in other than mud and brackish water”. Since all other eastern Australian populations of A. richardsoni occur to the south of Moreton Bay, Queensland (Figs. 17, 51C), the material from Arlington Reef and Green Island is also somewhat isolated geographically. Whatever the case may be, the taxonomic status of these specimens can only be determined after collection and examination of fresh material from reefs off or near Cairns., Published as part of Anker, Arthur, 2023, Revision of Alpheus euphrosyne De Man, 1897 and A. microrhynchus De Man, 1897, with description of three new species and taxonomic remarks on several other morphologically and ecologically similar snapping shrimps (Malacostraca: Decapoda: Alpheidae), pp. 1-115 in Zootaxa 5282 (1) on pages 34-38, DOI: 10.11646/zootaxa.5282.1.1, http://zenodo.org/record/7912292, {"references":["Yaldwyn, J. C. (1971) Preliminary descriptions of a new genus and twelve new species of natant decapod Crustacea from New Zealand. Records of the Dominion Museum, 7, 85 - 94.","Fransen C. H. J. M., Holthuis, L. B. & Adema, J. P. H. M. (1997) Type-catalogue of the decapod Crustacea in the collections of the Nationaal Natuurhistorisch Museum, with appendices of pre- 1900 collectors and material. Zoologische Verhandelingen, Leiden, 311, 1 - 344.","Poore, G. (2004) Marine decapod Crustacea of southern Australia. A guide to identification. Museum Victoria and CSIRO, Collingwood, 576 pp., 32 pls. https: // doi. org / 10.1071 / 9780643092129","Gowlett-Holmes, K. (2008) A field guide to the marine invertebrates of South Australia. Notomares, Sandy Bay, 333 pp.","De Grave, S. & Fransen, C. H. J. M. (2011) Carideorum catalogus: the recent species of the dendrobranchiate, stenopodidean, procarididean and caridean shrimps (Crustacea: Decapoda). Zoologische Mededelingen, Leiden, 85, 195 - 589.","Hutchings, P. A. & Recher, H. F. (1974) The fauna of Careel Bay with comments on the ecology of mangrove and sea-grass communities. Australian Zoologist, 18, 99 - 128.","De Man, J. G. (1897) Bericht u ¨ ber die von Herrn Schiffscapitan Storm zu Atjeh, an den westlichen Ku ¨ sten von Malakka, Borneo und Celebes sowie in der Java-See gesammelten Decapoden und Stomatopoden. Fu ¨ nfter Theil. Z oologische Jahrbu ¨ cher. Abtheilung fu ¨ r Systematik, Geographie und Biologie der Thiere, 9, 725 - 790, pls. 34 - 37 (in volume 10).","Banner, D. M. & Banner, A. H. (1982) The alpheid shrimp of Australia. Part III: The remaining alpheids, principally the genus Alpheus and the family Ogyrididae. Records of the Australian Museum, 34, 1 - 357. https: // doi. org / 10.3853 / j. 0067 - 1975.34.1982.434","Morgan, G. J. & Jones, D. S. (1991) Checklist of marine decapod Crustacea of southern Western Australia. In: Wells, F. E., Walker, D. I., Kirkman, H. & Lethbridge, R. (Eds.), The marine flora and fauna of Albany, Western Australia. Vol. 2. Western Australian Museum, Perth, pp. 483 - 497.","Morrison, M. A., Jones, E. G., Consalvey, M. & Berkenbusch, K. (2014) Linking marine fisheries species to biogenic habitats in New Zealand: a review and synthesis of knowledge. New Zealand Aquatic Environment and Biodiversity Report, 130, 1 - 156.","Richardson, L. R. & Yaldwyn, J. C. (1958) A guide to the natant decapod Crustacea (shrimps and prawns) of New Zealand. Tuatara, 7, 17 - 41.","Miya, Y. (1995) Four species of Alpheus from intertidal and shallow water mudflats in the Sea of Ariake, Kyushu, Japan (Crustacea, Decapoda, Alpheidae). Bulletin of the Faculty of Liberal Arts, Nagasaki University, Natural Science, 35, 271 - 288.","Yang, B. P. & Kim, C. H. (1996) Zoeal stages of Alpheus euphrosyne richardsoni Yaldwyn, 1971 (Decapoda: Macrura: Alpheidae) reared in the laboratory. Korean Journal of Zoology, 39, 106 - 114.","Hayashi, K. I. (1998) Prawns, shrimps and lobsters from Japan (101). Family Alpheidae - Genus Alpheus (10). Aquabiology, 117 (20), 289 - 293.","Binh, N. T. & Van, N. C. (2021) Research on species composition and distribution of benthic fauna in the coastal area of Ninh Binh province. Science Journal of Tan Trao University, 22, 128 - 141.","De Man, J. G. (1920) Diagnoses of some new species of Penaeidae and Alpheidae with remarks on two known species of the genus Penaeopsis A. M. - Edw. from the Indian Archipelago. Zoologische Mededelingen, Leiden, 5, 103 - 109.","Kunishima, T., Palla, H. P., Tachikara, K. & Maeda, K. (2022) First records of an estuarine goby Acentrogobius ocyurus (Gobiiformes: Gobiidae) from Japan and the Sulu Sea in the Philippines. Species Diversity, 27, 129 - 138. https: // doi. org / 10.12782 / specdiv. 27.129","Ortmann, A. (1890) Die Decapoden-Krebse des Strassburger Museums, mit besonderer Beru ¨ cksichtigung der von Herrn Dr. D ˆ derlein bei Japan und bei den Liu-Kiu-Inseln gesammelten und z. Z. im Strassburger Museum aufbewahrten Formen. I. Theil. Die Unterordnung Natantia (Boas) (Abtheilungen: Penaeidae und Eucyphidea = Caridae der Autoren). Zoologische Jahrbu ¨ cher. Abtheilung fu ¨ r Systematik, Geographie und Biologie der Thiere, 5, 437 - 542, pls. 36, 37."]}

Alpheus cyanoteles Yeo & Ng, 1996 (Figs. 23, 24A–D, 51E) Alpheus cyanoteles Yeo & Ng 1996: 46, figs. 4–7; Hurt et al. 2021: 7. Alpheus paludosus (lap. cal.).— Johnson 1965: 9 (part.?); Johnson 1966: 430; Lovett 1981: 67 (not A. paludicola Kemp, 1915). Alpheus paludicola. — Banner & Banner 1966: 135 (part.?) (not A. paludicola Kemp, 1915). Type material. Holotype, male (cl 13.7 mm, tl 35.7 mm, chl 14.0 mm), ZRC 1996.1, Malaysia, Johor, Kota Tinggi, Sungei Tementang, 01°52’0.81 N, 103°55’49.2 E, freshwater stream, leg. P.K.L. Ng et al., 05.06.1995. Additional material. Malaysia: 1 male (cl 10.2 mm), 1 female (cl 15.6 mm, tl 42.5 mm, chl 22.0 mm), MNHN-IU-2018-5734, Johor, Sungei Tementang, leg. H.H. Tan et al., 23.10.2009; 1 male (cl 12.8 mm), OUMNH. ZC. 2013.05.5, Johor, Mawai, Sungei Tementang, Kota Tinggi —Kuantan road, km 268.5 Kuantan / km 60.5 Johor Bahru, 01°52.846’N, 103°55.799’E, 19 m above sea level, leg. H.H. Tan, S. De Grave & A. Anker, 29.05.2013; 1 male (cl 9.5 mm), 2 females (cl 9.5, 9.8 mm), OUMNH. ZC. 2013.05.6 –8, Johor, Mawai, Sungei Kayu, Kota Tinggi —Kuantan road, km 262 Kuantan / km 67 Johor Bahru, 0155.557’N, 10354.400’E, 26 m above sea level, leg. H.H. Tan, S. De Grave & A. Anker, 29.05.2013; 2 males (cl 7.8, 8.3 mm), 1 female (cl 8.2 mm), ZRC 2014.0686, Johor, Sedili, Sungei Kayu, swamp forest, 02.1937; 1 male (cl 9.5 mm), 1 female (cl 10.3 mm), ZRC 1996.46, Johor, no further data; 1 male (cl 7.5 mm), ZRC 1996.5, Johor, Kota Tinggi, Sungei Selangi, stream banks, leg. P.K.L. Ng et al., 1994; 1 male (cl 14.2 mm, tl 36.0 mm, chl 19.5 mm), ZRC 2000.2131, Sarawak, Serian, fish market, O. Chia, 23.06.1998;> 40 specimens of both sexes, (cl indet., no ov. females, small vial with detached eggs present), ZRC 2000.2185, Sarawak, Serian, fish market, H.H. Tan, 29– 31.10.1997; 2 males (cl 11.5, 12.5 mm), 1 female (cl 12.2 mm), MNHN-IU-2018-5591, same collection data as for previous specimens. Description. See Yeo & Ng (1996) for original description and illustrations; a complementary, revised diagnosis is provided below. Diagnosis. Medium-sized species of Alpheus (maximal cl 16.7 mm, tl ~43.0 mm). Carapace and pleon smooth, without pubescence. Rostrum short, subtriangular, wider than long, reaching 0.3 length of first article of antennular peduncle, sometimes obliquely ascendant; rostral carina low, short, gently sloping into very shallow rostro-orbital furrows, flattening between orbital hoods. Orbital hoods somewhat angular in dorsal view. Pterygostomial angle broadly rounded. Telson ovate-rectangular, gently tapering towards posterior margin, about 1.3 times as long as maximal width; dorsal surface with faint, longitudinal median depression, latter without setae, and two pairs of small spiniform setae inserted at some distance from margin; posterior margin broadly rounded; posterolateral angle with two very small spiniform setae. Antennular peduncle with stylocerite slightly convex laterally, with tip acute, not overreaching distal margin of first article; ventromesial carina with reduced tooth, latter without acute point; second article about 2.1 times as long as wide. Antennal basicerite armed with sharp distoventral tooth; scaphocerite with lateral margin nearly straight; blade broad, separated from strong distolateral tooth by deep cleft; anterior margin of blade rounded, reaching well beyond distolateral tooth. Third maxilliped with antepenultimate article not broadened; penultimate article about four times as long as wide. Chelipeds with ventromesial margin of meri unarmed distally. Major chela of A. edwardsii - type, with fingers about 0.6–0.7 length of palm; mesial face of palm smooth, with low mesial transverse ridge and very shallow and narrow mesial longitudinal groove; lateral face of palm with subrectangular lateral longitudinal groove; dorsal shoulder rounded, sloping gently with angle inferior to 45° into dorsal transverse groove, not overhanging; ventral shoulder pronounced, slightly protruding in lateral view, broadly rounded; mesial and lateral surfaces of both fingers smooth, without granules; mesial face of pollex with well-developed mesial subdistal ridge extending from low proximal protuberance and bordered by two shallow depressions; dactylus with dorsal ridge slightly curved mesially, not protruding; dactylar plunger large, stout, distinct from dactylar ventral margin; distal portion of plunger with obliquely flattened ventromesial area; adhesive discs small. Male minor chela moderately stout, strongly balaeniceps, with fingers about 0.9 length of palm; all surfaces smooth, without granulation; mesial face of palm with poorly developed mesial longitudinal groove; lateral face of palm with narrow lateral longitudinal groove; dorsal transverse groove and shoulder not distinct; ventral transverse groove and shoulder at most indicated by shallow sinus; dactylus with proximal balaeniceps expansion about 2.2 times as long as maximal dactylar width, with rows of balaeniceps setae. Female minor chela not balaeniceps; palm smooth, without sculpture; fingers subequal to palm, simple, with sharp cutting edges, not gaping. Second pereiopod with ratio of carpal subarticles approximately equal to 5: 2: 1: 1: 1.5. Third pereiopod with ischium unarmed; merus more than six times as long as maximal width, unarmed; propodus with about eight stout spiniform setae on ventral margin and one pair of spiniform setae on distoventral margin adjacent to dactylus; dactylus about 0.3 length of propodus, trigonal-conical, tapering. Fifth pereiopod with ischium unarmed. Male second pleopod with appendix masculina about same length as appendix interna or up to 1.3 times longer. Uropod with both protopod lobes ending in sharp tooth distally. Eggs not numerous (usually less than 30), very large; egg diameter about 1.5 mm. Colour pattern. Carapace pale brownish orange, with darker grey-brown saddle and some orange mottling, posterior margin with narrow, transverse, dark blue band; pleon with whitish or pale yellow ground colour; each pleonite with one broad, transverse, dark greyish blue band on posterior half; middle section of each pleonite occupied by diffuse, transverse, pale orange band; antennular peduncles and scaphocerite yellowish, marbled with orange brown; antennular and antennal flagella pale bluish green; mesial face of major chela orange-brown or olive-brown, with paler orange or yellowish areas, especially on dorsal and ventral transverse grooves; dorsal margin of pollex and distomesial ridge area conspicuous blue; dactylus orange-brown with olive tinge and with bright blue markings on margins; distal portion of both fingers pink; minor chela brown- or olive-orange, fingers darker; second to fifth pereiopods reddish; telson pale yellow-orange proximally, turning bluish distally; uropodal exopod whitishyellow proximally, conspicuously deep blue posterior to transverse suture; uropodal endopod pale orange with some blue distally (Figs. 23, 24A–D). Type locality. Sungei Tementang, Kota Tinggi, Johor, Malaysia. Distribution. Restricted to Sunda area (Fig. 51E): southern Johor in peninsular Malaysia (Sedili basin, e.g., Sungei Tementang, Sungei Kayu); Sarawak, Malaysian part of Borneo (vicinity of Serian); southern Thailand (Pattalung Province: Tapi River, see remarks below). Common name proposed. Freshwater snapping shrimp. Ecology and biology. Alpheus cyanoteles is a true freshwater snapping shrimp living in small rivers and streams above tidal influence (pH 4.7–5.8), shaded by healthy or degraded rainforests (Fig. 23E). The syntopic fauna consists of primary and secondary freshwater fishes (e.g., Cyprinidae, Balitoridae, Cobitidae, Siluridae, Phallostethidae, Synbranchidae, Nandidae, Channidae) as well as palaemonid (Macrobrachium spp.) and atyid (Caridina spp.) shrimps, and freshwater crabs (Parathelphusa sp.) (see Yeo & Ng 1996 for extensive description of the type locality). The low number of eggs and their large size suggest that the larval development of A. cyanoteles is highly abbreviated. Abbreviated, semi-direct and direct larval development is part of a series of physiological and reproductive adaptations of various decapod crustaceans to life in freshwater environments. However, the material from Serian (Sarawak) may have been collected in slightly brackish conditions (see comments below). Alpheus cyanoteles becomes increasingly popular in aquarium trade as the world’s only true freshwater snapping shrimp and is occasionally sold in pet shops in Malaysia and Singapore. In Borneo, A. cyanoteles is apparently also sold as food on some traditional markets (see below). Taxonomic remarks. Alpheus cyanoteles appears to be most closely related to A. microrhynchus. These two species share a great number of morphological features and have abbreviated larval development.The only consistent morphological difference between them is the presence of a moderately large, sharp tooth on the distolateral margin of the antennal basicerite in A. cyanoteles (Yeo & Ng 1996: fig. 5b); this tooth is either absent or minute in A. microrhynchus (De Man 1897). In life, however, A. cyanoteles and A. microrhynchus can be easily distinguished from each other by their colour patterns, especially by the pleon decorated with broad, dark blue bands, alternating with diffuse orange bands, in A. cyanoteles (Figs. 23, 24) vs. with less conspicuous, narrower dusky grey-blue bands, without intercalated orange bands, in A. microrhynchus (Figs. 21, 22). Yeo & Ng (1996: fig. 5e) did not illustrate or mention the rather conspicuous, sharp, mesial subdistal ridge on the pollex of the major chela in A. cyanoteles (well visible in Figs. 23B, 24A, D). This ridge is also present in A. microrhynchus (Figs. 19D, 21E), A. euphrosyne (abbreviated as “msr in Fig. 4; see also Figs. 5C, 9B), A. eurydactylus (Figs. 11C, D, 16C), and several other species of this group, including herein described new taxa (see below and refer to Table 1). Alpheus cyanoteles can be easily separated from A. euphrosyne, A. eurydactylus and A. richardsoni by the non-expanded, more trigonal-conical dactyli on the third to fifth pereiopods; and the absence of a spiniform seta on the ischium of the third and fourth pereiopods (although it may be obsolete or absent in large specimens of these three species); from A. euphrosyne and A. eurydactylus by the mesially non-granulated major chela; the much less sculptured male minor chela; and the presence of a distolateral tooth on the antennal basicerite (absent in A. eurydactylus, absent or minute in A. euphrosyne); from A. eurydactylus and A. richardsoni and by the shorter rostrum; from A. euphrosyne by the non-overhanging dorsal shoulder and the much stouter dactylar plunger of the major chela; and from A. richardsoni by the much stronger distomesial ridge on the major chela and the noticeably slenderer male minor chela (Yeo & Ng 1996: figs. 5–7; cf. Figs. 6, 7, 14, 15 in the present study; Banner & Banner 1982: fig. 74). In life, A. cyanoteles is easily distinguishable from A. euphrosyne, A. eurydactylus and A. richardsoni by several marked differences in the colour pattern (cf. Figs. 8, 9, 16–18, 23, 24). For separation of A. cyanoteles from other species see below or refer to Table 1. The record of A. cyanoteles from Tapi River in Thailand is based on colour photographs supplied by Jens Ķhne in 2013 (A. Anker, pers. obs.). This interesting photographic record extends the previously known range of A. cyanoteles from Johor to southern Thailand and suggests that the species may also occurs in other river systems along the eastern coast of Malay Peninsula. The material from the Serian fish market in Sarawak (Malaysian Borneo) was identified as A. cyanoteles based on the combination of morphological characters, including the presence of a stout, sharp tooth on the antennal basicerite, which seems to be the only morphological feature reliably separating A. cyanoteles from the closely related A. microrhynchus, its presumed oligohaline sister species. Serian is situated approximately 45 km south of Kuching and most likely these specimens were collected in one of the tributaries of Sungai Sabang or Batang Sadong. Taking into account the marine / brackish origin of both A. cyanoteles and A. microrhynchus and the presumably relatively recent invasion of freshwater habitats, it is possible that A. cyanoteles is able to tolerate slightly brackish conditions. However, this assumption needs to be confirmed by collection of fresh material from Serian and other localities in Borneo, with colour photographs and more reliable ecological data (salinity, pH). A molecular comparison between the Johor and Sarawak material of A. cyanoteles is highly desirable as these populations may well be genetically isolated, possibly as a result of independent freshwater invasion events. This hypothesis is supported by the possibly semi-direct or at least highly abbreviated larval development in A. cyanoteles, which presumably has some limiting effects on the circulation of larvae in both fresh and brackish waters. Hurt et al. (2021) recovered A. cyanoteles very distant from the remaining members of the A. edwardsii group. This rather unexpected result can be explained by the generally unsatisfactory sampling of the large and heterogeneous A. edwardsii group, which was confirmed to be non-monophyletic. The fact that several important taxa, such as A. euphrosyne, A. microrhynchus, A. richardsoni and other species of the A. edwardsii group associated with estuaries, mangroves, mudflats and brackish lagoons, were not included in the molecular analysis of Hurt et al. (2021), certainly created strong biases for other groups, and may have resulted in a long branch attraction artifact for A. cyanoteles., Published as part of Anker, Arthur, 2023, Revision of Alpheus euphrosyne De Man, 1897 and A. microrhynchus De Man, 1897, with description of three new species and taxonomic remarks on several other morphologically and ecologically similar snapping shrimps (Malacostraca: Decapoda: Alpheidae), pp. 1-115 in Zootaxa 5282 (1) on pages 46-50, DOI: 10.11646/zootaxa.5282.1.1, http://zenodo.org/record/7912292, {"references":["Yeo, D. C. J. & Ng, P. K. L. (1996) A new species of freshwater snapping shrimp, Alpheus cyanoteles (Decapoda: Caridea: Alpheidae) from Peninsular Malaysia and a redescription of Alpheus paludicola Kemp, 1915. Raffles Bulletin of Zoology, 44, 37 - 63.","Hurt, C., Hultgren, K. M., Anker, A., Lemmon, A. R., Moriarty Lemmon, E. & Bracken-Grissom, H. (2021) First worldwide molecular phylogeny of the morphologically and ecologically hyperdiversified snapping shrimp genus Alpheus (Malacostraca: Decapoda). Molecular Phylogenetics and Evolution, 158 (197080), 1 - 11. https: // doi. org / 10.1016 / j. ympev. 2021.107080","Johnson, D. S. (1965) A review of brackish water prawns of Malaya. Bulletin of the National Museum of Singapore, 33, 7 - 11.","Lovett, D. L. (1981) A guide to the shrimps, prawns, lobsters and crabs of Malaysia and Singapore. Faculty of Fisheries & Marine Science, Universiti Pertanian Malaysia, Serdang, Selangor Malaysia, Occasional Publication, 2, 1 - 156.","Kemp, S. (1915) Fauna of the Chilka Lake. No. 3. Crustacea Decapoda. Memoirs of the Indian Museum, 5, 201 - 325, pls. 12 - 13. https: // doi. org / 10.5962 / bhl. title. 10414","Banner, A. H. & Banner, D. M. (1966) The alpheid shrimp of Thailand. The Siam Society Monograph Series, 3, 1 - 168.","De Man, J. G. (1897) Bericht u ¨ ber die von Herrn Schiffscapitan Storm zu Atjeh, an den westlichen Ku ¨ sten von Malakka, Borneo und Celebes sowie in der Java-See gesammelten Decapoden und Stomatopoden. Fu ¨ nfter Theil. Z oologische Jahrbu ¨ cher. Abtheilung fu ¨ r Systematik, Geographie und Biologie der Thiere, 9, 725 - 790, pls. 34 - 37 (in volume 10).","Banner, D. M. & Banner, A. H. (1982) The alpheid shrimp of Australia. Part III: The remaining alpheids, principally the genus Alpheus and the family Ogyrididae. Records of the Australian Museum, 34, 1 - 357. https: // doi. org / 10.3853 / j. 0067 - 1975.34.1982.434"]}

Alpheus firmus Kim & Abele, 1988 (Figs. 49, 50A, B, 52F) Alpheus firmus Kim & Abele 1988: 93, fig. 39 (part.?); Abele & Kim 1989: 15; Wicksten & Hendrickx 1992: 4; Wicksten & Hendrickx 2003: 64 (part.?). (?) Alpheus firmus.— Alvarez et al. 1996: 716, fig. 2 [= A. cf. firmus Kim & Abele, 1988; see below]. Material examined. Panama: 1 male (cl 6.9 mm), MNHN-IU-2018-5574, Veraguas, Coiba Archipelago, Coiba, Bahía Santa Cruz, intertidal mudflat with rocks, leg. A. Anker et al., 21.03.2007 [AA-07-145]; 1 male (cl 11.0 mm), 1 ov. female (cl 12.8 mm), USNM 237798, Panama Canal, Pacific side entrance, Miraflores Locks, NMNH-STRI sta. 203, 26.08.1974; 1 male, 1 female (cl indet., specimens in poor condition, not deposited, see below), Panama Canal, Pacific side entrance, Miraflores Locks, leg. J. Jara et al., 21.07.2005 [AA-05-189]. Tentative identification. Alpheus cf. firmus. Panama: 1 male (cl 10.0 mm), MNHN-IU-2018-5577, Veraguas, Puerto Vidal, mangrove creek, in mud under rocks at low tide, leg. A. Anker, 19.03.2007 [AA-07-092B]; 1 ov. female (cl 10.5 mm), MNHN-IU-2018-5576, same collection data as for previous specimen [AA-07-092A]; 1 male (cl 9.3 mm), MNHN-IU-2018-5575, same collection data as for previous specimens [AA-07-091]. Description. See Kim & Abele (1988) for detailed description and illustrations. Colour pattern. Type A colour morph (Alpheus firmus sensu Kim & Abele 1988): body uniform purplish to pinkish brown; posterior edges of pleonites and rostrum darker reddish-brown; antennular and antennal flagella greenish yellow; major and minor chelae greenish to olive-brown with paler green-yellow areas, fingers distally pinkish; walking legs reddish; uropod with bluish tinge distally (Figs. 49, 50A, B). Type B colour morph (Alpheus cf. firmus): greenish to olive-brown with whitish saddle on carapace and transverse whitish areas on pleon resulting in banded pattern; antennular and antennal flagella greenish yellow; major and minor chelae green-brown with paler greenish areas, fingers distally reddish; walking legs reddish; uropod with deep blue and brick-red areas distally (Fig. 50D–F). See discussion of the two colour morphs below. Type locality. Miraflores Locks, Pacific entrance to Panama Canal, Panama. Distribution. Tropical eastern Pacific: Panama (Punta Paitilla, Panama Canal, Puerto Vidal, Coiba Island) to Mexico (Nayarit) (Fig. 52F) (Kim & Abele 1988; Alvarez et al. 1996; present study); however, A. firmus sensu Kim & Abele (1988) may be restricted to Panama (see discussion below). Common name proposed. Panama Canal snapping shrimp. Ecology and biology. Alpheus firmus is confined to mudflats and mangroves with abundant wood debris and rocks, living in self-made burrows, sometimes dug under large rocks or decaying logs, mainly in the intertidal and adjacent shallow subtidal areas (a few meters in depth). It appears to be a euryhaline species, capable of tolerating brackish conditions. The large number of eggs and their small size suggest an extended larval development. Taxonomic remarks. The type series of A. firmus (USNM) is composed of 82 adult specimens collected at Miraflores Locks (Panama Canal) and Punta Paitilla, Panama City. In July 2005, a male-female pair of A. firmus (Fig. 49) was collected at the type locality of the species and kept in the running sea water tanks of the Naos Laboratories of the Smithsonian Tropical Research Institite (STRI). Unfortunately, due to an accidental rupture of sea water supply, both shrimps died and were found partly disintegrated; therefore, they were preserved in ethanol, but not deposited in a museum collection due to their poor condition. Another specimen, a smaller male, was collected in 2007 on a mudflat of Santa Cruz Bay, Coiba Island (Fig. 50A–C). These three specimens correspond well to the description and illustrations of the holotype of A. firmus in Kim & Abele (1988). Kim & Abele (1988) noted some variation in the presence / absence of a small, sharp tooth on the antennal basicerite, as well as in the relative length ratio of the penultimate and ultimate articles of the third maxilliped in the Panamanian material of A. firmus. The variation in these two features seems to be correlated: all specimens with a tooth on the basicerite also had a relatively shorter penultimate article of the third maxilliped (ratio ultimate vs. penultimate article 1.3–1.6), compared to the specimens with an unarmed basicerite (ratio ultimate vs. penultimate article 1.2). For instance, in the illustrated holotype of A. firmus, the basicerite has a small tooth and the ratio between the ultimate and penultimate articles is approximately 1.3 (Kim & Abele 1988: fig. 39d). In the specimen of A. firmus from Coiba, Panama, the basicerite is armed with a small tooth and the ratio between the ultimate and penultimate articles is about 1.5, whereas in the specimens tentatively identified as A. cf. firmus from Puero Vidal, Panama, the basicerite is unarmed and the ratio between the ultimate and penultimate articles is at most 1.2, thus confirming observations by Kim & Abele (1988). Alvarez et al. (1996), reporting A. firmus from Nayarit, Mexico, noted that their specimens also were lacking a tooth on the antennal basicerite and had a long ultimate article of the third maxilliped (twice as long as the penultimate instead of slightly longer, as in the type material). However, in the figure of the third maxilliped provided by these authors, the ultimate article is only ~1.6 times longer than the penultimate article (Alvarez et al. 1996: fig. 2b), although both articles are noticeably broader than their counterparts in the holotype (Kim & Abele 1988: fig. 39d). In addition, the colour pattern of the specimens from Miraflores Locks and Coiba Island identified as A. firmus (type A, with a non-banded pleon, Figs. 49, 50A, B), is somewhat different from that of the specimens assigned to A. cf. firmus (type B, with a banded pleon, Fig. 50D–F). The variation in both morphology (presence vs. absence of a tooth on the antennal basicerite, ratio and proportions of the penultimate and ultimate articles of the third maxilliped) and colour pattern (non-banded vs. banded pleon) in the Panamanian material and the presence of only morphotype in the Mexican material of A. firmus (Alvarez et al. 1996), suggests that a second species may be present in the tropical eastern Pacific, ranging from Mexico (Alvarez et al. 1996) to Panama (Kim & Abele 1988; present study), where it overlaps with the distributional range of A. firmus. This taxonomic problem will be addressed by the author in the near future, after a careful examination of the entire type series of A. firmus in the USNM (possible only after the end of the Covid-19 pandemic), DNA analyses of the Panamanian material collected in 2007, and a thorough morphological and genetic comparison of A. cf. firmus with A. zarenkovi Anker & Pachelle, 2015, described from the same geographical area (Anker & Pachelle 2015). For the time being, it seems more prudent to treat A. cf. firmus separately from A. firmus and to restrict the distributional range of the latter species to Panama (as shown in Fig. 52F). Alpheus firmus appears to be most closely related to the amphi-Atlantic Atlantic A. pontederiae and several members of the transisthmian A. heterochaelis species complex (possibly non-monophyletic grouping), which includes the western Atlantic A. heterochaelis Say, 1818, A. buckupi Almeida, Terossi, Araújo-Silva & Mantelatto, 2013 and A. petronioi Almeida, Terossi & Mantelatto, 2014, and the eastern Pacific A. distinctus Kim & Abele, 1988, all characterised by the presence of a well-developed balaeniceps condition on the male minor chela. However, A. firmus (sensu Kim & Abele 1988) can be separated from all these species by the longer and relatively slenderer penultimate article of the third maxilliped (Kim & Abele 1988: fig. 39d); specifically from A. pontederiae by the absence of a distinct mesial subdistal ridge on the major chela pollex (idem: fig. 39f; cf. Fig. 47D, 48C; Christoffersen 1984: fig. 4a) and the third and fourth pereiopods with a trigonal, feebly broadened dactylus (idem: fig. 39k; cf. Christoffersen 1984: fig. 4h); from A. heterochaelis by the distinctly shorter rostrum and the more gently sloping dorsal shoulder of the major chela (idem: fig. 39b, f; cf. Williams 1984: fig. 65a, b); from A. buckupi by the much shorter rostrum, the less marked rostral carina, the less concave lateral margin of the antennal scaphocerite, and the distomesially unarmed cheliped merus (idem: fig. 39b, f; cf. Almeida et al. 2013: figs. 1A, 2A); from A. petronioi by the shorter rostrum and less marked rostral carina, and the longer second article of the antennular pedunce (idem: fig. 39b; cf. Almeida et al. 2014: fig. 1B); and from A. distinctus by the more gently sloping dorsal shoulder of the major chela, the much less sculptured male minor chela, and the different ratio of the first and second carpal subarticle of the second pereiopod (idem: fig. 39f, h, j, 40f, h, i). Alpheus firmus also has affinities to some Indo-West Pacific species of the A. euphrosyne — A. microrhynchus complex, but can be easily separated from all of them by the elongated penultimate article of the third maxilliped, in combination with other morphological features (e.g., trigonal dactylus of the walking legs, major chela pollex without mesial subdistal ridge, weak sculpture of the minor chela, several features of the rostro-orbital region, antennules, antennae, etc.) and colour pattern., Published as part of Anker, Arthur, 2023, Revision of Alpheus euphrosyne De Man, 1897 and A. microrhynchus De Man, 1897, with description of three new species and taxonomic remarks on several other morphologically and ecologically similar snapping shrimps (Malacostraca: Decapoda: Alpheidae), pp. 1-115 in Zootaxa 5282 (1) on pages 90-93, DOI: 10.11646/zootaxa.5282.1.1, http://zenodo.org/record/7912292, {"references":["Kim, W. & Abele, L. G. (1988) The snapping shrimp genus Alpheus from the eastern Pacific (Decapoda: Caridea: Alpheidae). Smithsonian Contributions to Zoology, 454, 1 - 119. https: // doi. org / 10.5479 / si. 00810282.454","Abele, L. G. & Kim, W. (1989) The decapod crustaceans of the Panama Canal. Smithsonian Contributions to Zoology, 482, 1 - 50. https: // doi. org / 10.5479 / si. 00810282.482","Wicksten, M. K. & Hendrickx, M. E. (1992) Checklist of penaeoid and caridean shrimps (Decapoda: Penaeoidea, Caridea) from the eastern tropical Pacific. Proceedings of San Diego Society of Natural History, 9, 1 - 11.","Wicksten, M. K. & Hendrickx, M. E. (2003) An updated checklist of benthic marine and brackish water shrimps (Decapoda: Penaeoidea, Stenopodidea, Caridea) from the eastern tropical Pacific. In: Hendrickx, M. E. (Ed.), Contribuciones al estudio de los crustaceos del Pacifico Este. Instituto de Ciencias del Mar y Limnologia, Universidad Nacional Autonoma de Mexico, Mexico City, pp. 49 - 76.","Alvarez, F., Camacho, M. E. & Villalobos, J. L. (1996) The first species of Prionalpheus from the eastern Pacific, and new records of caridean shrimp (Crustacea: Decapoda: Caridea) from the western coast of Mexico. Proceedings of the Biological Society of Washington, 109, 715 - 724.","Anker, A. & Pachelle, P. P. G. (2015) Two new snapping shrimps (Decapoda, Caridea, Alpheidae, Alpheus) from the tropical eastern Pacific. Arthropoda Selecta, 24, 247 - 258.","Say, T. (1818) An account of the Crustacea of the United States, part 5. Journal of the Academy of Natural Sciences at Philadelphia, 1, 235 - 253.","Almeida A. O. de, Terossi M., Araujo-Silva C. L. & Mantelatto, F. L. (2013) Description of Alpheus buckupi spec. nov., a new amphi-Atlantic snapping shrimp (Caridea: Alpheidae), based on morphological and molecular data. Zootaxa, 3652 (4), 437 - 452. https: // doi. org / 10.11646 / zootaxa. 3652.4.3","Almeida, A. O. de, Terossi, M. & Mantelatto, F. L. (2014) Morphology and DNA analyses reveal a new cryptic snapping shrimp of the Alpheus heterochaelis Say, 1818 (Decapoda: Alpheidae) species complex from the western Atlantic. Zoosystema, 36, 53 - 71. https: // doi. org / 10.5252 / z 2014 n 1 a 4","Christoffersen, M. L. (1984) The western Atlantic snapping shrimps related to Alpheus heterochaelis Say (Crustacea, Caridea) with the description of a new species. Papeis Avulsos de Zoologia, 35, 189 - 208.","Williams, A. B. (1984) Shrimps, lobsters, and crabs of the Atlantic coast of the eastern United States, Maine to Florida. Smithsonian Institution Press, Washington, D. C., 550 pp."]}

Alpheus dingabadi sp. nov. (Figs. 1–3) Type material. Holotype: male (cl 5.4 mm), FLMNH UF 39660, Maldives, Faafu Atoll, 1.6 km east of Magoodhoo Island, Dhign Reef, 3.0803 72.9827 [3°04’49.1” N 72°58’57.7”E], high impact forereef, 10–30 m, leg. J. Moore, 13 May 2014 [MALD-282]. Description. Carapace glabrous, with short, scattered setae (Fig. 1A, B). Rostrum short, narrow, about twice as long as wide at base, slightly descendent in lateral view, acute distally, barely reaching mid-length of first article of antennular peduncle, posteriorly continued by well-developed, narrow, blunt rostral carina, latter abruptly flattening between eyes (Fig. 1A, B). Orbital hoods swollen, somewhat projecting anteriorly in lateral view, with small, acute teeth, latter not reaching beyond tip of rostrum, directed anteriorly or slightly anteromesially; frontal margin between rostrum and orbital hood slightly flattened, almost straight (Fig. 1A, B). Pterygostomial angle broadly rounded (Fig. 1B); cardiac notch deep. Telson very broad, strongly tapering distally, about 1.7 times as long as maximal width; lateral margins slightly convex, more noticeably converging in posterior half; dorsal surface with two pairs of very stout spiniform setae inserted far from lateral margin, first pair at 0.4 of telson length, second pair between 0.6 and 0.7 of telson length; posterior margin broadly rounded; posterolateral angles each with one pair of spiniform setae, mesial ones between double and triple length of lateral ones (Fig. 1C). Eyes completely concealed in lateral view, partly concealed in frontal view; cornea large, well-pigmented (Figs. 1A, B, 3). Antennular peduncle moderately elongate; stylocerite narrow, feebly swollen, acuminate distally, tip not reaching distal margin of first article; ventromesial carina with strong, anteriorly directed tooth; second article about 2.5 times as long as wide; lateral antennular flagellum with secondary ramus fused to main ramus over most of its length, free portion represented by small stump; numerous groups of aesthetascs present from about 19th subdivision of flagellum to distal end of free portion (Fig. 1A, B, D–F). Antenna with basicerite very stout, much higher than long, armed with prominent, sharp tooth on distoventral margin; scaphocerite relatively small, with reduced blade and broadly concave lateral margin; distolateral tooth very strong, slightly overreaching distal margin of second article of antennular peduncle and distinctly reaching beyond mid-length of carpocerite; carpocerite very stout, subcylindrical, tapering distally, reaching or slightly overreaching distal end of antennular peduncle; flagellum moderately stout, furnished with long setae (Fig. 1A, B, D). Mouthparts typical for genus.Third maxilliped moderately slender, elongate, setose; coxa with bluntly projecting lateral plate; antepenultimate article flattened ventrolaterally, slightly twisted, with blunt ridge running parallel to dorsal margin on lateral surface, about four times as long as maximal width, with somewhat rugose ventral margin; penultimate article distally widening, about 2.8 times as long as distal width, with long setae on ventral margin; ultimate article 1.7 times as long as penultimate article, distally tapering, densely furnished with transverse rows of serrulate setae on dorsomesial and mesial surfaces and variously long, slender setae; exopod well developed, overreaching distal margin of antepenultimate article; arthrobranch well developed (Fig. 1G). Major cheliped robust; ischium short, stout, with two movable spinules on ventromesial margin; merus stout, trigonal in cross-section, slightly more than twice as long as maximal width of lateral surface; distodorsal margin terminating in blunt tooth; ventromesial margin straight, terminating in blunt distoventral lobe, armed with row of movable spinules; carpus very short, wider than long, cup-shaped; chela of basic macrocheles - type, not particularly elongate, rather stout, swollen, with numerous setae along dorsal and ventral margins and denser setal cover on most of mesial surface, especially on distal portion of palm and fingers; palm somewhat compressed, distally narrowing, slightly less than twice as long as maximal height (in lateral view of chela); distal area of lateral surface with strong longitudinal crest between two deep depressions, terminating in large acute tooth; dorsomesial margin with deep transverse groove extending somewhat to mesial surface, ending in large, blunt crest overhanging base of dactylus in mesial view; dorsomesial surface posterior to transverse groove and mesial to shallow dorsal groove crenulate; fingers, especially dactylus, noticeably deviating from main chela axis; pollex short, ending bluntly; dactylus exceeding pollex, distally strongly curved, rounded; plunger short, stout, distally truncate, well demarcated from anterior cutting edge of dactylus; adhesive disks very large and conspicuous (Fig. 2A–F). Minor cheliped relatively robust; ischium short, stout, with several movable spinules on ventromesial margin; merus slenderer than that of major cheliped, trigonal in cross-section, about 2.3 times as long as maximal width of lateral surface; distodorsal margin terminating in blunt tooth; ventromesial margin straight, terminating in blunt distoventral tooth, armed with row of movable spinules; carpus short, cup-shaped; chela swollen, stout, with numerous setae along dorsal and ventral margins and denser setal cover on mesial surface of palm and fingers; palm somewhat compressed, about 1.6 times as long as maximal height (in lateral view of chela); lateral surface without sculpture, terminating distally in small blunt tooth; dorsomesial margin with shallow longitudinal sulcus distally, delimiting very large, triangular, distally subacute, anteriorly protruding crest, latter extending well beyond base of dactylus; dorsomesial surface posterior to this crest somewhat crenulate; ventral surface between palm and fingers with broad, deep sinus; fingers slightly deviating from main chela axis, subequal in length, with blade-like cutting edges and fingertips curved and crossing distally; dactylus not expanded dorsally, not lamellar, without ridges, with strongly curved tip; adhesive disks small (Fig. 2G–I). Second pereiopod moderately slender; ischium and merus subequal in length; carpus with five subarticles, first longest, ratio of carpal subarticles approximately equal to 4.5: 2: 1.3: 1: 2; chela slender, about as long as first carpal subarticle (Fig. 1H). Third pereiopod robust; ischium with large spiniform seta on ventrolateral surface; merus about 3.7 times as long as maximal width, with prominent, acute, distoventral tooth; carpus about 0.6 times length of merus, much slenderer than merus, with small, subacute, distoventral tooth; propodus noticeably slenderer than carpus, about 1.7 times length of carpus, with row of about 10 small spiniform setae on ventral margin and one pair of spiniform setae on distoventral margin adjacent to base of dactylus; dactylus 0.35 times length of propodus, rather stout, strongly curved, distinctly biunguiculate, with secondary unguis shorter and slightly divergent from axis of main unguis (Fig. 1I, J). Fourth pereiopod generally similar to third pereiopod, slenderer; ischium more elongate, with large spiniform seta on ventrolateral surface; carpus slenderer than that of third pereiopod, otherwise similar; merus about 4.1 times as long as maximal width, with prominent, subacute, distoventral tooth, latter distinctly smaller than tooth of merus of third pereiopod; propodus with eight small spiniform setae on ventral margin and one pair of spiniform setae on distoventral margin adjacent to base of dactylus; dactylus similar to that of third pereiopod (Fig. 1K). Fifth pereiopod not noticeably slenderer than fourth pereiopod; ischium elongate, unarmed; merus about 4.2 times as long as wide; carpus slenderer than merus, about same length as merus, unarmed; propodus subequal to carpus in length, with propodal grooming brush composed of four rows of serrulate setae on distal fourth of ventrolateral surface, ventral margin with two spiniform setae both situated in distal fourth of article, distoventral margin with pair of slender spiniform setae adjacent to base of dactylus; dactylus similar to that of third and fourth pereiopods (Fig. 1L, M). Second pleopod with appendix masculina almost double length of appendix interna, reaching beyond distal margin of endopod, densely furnished with slender spiniform setae (Fig. 1N, O). Uropod with mesial and lateral lobes of protopod each distally produced as blunt or acute tooth, respectively; exopod with straight lateral margin ending in blunt distolateral tooth; adjacent spiniform seta very stout, reaching far beyond distal margin of exopod; distal margin armed with row of small spiniform setae; diaeresis sinuous, with subtriangular tooth near distolateral spiniform seta; endopod slightly narrower than exopod, oval-shaped; distal margin with row of spiniform setae increasing in size towards distolateral margin (Fig. 1P). Colour pattern. Body background whitish, almost opaque; carapace and pleon covered with bright red chromatophores disposed as following: carapace with large semicircular area of red chromatophores on gastric and postrostral regions, extending ventrally as oblique band on each side, and U-shaped posterior band consisting of two obliquely running bands, one on each side, joining each other along posterior margin; rostral area red; pleon with narrow transverse bands of red chromatophores, one per pleonite, joining each other on lateral surface of pleura; eyes with bluish-grey cornea; antennular peduncles mostly red; antennal peduncle mostly whitish to pale orange, with some red dots on basicerite and distal part of carpocerite; antennular and antennal flagella pale orange; third maxillipeds mainly red distally; major and minor chelipeds with merus mostly white with few scattered chromatophores, carpus pale orange-red, and mesial surface of chela decorated with large, irregularly shaped, redorange blotches and deep-red spots between whitish patches; second to fifth pereiopods largely colourless, except for reddish tinge on second pereiopod carpus; pleopods also colourless; telson and uropods with some reddish chromatophores (exact pattern not clearly seen in the only photograph available) (Fig. 3). Etymology. The specific epithet is a combination of two Maldivian (Dhivehi) words, din’gaa (shrimp) and badi (gun) (Abdulla & O’Shea 2005), with one letter omitted for euphony, and obviously referring to the fact that A. dingabadi sp. nov. is a pistol shrimp. The name is used as a noun in apposition. Common name proposed. Dingabadi snapping shrimp. Distribution. Central Indian Ocean: presently known only from the type locality in the Maldives. Ecology. The holotype was collected on an exposed forereef, between 10 and 30 m, most likely by breaking up coral rubble. Remarks. The combination of seven diagnostic characters of Alpheus dingabadi sp. nov., as following: (1) frontal margin of carapace with distinctly carinate rostrum and small, acute orbital teeth (Fig. 1A); (2) antennal scaphocerite short, with reduced blade and distolateral tooth not reaching distal fourth of carpocerite, reaching only slightly beyond distal margin of second article of antennular peduncle (Fig. 1A, B, D); (3) major chela swollen, not elongate, with two deep grooves and distally sharp crest between them on lateral surface, deep transverse groove proximally delimiting large blunt crest on dorsomesial surface, and area posterior to transverse groove conspicuously crenulate (Fig. 2A–C); (4) minor chela swollen, deeply concave ventrally between pollex and palm, and with dorsomesially margin of palm ending in prominent tooth (Fig. 2G, H); (5) both major and minor chelae densely setose, especially on mesial surfaces (Fig. 2C, G); (6) third pereiopod with relatively stout merus, distoventrally armed with strong, sharp tooth, and with strongly biunguiculate dactylus (Fig. 1I, J); and (7) second pleopod with appendix masculina reaching far beyond appendix interna and overreaching distal margin of endopod (Fig. 1N, O), separates the new species from all other members of the A. macrocheles group, including the morphologically closest species, A. collumianus (species complex) and A. deuteropus. The elongation of the appendix masculina in A. dingabadi sp. nov. appears to be unique within the A. macrocheles group. The degree of setation of the major and minor chelae, as seen in A. dingabadi sp. nov. is also uncommon, although a dense cover of very thin, almost fluffy setae is also present on the chelae of A. deuteropus (cf. Miya 1974: pl. 15, fig. C; Banner & Banner 1982: fig. 8d, g). The two species also share a similar frontal margin of the carapace and the presence of a prominent crest on the distomesial margin of the minor chela palm (cf. Banner & Banner 1982: fig. 8a, g). On the other hand, the well-known colour pattern of A. deuteropus, with its uniform dark brownred or grey-blue body and conspicuously red-and-white banded antennular and antennal flagella (e.g., Dehghani et al. 2019: fig. 2A, B), is very different from that of A. dingabadi sp. nov. Similarly, none of the species of the A. collumianus complex (A. Anker, pers. obs.) has a colour pattern that approaches that of A. dingabadi sp. nov. In addition, they appear to be morphologically more distinct from the new species than A. deuteropus (cf. Banner 1956; Miya 1974; Banner & Banner 1982)., Published as part of Anker, Arthur, 2023, A conspicuously coloured new species of the Alpheus macrocheles group from the central Indian Ocean (Decapoda: Alpheidae), pp. 155-162 in Zootaxa 5271 (1) on pages 156-160, DOI: 10.11646/zootaxa.5271.1.6, http://zenodo.org/record/7864449, {"references":["Abdulla, F. & O'Shea, M. (2005) English - Dhivehi. Dictionary. Dhivehi - English. A guide to the language of the Maldives. Version 1.0. Michael O'Shea, Belconnen, 157 pp.","Miya, Y. (1974) The Alpheidae (Crustacea Decapoda) of Japan and its adjacent waters, part II. Publications from the Amakusa Marine Biological Laboratory, Kyushu University, 3, 103 - 195, pls. 15 - 31.","Banner, D. M. & Banner, A. H. (1982) The alpheid shrimp of Australia, part III: the remaining alpheids, principally the genus Alpheus, and the family Ogyrididae. Records of the Australian Museum, 34, 1 - 357. https: // doi. org / 10.3853 / j. 0067 - 1975.34.1982.434","Dehghani, A., Sari, A. & Naderloo, R. (2019) Annotated checklist of the snapping shrimps of the genus Alpheus Fabricius, 1798 (Crustacea: Decapoda: Alpheidae) from the Persian Gulf and Gulf of Oman, Iran. Zootaxa, 4544 (4), 479 - 504. https: // doi. org / 10.11646 / zootaxa. 4544.4.2","Banner, A. H. (1956) Contributions to the knowledge of the alpheid shrimp of the Pacific Ocean, part I: collections from the Mariana Archipelago. Pacific Science, 10, 318 - 373."]}

Alpheus sulcipalma sp. nov. (Fig. 1–4) Material examined Holotype: male, CL 9.0 mm, Accession No.: NBFGR/ALPASUL–00, ID No. NBFGR: DBTLD220, Rocky dead coral, Agatti Island, Lakshadweep, India (10° 50ʹ 38.6" N, 072° 11ʹ 22.1" E), 0.5–1.0 m, hand net, December 2018. Paratypes: 1 male, CL 7.0 mm, Accession No: NBFGR/ALPASUL–01, ID No. NBFGR: DBTLD:42; 1 female, CL 4.5 mm, Accession No: NBFGR/ALPASUL–02, ID No. NBFGR: DBTLD:138, rocky dead coral, Agatti Island, Lakshadweep, India, 0.5–1.0 m, hand net, January 2019. 7 males, 10 females (CL 4.5–9.0 mm; Table 2) deposited in the Peninsular and Marine Fish Genetic Resources (PMFGR) centre, ICAR–NBFGR, India. ......Continued on the next page IN—India; FB—French Polynesia; US—United States Description Small-sized species (CL 3.5–9.0 mm). Carapace (Figs. 1A, B) glabrous, smooth, not setose or tubercle. Rostrum slender, acute, straight directed forward, overreaching mid-length of first article of antennular peduncle, lateral margin bearing few small setae on each side, without rostral carina. Orbital hood moderately swollen, slightly higher than rostrum; anterior margin unarmed; margin between rostrum and orbital hood slightly concave. Pterygostomial angle broadly rounded. Cardiac notch well developed and V-shaped. Pleon smooth, subcylindrical, and slightly compressed laterally; pleura of pleomeres 1–4 marginally rounded, pleuron of pleomere 2 subtriangular, pleuron of pleomere 5 with leaf-like process in posteroventrally (Fig. 3H). Telson (Fig. 1C) glabrous, trapezoidal, tapering posteriorly, about 2.6–3.1 times as long as posterior width and 2.0- 2.2 times as long as proximal width, dorsal surface smooth with 2 pairs of movable spines, proximal and second pairs located at 0.4 and 0.65 of telson length, respectively; posterior margin convex, fringed with numerous long plumose setae, posterolateral angle with 2 pairs of movable spines, mesial spine slender and about 2.0 times as long as a lateral one. Eyes (Figs. 1A, B) small, fully covered with orbital hood in both dorsal and lateral view; cornea dark in colour with globular shaped; anteromesial margin moderately produced and rounded in anteriorly. Antennular peduncle (Fig. 1B) moderately slender, almost reaching distal margin of antennal scaphocerite; stylocerite flatted, terminating an acute spine, overreaching distal margin of first article of peduncle; first article short, with broad ventromesial carina; second article long, about 2.1–2.4 times as long as maximum width and 1.6–1.7 times of first article; third article slightly longer than first article; antennular flagella asymmetrical in length; outer flagellum longest, slender, gradually tapering towards terminal units; outer flagellum short and thicker than inner flagellum with numerous aesthetascs on about 10–12 segments. Antenna (Fig. 1A, B) with basicerite rather moderate, ventrolateral margin armed with a slender spine; scaphocerite subovate, thin, tapering distally, about 3.1–3.2 times as long as maximum width, lateral margin straight and barely concave proximally, distolateral tooth well- developed, sharp, overreaching well beyond distal margin of lamella and third article of antennular peduncle; carpocerite long, thick, overreaching distal margin of scaphocerite; flagellum thick, tapering towards to terminal end and longer than antennular flagella. Mouthparts not dissected and typical for this genus in external observation. Third maxilliped (Fig. 1E) slender, slightly overreaching distal margin of antennal scaphocerite; antepenultimate article broad, longest, about 4.1–4.4 times as long as maximum width, dorsolateral margin with thick longitudinal ridge with numerous long setae, ventral surface with a sharp carina mesial to distal margin with numerous small setae; penultimate article short, about 0.42 times of antepenultimate and 2.3 times of maximum width, slightly widen distally, numerous long setae on dorsal and lateral surface; ultimate article about 4.6 times as long as maximum width and 1.8 times of penultimate article, tapering gradually towards terminal end, lateral and dorsal surface furnished with numerous rows of long serrulate setae, distal margin with stout long setae; exopod long, overreaching distal margin of antepenultimate article with numerous small setae on distally. Arthrobranch large with a faint terminal tooth with few setae on laterally (Figs. 1E, F). Male major cheliped (first pereiopod: Figs. 2A–D) broad, larger, laterally compressed, extending beyond distal margin of antennal scaphocerite by half length of palm and dactylus; ischium very short and stout; merus stout, broad, about 1.9 times as long as broad, distodorsal margin rounded and extended somewhat angled; mesial margin smooth, with few simple small and long setae, distomesial angle blunt, without tooth; carpus very short, cup-shaped, distodorsal margin with few long setae; chela broad, stout, slightly compressed, about 2.3 times as long as broad, 1.5 times of CL, surface almost smooth with numerous minute hairy form setae, minute granulated with numerous setae on distal half of palm and fingers; dorsal shoulder rounded, strongly overhanging, latter deep notch, moderately broad; dorsolateral surface of palm with deep shallow triangular groove extending from posterior of dorsal notch, mixing with linea impressa and continued posteroventral of lateral surface; dorsomesial surface with longitudinal groove extending from dorsal notch to near linea impressa anteriorly; ventral shoulder blunt, rounded, protruding forward, with deep ventral notch, somewhat 'V' shaped; ventromesial and ventrolateral surface with broad and deep groove on each side; fingers about 0.63 times as long as palm; pollex stout, terminating acute tip and curved, distolateral margin of socket for plunger sinuous with a small tooth like process, distomesial margin with sinuous; dactylus slightly longer than pollex, distal margin broadly rounded, acute tip, with few stout setae; plunger stout, broad, smooth, anteriorly rounded and thick in distally; adhesive discs well found on dactylus and palm. Female major cheliped (first pereiopod Figs. 2E, F) broad, extending beyond the distal antennal scaphocerite by three fourth of chela; ischium, merus and carpus similar with male; chela broad, stout, slightly swollen, about 2.4 times as long as broad, 1.1 times of CL, surface with numerous minute and long hairy form setae; dorsal shoulder rounded, strongly overhanging with deep broad notch; dorsolateral surface of palm with deep shallow and narrow triangular groove extending from posterior of dorsal notch, mixing with linea impressa and continued posteroventral of lateral surface; dorsomesial surface with narrow triangular groove extending from dorsal notch to near linea impressa; ventral shoulder blunt, rounded, protruding forward, with deep ventral notch, more or less 'U' shaped; ventromesial and ventrolateral surface with broad and deep groove on each side; fingers compressed, about 0.51 times as long as palm; pollex stout, terminating acute tip and curved, distomesial margin with sinuous; dactylus slightly longer than pollex, distal margin very broadly rounded, with few small setae; plunger stout, smooth, rounded anteriorly and thick in distally; adhesive discs well found on both dactylus and palm. Minor cheliped (Figs. 2G, H) slender, simple, and non-balaeniceps in males and females. Minor cheliped of male very smaller than major cheliped; ischium very short; merus stout, about 2.1–2.9 times as long as broad, distodorsal margin somewhat angle sharped in lateral view; mesial margin with few simple small and long setae, distomesial angle blunt and rounded; carpus short, cup-shaped; chela moderately stout, about 3.8 times as long as broad, surface smooth with few setae, especially dorsolateral surface with very setose; palm about 1.6 times as long as height with a prominent form of linea impressa proximally, distodorsal margin slightly depressed; fingers equal in length, about 1.3 times as long as palm, without any ridges or grooves, with a strong hook-like tooth and crossing distally, covered with small and long setae; dactylus slightly slender than pollex, a proximal site with two small crests, cutting edges simple, thin, blade-like with small setae throughout the edges; adhesive discs small on dactylus and palm. Female minor cheliped similar with the male. Second pereiopod (Fig. 3A) slender, simple, overreaching distal margin of scaphocerite by the length of carpus and chela; ischium long, about 6.0 times as long as broad; merus equal length to ischium without any spine; carpus subdivided into five segments, proximal segment longest and about 5.5 times as long as width, fifth segment twice the length of fourth; a ratio of carpal segments 4.0: 2.7: 1.0: 1.0: 2.0; chela simple, about 1.8 times as long as a terminal segment of carpus; fingers slightly longer than palm furnished with several groups of simple setae. Third pereiopod (Figs. 3B, C) moderately slender, overreaching distal margin of scaphocerite by length of dactylus; ischium with a movable spine; merus broad, about 4.0 times as long as maximum width with minute several setae on dorsally, distomesial margin unarmed; carpus about 4.1 times as long as width, 0.6 times of merus, and slender than merus; propodus about 6.0 times of maximum width, 0.7 times of merus, ventral margin with a row of 7–8 spiniform setae and a pair of spiniform setae in distoventral margin; dactylus simple, conical, long, curved, with acute terminal end, about 0.4 times as long as propodus. Fourth pereiopod (Figs. 3D, E) similar to third one, slightly slender, overreaching scaphocerite by length of dactylus; ischium with a movable spine; propodus about 6.1 times as long as width, ventral margin armed with 7–8 stout spiniform setae and a pair in distoventral margin with several simple and long setae; dactylus similar with third pereiopod. Fifth pereiopod (Figs. 3F, G) more slender than third and fourth, reaching mid-length of scaphocerite; ischium without ventrolateral spine; merus about 6.7 times as long as maximum width, 0.75 times of carpus; carpus slender than merus about 6.3 times of width; propodus about 7.3 times as long as width, with numerous long setae in dorsal and ventral surface, a row of 6–7 spiniform setae in ventral margin with a pair in distally, and distoventral half covered with 8–10 transverse rows of serrulate setae found as cleaning brush; dactylus about 0.38 times of propodus and similar with third pereiopod. Male first pleopod (Fig. 1G) with a reduced form of endopod, much smaller than exopod, bearing few stiff setae along the distal margin, about 0.2 lengths of exopod. Male second pleopod with similar length of exopod and endopod, along the distal margin covered with long setae; appendix masculina small and thin about 0.82 times as long as appendix interna, lateral and distal margin covered with long stiff setae (Fig. 1H). Female first pleopod (Fig. 1I) endopod reduced, spatulate shaped, with few distal setae; second pleopod similar to those of male; protopod broad, thick with several long setae on inner margin (Fig. 1J). Uropod (Fig. 1D) broadly rounded and slightly overreaching distal margin of telson; protopod with inner and outer lobes each produced into a triangular acute tooth, distolateral margin with few long tuft setae; exopod broad and almost equal length to endopod, bearing a small distolateral tooth and a stout spiniform distolateral seta; diaeresis barely trilobed, distal margin fringed with long plumose setae; endopod fringed with long setae from inner to distal margins. Colour pattern The live colour pattern (Fig. 4A) of the new species shows exhibits sexual dimorphism. Generally, males with carapace, antennal segments, abdomens, pereiopods, pleopods, uropod, and telson translucent light greenish; major pereiopod fingers with light violet in distally. In dead condition, body with a translucent light brown ( Fig. 4B). Female body and appendages are generally translucent light violet; distal of half of the fingers in major pereiopod with light violet. Habitat All the specimen were found in the crevices of dead and live coral rocks in the intertidal zone of Agatti Island at 0.5 to 1.0 m depths (see Table 2). M—Male; F B—Female Berried; F—Female Distribution Presently known only from waters of Agatti Island of Lakshadweep, India. Parasite Some of the individuals of A. sulcipalma sp. nov. were found infested with bopyrid parasite, Argeiopsis inhacae Kensley, 1974, in branchial region of cephalothorax. Carapace shows a bulbous structure which can be seen in lateral view (fig 4B–D). Most of the boyprid parasites were adult females carrying numerous matured eggs. Etymology The species name is derived from Latin, Sulcus (= groove or depression) and palma (palm), which refer to the strong triangular structure of groove on the dorsolateral surface of palm in major cheliped of both the sexes.

Alpheus sulcipalma sp. nov. (Fig. 1–4), Published as part of Purushothaman, P., Bharathi, S., Damodhar, A. T., Ajith Kumar, T. T. & Lal, K. K., 2023, Morphological and molecular approaches revealed a new species of snapping shrimp genus Alpheus Fabricius, 1798 (Decapoda: Caridea: Alpheidae) from Lakshadweep Islands, India, pp. 426-442 in Zootaxa 5227 (4) on page 428, DOI: 10.11646/zootaxa.5227.4.2, http://zenodo.org/record/7518860

Alpheus nuttingi (Schmitt, 1924) Examined material. Munipality of Amapá. Maracá-Jipioca Ecological Station, beach from Jacal (02°08’20.24”N, 50°25’10.35”W), 22.xi.1995, H.P. Belo and J.O. Cardoso, 1 male and 1female (IEPA 54); idem, stream Herculano (02°06’34.21”N, 50°24”29.78”W), 26.xi.1995, H.P. Belo and J.O. Cardoso, 4 males, 4 females and 4 females with eggs (IEPA 55); left margin from Igarapé Herculano, 26.xi.1995, H.P. Belo and J.O. Cardoso, 3 males and 1 female (IEPA 56); estuary of Sucuriju River (01°40’04.21”N, 49°55101.45”W), 25.vii.1996, H.P. Belo, J.O. Cardoso and I.M. Vieira, 1 male, 4 females and 4 females with eggs (IEPA 95); estuary of Sucuriju River (01°40’17.0”N, 49°55’06.0”W), 25.ix.1996, H.P. Belo, J.O. Cardoso and I.M. Vieira, 14 males, 6 females and 15 females with eggs (IEPA 110) estuary of Sucuriju River, xi.1996, H.P. Belo, J.O. Cardoso and I.M. Vieira, 12 males and 8 females (IEPA 122); estuary of Sucuriju River, 03.iv.1997, H.P. Belo, J.O. Cardoso and I.M. Vieira, 10 males, 2 females and 13 females with eggs (IEPA 128) (Figure 2). Geographic distribution. Western Atlantic, South Florida, Southwest Gulf of Mexico and West Indies to Brazil (States of Ceará to Santa Catarina) (Coelho et al. 2006; Anker 2007; Santos et al. 2012; Soledade & Almeida 2013). Distribution in Amapá. Oceanic island of Maracá-Jipioca and mouth of the Sucuriju River. Previous records. present study Ecological notes. Species found in small tunnels built in more compacted soil mixed with decomposing vegetation in the mangrove, in the intertidal bands. Its presence in the environment is noticed by the emission of clicks caused by its fingers. Remarks. The morphology of the specimens in this study corresponds to the descriptions by De Grave et al. (2008)., Published as part of Vieira, Inacia Maria, Silva, Luiz Maurício Abdon, Almeida, Alini Gomes Santiago De, Almeida, Danilo Pelaes De, Silva-Júnior, Orleno Marques & Tavares-Dias, Marcos, 2022, Diversity, distribution and new records of freshwater and estuarine shrimp in the state of Amapá, eastern Brazilian Amazon region, pp. 41-71 in Zootaxa 5178 (1) on page 44, DOI: 10.11646/zootaxa.5178.1.3, http://zenodo.org/record/7021864, {"references":["Coelho, P., Almeida, A., Fidelis, J., Bezerra, L. & Giraldes, B. (2006) Diversity and distribution of the marine and estuarine shrimps (Dendrobranchiata, Stenopodidea and Caridea) from North and Northeast Brazil. Zootaxa, 1221 (1), 41 - 62. https: // doi. org / 10.11646 / zootaxa. 1221.1.5","Santos, P. S., Soledade, G. O. & Almeida, A. O. (2012) Decapod crustaceans on dead coral from reef areas on the coast of Bahia, Brazil. Nauplius, 20, 145 - 169. https: // doi. org / 10.1590 / S 0104 - 64972012000200007","Soledade, G. O. & Almeida, A. O. (2013) Snapping shrimps of the genus Alpheus Fabricius, 1798 from Brazil (Caridea: Alpheidae): updated checklist and key for identification. Nauplius, 21, 89 - 122. https: // doi. org / 10.1590 / S 0104 - 64972013000100010","De Grave, S., Cai, Y. & Anker, A. (2008) Global diversity of shrimps (Crustacea: Decapoda: Caridea) in freshwater. Hydrobiologia, 595, 287 - 293. https: // doi. org / 10.1007 / s 10750 - 007 - 9024 - 2"]}

Alpheus thompsoni Anker, 2022 (Figs. 6–8; 9A) Alpheus thompsoni Anker 2022: 274, figs. 1–3, 8A, B. Alpheus ochrostriatus.— Karplus et al. 1981: 6, fig. 2F (black-and-white photograph); Debelius 1997: 203 (colour photograph); Debelius 2001: 150 (part.), 1 colour photograph (yellow morph); Randall et al. 2003: 512, fig. 8 (colour photograph); Randall 2005: 513; Poupin 2010: 33; Karplus & Thompson 2011: 591, fig. 4.4.10-B (black-and-white photograph) [nomen nudum]. Alpheus “ ochrostriatus ”. — Anker 2000: 3, fig. 2 (colour photograph); Jaafar & Randall 2009: 29, pl. 2A (colour photograph) [nomen nudum]. Alpheus sp. ‘ ochrostriata ’.— Kuiter & Debelius 2009: 151, 3 colour photographs [nomen nudum]. Alpheus djiboutensis.— Yu et al. 1996: 35, figs. 19, 20 (colour photographs) [not A. djiboutensis De Man, 1909]. Alpheus sp. 4.— Minemizu 2013: 102, colour photograph. Material examined. 1 male (cl 10.0 mm), FLMNH UF 37002, Saudi Arabia, Red Sea, Farasan Islands, Zahrat Durakah, 16°50’09.2”N, 42°18’22.7”E, fringing reef slope around sandy island, depth 2–6 m, leg. A. Anker, P. Norby, G. Paulay, 11.03.2013 [fcn BDJRS-2689]. Description. See Anker (2022). Complementary illustrations of the general morphology of the Red Sea male are provided in Figs. 6, 7. The variation in the antennal scaphocerite and major and minor chelipeds are discussed below. Colour pattern. Anker (2022) described in detail the colour pattern of the female holotype of A. thompsoni from Madang, Papua New Guinea. This colour pattern is also present in shrimps from numerous other localities in Papua New Guinea and Indonesia (e.g. Kuiter & Debelius 2009: Flores and Bali; Fig. 9A, Sulawesi). In the male from Saudi Arabia (Fig. 8A, B), the red parallel-running longitudinal bands are simple, i.e. they form simple, elongate, red strings closed at both ends, which is in marked contrast to the more complex bands of the holotype of A. thompsoni (cf. Anker 2022: fig. 3A, B, 8A, B), in which they are broader and filled with a wealth of finer anastomosing bands and streaks (also visible in Fig. 9A). See also discussion below. Distribution. Indo-West Pacific from the Red Sea to southern Japan, Indonesia and New Caledonia (Karplus et al. 1981; Debelius 1997, 2001; Anker 2000, 2022; Randall 2005; Kuiter & Debelius 2009; Poupin 2010; present study). Ecology. Coral reefs and adjacent reef flats; associated with several species of gobiid fishes, including Amblyeleotris wheeleri (Polunin & Lubbock, 1977), A. steinitzi (Klausewitz, 1974), A. guttata (Fowler, 1938), A. fasciata (Herre, 1953), Ctenogobiops pomastictus Polunin & Lubbock, 1977, and C. tangaroai Lubbock & Polunin, 1977 (Karplus et al. 1981; Randall et al. 1998, 2003, 2007; Anker 2000, 2022; Debelius 2001; Randall 2005; Kuiter & Debelius 2009; Jaafar & Randall 2009; Karplus & Thompson 2011). Remarks. The present complete male specimen from Saudi Arabia was initially considered to belong to an undescribed species closely related to A. thompsoni due to some discrepancies in the colour pattern (see above) and a small difference in the shape of the scaphocerite (see below). Most shrimps identified as “ A. ochrostriatus ” or “ Alpheus sp. ” in the published or unpublished colour photographs analysed by the author could be classified as either the species-diagnostic type A pattern, with broader red bands containing fine, often anastomosing lines (Fig. 9A; see also Anker 2022: fig. 3), or the type B pattern, with narrower red bands not filled with fine lines (Fig. 8A, B, D). It is important to mention that the holotype female at cl 13.3 mm is only slightly larger than the present male at cl 10.0 mm; thus, these two colour pattern types are comparable since they are present in adult individuals. However, after a more detailed analysis and inclusion of more photographic material, the author came to the conclusion that the type A and type B patterns are part of the same general colour pattern of A. thompsoni. The most pivotal observation for this conclusion was an in situ colour photograph in Kuiter & Debelius (2009: 151, upper photograph), showing two individuals cohabiting the same burrow, a larger female at the burrow entrance apparently (slightly out of focus) with the type A pattern and a smaller male in front of her clearly with the type B pattern. In addition, an indication of an intermediate state between the type A and type B pattern can be seen on the pleon of the specimen from southern Taiwan (Fig. 8C; same photograph as in Yu et al. 1996: fig. 19). Differences in the proportions of the major and minor chelipeds between males and females are expected. As is the case of many other species of the A. brevirostris group, both chelipeds of the Saudi Arabian male of A. thompsoni (Fig. 7A, B, E) are larger and stronger than their homologs in the female holotype from Papua New Guinea (Anker 2022: fig. 2). The male minor chela is also lacking balaeniceps ridges and setae (Fig. 7D, E), thereby confirming their absence in both sexes in A. thompsoni. The ventromesial margin of the merus of both chelipeds is unarmed in the present male (Fig. 7A, D, F) and also has a reduced armature, with only one spiniform seta present on the minor cheliped merus, in the holotype female (Anker 2022: fig. 2C, F). The presence of spiniform setae on the cheliped merus is usually a consistent character, although their number may be variable; in addition, they break off easily or may be missing entirely in regenerated chelipeds. The relative proportions of the carpal subarticles of the second pereiopod is only slightly different, with the first subarticle being slightly than the second in the holotype female (Anker 2022: fig. 1F) vs. the two being almost equal in length in the present male (Fig. 7). The only remaining significant difference between the female holotype from Papua New Guinea and the male from Saudi Arabia lies in the shape of the distal region of the antennal scaphocerite. In the female, the blade extends slightly beyond the distolateral tooth (Anker 2022: fig. 1A), whereas in the male, it is noticeably shorter than the distolateral tooth (Fig. 6A). This difference can be explained either by the somewhat worn distolateral tooth of the holotype (which normally would be slightly longer) or by intraspecific variability. With only two specimens of A. thompsoni available, it is difficult to assess the variation observed in some morphological characters, as well as in the colour pattern. Although the involvement of a second species cannot be excluded at this stage, it seems to be less likely based on the presently available data. The planned DNA analyses of the entire A. djeddensis — A. djiboutensis group should eventually resolve this issue and confirm whether or not the assignment of the Saudi Arabian male to A. thompsoni is correct. As already pointed out by Anker (2022), several other types of colour pattern with parallel-running lines seem to correspond to further species, some of them closely related to A. thompsoni. For instance, a very distinctive colour pattern, hereafter type C pattern, exists in some snapping shrimps from Indonesia (Fig. 9B, C; see also Debelius 2001: 150, upper photograph under “ A. ochrostriatus ”). The type C pattern differs from both pattern types of A. thompsoni (types A and B, as defined above), in the colour of the second to fifth pereiopods, which are deep blue with bright yellow markings near articulations (not yellow, as in A. thompsoni), and in the background colour and general pattern of the major and minor chelae, which are dark greyish-brown or brown-green with pale bluish blotches and spots (not yellowish with longitudinal red lines / blotches, as in A. thompsoni). The shrimps of the type C pattern possibly correspond to Alpheus sp. from the northern Red Sea in Karplus et al. (1981: fig. 2H, black-andwhite photograph in low resolution) and may well represent a further undescribed taxon, although several other species of the A. brevirostris group need be examined and compared. In the same publication of Debelius (2001: 150, also under A. ochrostriatus), the middle photograph shows a pink-reddish shrimp with red longitudinal lines, which is generally similar to the type B pattern of A. thompsoni. This shrimp has a dull pink (instead of yellowish) background, pink-reddish legs (not yellow as in types A and B, or blue with yellow spots as in type C) and pinkish antennal flagella (Fig. 9D; see also Anker 2022: fig. 8E). This colour pattern, hereafter the type D pattern, probably represents either A. mannarensis or a closely related undescribed taxon with affinities to both A. mannarensis and A. thompsoni (Anker 2022; but see Miya’s view of colour pattern variation in Banner & Banner 1981). Further complicating the issue is the presence in the Red Sea of a dark red Alpheus sp. with a balaeniceps minor chela (Fig. 9E, F), and with a colour pattern similar to that of A. mannarensis and the above-defined type D pattern. This colour pattern, hereafter the type E pattern, is defined by numerous, closely parallel-running lines of dark-red colour, the purplelilac antennal flagella and the uniform dark red-purple chelae (Fig. 9F; see also Minemizu 2013: 103, Alpheus sp. 6). Although the species with the type E pattern was not included in Karplus et al. (1981), Dr. I. Karplus was aware of its presence and had some colour slides of it (I. Karplus, pers. comm.). The type E pattern is also reminiscent of the colour pattern of A. fenneri, although the latter species’ body is uniform orange brown, apparently without red lines (Bruce 1994: fig. 5). In summary, the identification of goby-associated Alpheus spp. with the colour pattern types C–E will only be possible after the collection of photo-vouchered material the Red Sea and elsewhere in the Indo-West Pacific., Published as part of Anker, Arthur, 2022, On two goby-associated snapping shrimps from the Red Sea, one of them new to science (Malacostraca: Decapoda: Alpheidae: Alpheus), pp. 421-438 in Zootaxa 5105 (3) on pages 431-436, DOI: 10.11646/zootaxa.5105.3.5, http://zenodo.org/record/6332829, {"references":["Anker, A. (2022) Description of two new species of goby-associated snapping shrimps from the tropical western Pacific (Decapoda: Alpheidae: Alpheus). Zootaxa, 5092 (3), 273 - 290. https: // doi. org / 10.11646 / zootaxa. 5092.3.2","Karplus, I., Szlep, R. & Tsurnamal, M. (1981) Goby-shrimp partner specificity. I. Distribution in the northern Red Sea and partner specificity. Journal of Experimental Marine Biology and Ecology, 51, 1 - 19. https: // doi. org / 10.1016 / 0022 - 0981 (81) 90151 - 9","Debelius, H. (1997) Indian Ocean reef guide. IKAN, Frankfurt, 320 pp.","Debelius, H. (2001) Crustacea guide of the world. IKAN, Frankfurt, 321 pp.","Randall, J. E., Shao, K. T. & Chen, J. P. (2003) A review of the Indo-Pacific gobiid fish genus Ctenogobiops, with descriptions of two new species. Zoological Studies, 42, 506 - 515.","Randall, J. E. (2005) Reef and shore fishes of the South Pacific. New Caledonia to Tahiti and the Pitcairn Islands. University of Hawaii Press, Honolulu, 707 pp.","Poupin, J. (2010) Biodiversite de l'Indo-Pacifique tropical francais. 2514 especes de crustaces decapodes et stomatopodes. Rapport Scientifique, Institut de Recherche de l'Ecole Navale, Brest, 76 pp.","Karplus, I. & Thompson, A. R. (2011) The partnership between gobiid fishes and burrowing alpheid shrimps. In: Patzner, R. A., Van Tassell, J. L., Kovacic, M. & Kapoor, B. G. (Eds.), The biology of gobies. Science Publishers, Enfield, New Hampshire, pp. 559 - 608.","Anker, A. (2000) Taxonomical problems of the goby-associated species of Alpheus (Decapoda, Alpheidae). IOP Diving News, 11 (8), 1 - 7.","Jaafar, Z. & Randall, J. E. (2009) A pictorial review and key to the shrimp gobies of the genus Amblyeleotris of the Red Sea, with description of a new species. Smithiana Bulletin, 10, 23 - 29.","Kuiter, R. H. & Debelius, H. (2009) World atlas of marine fauna. IKAN, Frankfurt, 725 pp.","Yu, H. P., Jeng, M. S., Chan, T. Y., Ho, P. H. & Shy, J. Y. (1996) Studies on the land and aquatic decapod crustacean fauna of the Kenting National Park. Kenting National Park, Ministry of the Interior, Pingtung, ii + 79 pp. [in Chinese]","De Man, J. G. (1909) Diagnoses of new species of macrurous decapod Crustacea from the \" Siboga-Expedition \". Tijdschrift der Nederlandse Dierkundige Vereeniging, 2, 99 - 125.","Minemizu, R. (2013) Coral reef shrimps of Indo-West Pacific. Bun-ichi Sogo Shuppan, Co. Ltd., Tokyo, 144 pp. [in Japanese]","Polunin, N. V. C. & Lubbock, R. (1977) Prawn-associated gobies (Teleostei: Gobiidae) from the Seychelles, western Indian Ocean: systematics and ecology. Journal of Zoology, London, 183, 63 - 101. https: // doi. org / 10.1111 / j. 1469 - 7998.1977. tb 04174. x","Randall, J. E., Allen, G. R. & Steene, R. C. (1998) Fishes of the Great Barrier Reef and Coral Sea. University of Hawaii Press, Honolulu, 594 pp.","Randall, J. E., Shao, K. T. & Chen, J. P. (2007) Two new shrimp gobies of the genus Ctenogobiops (Perciformes: Gobiidae), from the Western Pacific. Zoological Studies, 46, 26 - 34.","Purushothaman, P., Abhilash, C. P., Ajith Kumar, T. T. & Lal, K. K. (2021) A new alpheid shrimp, Alpheus mannarensis sp. nov. (Crustacea; Decapoda; Alpheidae) from Gulf of Mannar, southern India. Zootaxa, 5026, 127 - 135. https: // doi. org / 10.11646 / zootaxa. 5026.1.5","Banner, D. M. & Banner, A. H. (1981) Annotated checklist of the alpheid shrimp of the Red Sea and Gulf of Aden. Zoologische Verhandelingen, Leiden, 190, l - 99.","Bruce, A. J. (1994) Alpheus fenneri sp. nov. and A. williamsi sp. nov., two new Indo-West Pacific alpheid shrimps of the brevirostris species group. The Beagle, Records of the Museums and Art Galleries of the Northern Territory, 11, 15 - 28. https: // doi. org / 10.5962 / p. 264023"]}

Alpheus karplusi sp. nov. (Figs. 1–4) Alpheus rubromaculatus.— Karplus et al. 1981: 6, fig. 2C (black-and-white photograph); Karplus 1987: 514, fig. 2C; Debelius 2001: 153 (part.), 1 colour photograph (shrimp from Egypt only) [nomen nudum]. Alpheus djeddensis.— Vine 1996: 104 (colour photograph) [not A. djeddensis Coutière, 1897]. See discussion for other records of Alpheus rubromaculatus [not A. karplusi sp. nov.]. Type material. Holotype: male (cl 10.5 mm), FLMNH UF 37011, Saudi Arabia, Red Sea, Farasan Islands, Zahrat Durakah, 16°50’09.2”N, 42°18’22.7”E, fringing reef slope around sandy island, depth 2–6 m, leg. M. Berumen, A. Anker, P. Norby, 11.03.2013 [fcn BDJRS-2701]. Paratypes: 1 male (cl 11.4 mm), FLMNH UF 37016, same collection data as for holotype [fcn BDJRS-2706]; 1 female (cl 7.4 mm), FLMNH UF 37070, Saudi Arabia, Red Sea, off Thuwal, Shib Nazar East offshore reef, 22°19’19.2”N, 38°51’18.0”E, depth about 10 m, leg. A. Anker, P. Norby, J. Moore, 16.03.2013 [fcn BDJRS- 2827]. Description. Carapace glabrous, without setae, sparsely pitted (Fig. 1A, B). Rostrum well developed, moderately slender, about 1.8–2.2 times as long as wide at base, straight, subacute distally, almost reaching distal margin of first article of antennular peduncle; rostral carina well developed, rounded dorsally, gently sloping into shallow adrostral furrows, continuing well beyond base of orbital hoods, gradually widening and fading, not reaching midlength of carapace (Fig. 1A, B). Orbital hoods swollen, somewhat projecting anteriorly in lateral view, unarmed; frontal margin between rostrum and orbital hood shallowly concave (Fig. 1A, B). Pterygostomial angle rounded (Fig. 1B); cardiac notch deep. Telson broad, subrectangular, gently tapering distally, about 1.7 times as long as maximal width, with lateral margins slightly convex at about 0.7 of its length, more noticeably tapering in posterior third; dorsal surface with two pairs of stout short cuspidate setae both inserted at some distance from lateral margin, first pair near telson mid-length, second pair between 0.7 and 0.8 telson length; posterior margin broadly rounded, with row of slender spiniform setae above plumose setae; posterolateral angles each with one pair of spiniform setae, mesial ones stouter and almost three times as long as lateral ones (Fig. 1C). Eyes with large, well-pigmented silvery cornea (Figs. 1A, B; 3A, B; 4B, D). Antennular peduncle moderately long and stout; stylocerite slightly swollen laterally, ending in sharp point, latter reaching but not significantly overreaching distal margin of first article; ventromesial carina with anteriorly directed, shark-fin shaped tooth; second article about 2.7 times as long as wide; lateral antennular flagellum with secondary ramus fused to main ramus over most of its length, with numerous groups of aesthetascs distally, starting from about 13th subdivision (Fig. 1A, B, D). Antenna with basicerite moderately stout, armed with sharp tooth on distoventral margin; scaphocerite with lateral margin shallowly and broadly concave; blade moderately broad, not reaching beyond stout, subacute distolateral tooth; scaphocerite overreaching both end of antennular peduncle and end of carpocerite (Fig. 1A, B). Mouthparts not dissected, typical for genus in external observation. Third maxilliped relatively slender, elongate, distally setose; coxa with bluntly projecting lateral plate; antepenultimate article flattened ventrolaterally, with distinct ridge running parallel to dorsal margin on lateral surface and almost smooth mesial margin, almost six times as long as high; penultimate article cup-shaped, not bulging ventrally, distally widening, its distoventral margin with tuft of moderately long setae, latter not reaching half-length of ultimate article; ultimate article densely furnished with rows of elongate stiff setae, dorsal and apical setae longest; arthrobranch well developed (Fig. 1E). Major cheliped moderately robust; ischium stout, smooth; merus moderately stout, trigonal in cross-section, about three times as long as maximal width, distodorsal margin ending bluntly, ventromesial margin slightly rugose, with blunt distal tooth, armed with three spiniform setae; carpus short, wide, cup-shaped; chela not particularly elongate, with long setae along dorsal and ventral margins; palm strongly compressed, subrectangular in cross-section, length / maximal height ratio about 1.9–2.1; surfaces relatively smooth, not granulated; dorsal margin with well-marked transverse groove subdistally, without pronounced longitudinal ridges; ventral margin feebly convex; fingers somewhat unequal in length with dactylus slightly longer than pollex, both about half-length of palm, not twisted or significantly deviating from chela axis; dactylus distally rounded, with plunger greatly reduced, poorly demarcated from anterior cutting edge, latter feebly bulging; adhesive disks small (Fig. 2A–C). Minor cheliped relatively slender, not sexually dimorphic; ischium short, smooth; merus slightly slenderer than that of major cheliped, trigonal in cross-section, about 3.5 times as long as maximal width, distodorsal angle blunt, ventromesial margin slightly rugose, with small blunt distal protuberance, armed with four small spiniform setae; carpus longer than that of major chela, distally widening, vase-shaped; chela not particularly elongate or swollen, slender, fringed with long setae along dorsal and ventral margins; palm compressed, oval in cross-section, length / maximal height ratio about 2.2; surfaces smooth, not granulated; dorsal margin without groove; ventral margin straight; fingers subequal in length, approximately as long as palm, straight, slightly gaping when closed, not balaeniceps; cutting edges of each finger with broadly triangular protuberance proximally; adhesive disks reduced (Figs. 2D, E; 3A). Second pereiopod slender; ischium and merus subequal in length; carpus with five subarticles, first much longer than second, ratio of carpal subdivisions approximately equal to 2.3: 2.2: 1.0: 0.9: 1.5; chela longer than distalmost carpal subdivision (Fig. 2F). Third pereiopod relatively slender; ischium with stout spiniform seta on ventrolateral surface; merus almost five times as long as maximal width, ventromesial margin unarmed distally; carpus about half-length of merus, much slenderer than merus, unarmed; propodus slightly longer than carpus, moderately setose, with about six spiniform setae along ventral margin and one distal pair of spiniform setae on distoventral angle near base of dactylus; dactylus about 0.4 length of propodus, gradually curving distally, subspatulate, moderately broadened, flattened ventrally, with longitudinal keel fringed by several tufts of setae (Fig. 2G–I). Fourth pereiopod generally similar to third pereiopod, somewhat slenderer. Fifth pereiopod slenderer than third and fourth pereiopods; ischium with small spiniform seta on ventrolateral surface; merus about eight times as long as wide; carpus slenderer than merus, about 0.9 length of merus; propodus subequal to carpus, with rows of serrulate setae forming cleaning brush on distal third of ventrolateral surface, and eight or so spiniform setae along ventromesial margin, in addition to distal spiniform seta on mesial side of ventrodistal angle adjacent to base of dactylus; dactylus almost conical, only slightly broadened (Fig. 2J, K). Second pleopod with appendix masculina about 0.8 length of appendix interna, with long stiff setae, mainly along outer margin and on apex (Fig. 1F). Uropod with mesial and lateral lobes of protopod each ending in sharp distal tooth; exopod broad, somewhat truncate distally, with blunt triangular distolateral tooth; diaeresis straight for most part, except for blunt lobe adjacent to stout spiniform seta, latter not reaching level of distal margin of exopod; endopod narrower than exopod, with row of small spiniform setae above plumose setae on distal margin (Fig. 1H, I). Colour pattern. General background semi-opaque pale buff, sometimes with yellowish or greenish tinge; carapace and pleon with larger or smaller, irregularly shaped (some rounded) and sized spots, blotches and short transverse bands of rusty-brown to orange-brown colour, some more or less organised in longitudinal lines, others without particular organisation; spots more numerous on pleon, tail fan and chelipeds, being somewhat larger on pleon flanks and chelae; second pereiopods and walking legs also with rusty-brown spots or transverse bands; posterior region of carapace and first pleonite with more or less conspicuous white saddle; antennular flagella yellowish; fingertips of major chela pinkish; ovaries in females (visible by translucence) olive greenish (Figs. 3, 4). Etymology. This remarkable new species of Alpheus is named after Dr. Ilan Karplus for his important contribution to the knowledge of the biology and ecology of goby-shrimp symbioses (Karplus et al. 1981; Karplus 1987, 1992; Karplus & Thompson 2011 and references therein). Distribution. North-Western Indian Ocean: presently known only from the Red Sea, including Saudi Arabia (Farasan Islands and Thuwal, type material), Egypt (El Quseir, Safaga, Nuweiba; Fig. 4E–G) and Israel (Eilat, see Karplus et al. 1981; Fig. 4D). Ecology. Coral reefs and adjacent reef flats with abundance of silt-covered rubble, at depths of 5–40 m (usually less than 20 m); associated exclusively with the whitecap shrimp goby Lotilia graciliosa Klausewitz, 1960 (Vine 1996; Karplus 1987; Karplus & Thompson 2011; Debelius 2011; for the most recent review of the genus Lotilia Klausewitz, 1960 see Shibukawa et al. 2012). Remarks. Alpheus karplusi sp. nov. has a striking and unmistakable, and therefore highly diagnostic, colour pattern, which is characterised by the presence of rusty-brown spots on a generally pale buff background (Figs. 3, 4). This colour pattern is in sharp contrast to all other known patterns among the goby-associated species of the A. djeddensis — A. djboutensis complex, separating A. karplusi sp. nov. at once from A. djiboutensis (sensu Holthuis 1958), A. bellulus, A. macellarius, A. fenneri, A. mannarensis, A. thompsoni, A. sciolii (see Miya & Miyake 1969; Bruce 1994; Debelius 2001; Kuiter & Debelius 2009; Minemizu 2013; Anker et al. 2015; Anker 2022), as well as the species called “ A. purpurilenticularis ” [nomen nudum] by Karplus et al. (1981). The colour pattern of A. djeddensis currently remains unknown, but the preliminary results of a morphological analysis of the Red Sea material morphologically corresponding best to A. djeddensis show that this species’ colour pattern is also very different from that of A. karplusi sp. nov. (A. Anker, in prep.). It is perhaps noteworthy that two more distantly related species of the A. brevirostris group, namely A. pubescens De Man, 1909 and A. williamsi Bruce, 1994, also have brown spots and blotches, but not as distinct and well-separated as in the new species (cf. Bruce 1994; Anker & De Grave 2016). Based on morphological grounds alone, A. karplusi sp. nov. can be easily separated from A. djeddensis, A. djiboutensis, A. bellulus, A. fenneri and A. sciolii by the minor chela being much slenderer and lacking balaeniceps setae; these are present at least in males (or possibly in both sexes) in all the other species (cf. Coutière 1899; De Man 1909; Miya & Miyake 1969; Bruce 1994; Anker 2022). In addition, A. karplusi sp. nov. differs from A. djiboutensis, A. bellulus and A. sciolii by the noticeably slenderer cheliped meri, especially that of the minor cheliped; from A. djeddensis and A. fenneri by the strongly concave lateral margin of the antennal scaphocerite (vs. almost straight in the other two species); from A. djiboutensis and A. djeddensis by the glabrous carapace and pleon (vs. with more or less developed pubescence in the other two species); from A. djiboutensis and A. sciolii by the distolateral tooth of the antennal scaphocerite only slightly overreaching the blade (vs. reaching far beyond the blade in the other two species); from A. djiboutensis and A. bellulus by the noticeably slenderer major chela, with the palm twice as long as high (vs. at most 1.5 times in the other two species); from A. bellulus and A. fenneri by the less protruding rostro-orbital area and the second pereiopod with the first carpal subarticle clearly longer than the second (vs. the two being subequal in length in A. bellulus and A. fenneri); from A. bellulus by the broader rostral carina and slenderer third pereiopod (noticeable in the width-length proportions of the merus); from A. fenneri, a much larger species at cl 19.5–23.0 mm, by the smooth major chela palm (vs. strongly granulated in A. fenneri), the ventral margin of the male minor chela almost straight (vs. conspicuously convex in A. fenneri), and the distoventral margin of the penultimate article of the third maxilliped furnished with a tuft of relatively few, moderately long setae (vs. with a dense field of numerous, elongate setae in A. fenneri, see also below); and from A. sciolii by the significantly longer and slenderer penultimate and ultimate articles of the third maxilliped (cf. De Man 1909; Miya & Miyake 1969; Bruce 1994; Anker 2022). The remaining three described species of the A. djeddensis — A. djiboutensis complex, viz. A. macellarius, A. mannarensis, A. thompsoni, have similarly slender major and minor chelipeds, without balaeniceps setae or with a reduced balaeniceps condition. Nevertheless, A. karplusi sp. nov. may be morphologically separated from A. macellarius by the ventromesial margin of the chelipeds distally unarmed (vs. armed with a sharp tooth in A. macellarius) and the shorter carpocerite, the latter not reaching the distal margin of the scaphocerite blade (vs. overreaching it in A. macellarius) (cf. Chace 1988). The new species can be easily separated from both A. mannarensis and A. thompsoni by the second pereiopod carpus with the first subarticle clearly longer than the second (vs. with the second subarticle distinctly longer than the first in A. mannarensis or with the first and second subequal in length in A. thompsoni) and the penultimate article of the third maxilliped furnished with a tuft of relatively few, moderately long setae on its distoventral margin (vs. with a field of more numerous and much longer setae in A. mannarensis and A. thompsoni); individually from A. mannarensis by the mesial surface of the major and minor chelae not conspicuously granulated (vs. faintly granulated in A. mannarensis, although probably not as strongly as in A. fenneri); and individually from A. thompsoni by the presence of at least three spiniform setae on the cheliped meri (vs. their absence or presence of only one spiniform seta in A. thompsoni) (cf. Purushothaman et al. 2021; Anker 2022; Figs. 6, 7). The development of setae on the ventral margin of the penultimate article of the third maxilliped was briefly mentioned and illustrated by Banner & Banner (1981: 21, fig. 1i, j), who concluded that they represent a variable feature. However, it seems that the development of these setae is in fact species-specific and two groups can be distinguished. They are denser and extremely elongate, reaching almost to the end of the ultimate article, in some specimens identified as A. djiboutensis by Banner & Banner (1982), and also in A. bellulus, A. fenneri, A. mannarensis and A. thompsoni (Miya & Miyake 1969; Bruce 1994; Purushothaman et al. 2021; Anker 2022; Fig. 6E). In contrast, they are less somewhat numerous and only moderately elongate, i.e. barely reaching the distal half of the ultimate article, in A. djeddensis sensu Banner & Banner (1982), some specimens of A. djiboutensis sensu Banner & Banner (1982), as well as in A. macellarius, A. sciolii and A. karplusi sp. nov. (Anker 2022; A. Anker, pers. obs.; Fig. 1E). The distribution range of A. karplusi sp. nov. appears to be restricted to the Red Sea (Israel, Egypt, Saudi Arabia). All photographic records of “ A. rubromaculatus ” outside of the Red Sea most probably refer to a closely related, undescribed species, hereafter A. aff. karplusi (Anker, pers. obs.). These records include three colour photographs of “ Alpheus sp. 8 & 9” from Malaysia and Indonesia in Kuiter & Debelius (2009: 153); a blackand-white photograph of “ A. rubromaculatus ” from Malaysia (same photograph as in Kuiter & Debelius 2009) in Karplus & Thompson (2011: 591, fig. 4.4.10-C); two colour photographs of “ A. rubromaculatus ” from Malaysia (Mabul) and Indonesia (Bali) in Debelius (2001: 153); a colour photograph of “ Alpheus sp. 5 ” in Minemizu (2013: 103); and several previously unpublished colour photographs from the Philippines, Indonesia, Taiwan, Marshall Islands and Mauritius (Fig. 5). The main differences between the colour patterns of A. aff. karplusi (Fig. 5) and A. karplusi sp. nov. (Fig. 3, 4) are the clearly more yellow-green tinge of the body and appendages in the former species, including the more yellow-greenish chelipeds, bright yellow walking legs and conspicuously yellow or orange antennal flagella, as well as the spots on the carapace and pleon being more rounded and more dark red-purple in A. aff. karplusi instead of rusty-brown, as in A. karplusi sp. nov. In addition, A. aff. karplusi associates with different species of gobies and is allopatrically isolated from A. karplusi sp. nov. (see below). This widespread but as yet unnamed species, which is often incorrectly referred to as “ A. rubromaculatus ” (e.g. Debelius 2001; Pinault et al. 2015; marine aquarium trade), will be described when material becomes available for study. Both A. karplusi sp. nov. and A. aff. karplusi are typically (but not exclusively) encountered in association with the whitecap hovering gobies of the genus Lotilia. According to Shibukawa et al. (2012), Lotilia is composed of two morphologically nearly identical species, which have been confused with each other until recently. The type species of Lotilia, L. graciliosa, is restricted to the “Red Sea and adjacent areas”, whereas L. klausewitzi Shibukawa, Suzuki & Senou, 2012 occurs in the “West Pacific” (Japan, Australia, Papua New Guinea, Phoenix Islands), although based on numerous colour photographs, it is also widely dist, Published as part of Anker, Arthur, 2022, On two goby-associated snapping shrimps from the Red Sea, one of them new to science (Malacostraca: Decapoda: Alpheidae: Alpheus), pp. 421-438 in Zootaxa 5105 (3) on pages 423-431, DOI: 10.11646/zootaxa.5105.3.5, http://zenodo.org/record/6332829, {"references":["Karplus, I., Szlep, R. & Tsurnamal, M. (1981) Goby-shrimp partner specificity. I. Distribution in the northern Red Sea and partner specificity. Journal of Experimental Marine Biology and Ecology, 51, 1 - 19. https: // doi. org / 10.1016 / 0022 - 0981 (81) 90151 - 9","Karplus, I. (1987) The association between gobiid fishes and burrowing alpheid shrimps. Oceanography and Marine Biology Annual Review, 25, 507 - 562.","Debelius, H. (2001) Crustacea guide of the world. IKAN, Frankfurt, 321 pp.","Vine, P. (1996) Red Sea invertebrates. Sea Challengers, Monterrey, 224 pp.","Coutiere, H. (1897) Note sur quelques especes du genre Alpheus du Musee de Leyde. Notes from the Leyden Museum, 19, 195 - 207.","Karplus, I. (1992) Obligatory and facultative goby-shrimp partnerships in the western tropical Atlantic. Symbiosis, 12, 275 - 291.","Karplus, I. & Thompson, A. R. (2011) The partnership between gobiid fishes and burrowing alpheid shrimps. In: Patzner, R. A., Van Tassell, J. L., Kovacic, M. & Kapoor, B. G. (Eds.), The biology of gobies. Science Publishers, Enfield, New Hampshire, pp. 559 - 608.","Debelius, H. (2011) Red Sea reef guide. IKAN, Frankfurt, 322 pp.","Shibukawa, K., Suzuki, T. & Senou, H. (2012) Review of the shrimp-associated goby genus Lotilia (Actinopterygii: Perciformes: Gobiidae), with description of a new species from the West Pacific. Zootaxa, 3362 (1), 54 - 64. https: // doi. org / 10.11646 / zootaxa. 3362.1.3","Holthuis, L. B. (1958) Contributions to the knowledge of the Red Sea. No. 8. Crustacea Decapoda from the northern Red Sea (Gulf of Aqaba and Sinai Peninsula). I. Macrura. Bulletin of Sea Fisheries Research Station, Haifa, 17, l - 40.","Miya, Y. & Miyake, S. (1969) Description of Alpheus bellulus sp. nov. associated with gobies from Japan (Crustacea, Decapoda, Alpheidae). Publications of the Seto Marine Biological Laboratory, 16, 307 - 314. https: // doi. org / 10.5134 / 175556","Bruce, A. J. (1994) Alpheus fenneri sp. nov. and A. williamsi sp. nov., two new Indo-West Pacific alpheid shrimps of the brevirostris species group. The Beagle, Records of the Museums and Art Galleries of the Northern Territory, 11, 15 - 28. https: // doi. org / 10.5962 / p. 264023","Kuiter, R. H. & Debelius, H. (2009) World atlas of marine fauna. IKAN, Frankfurt, 725 pp.","Minemizu, R. (2013) Coral reef shrimps of Indo-West Pacific. Bun-ichi Sogo Shuppan, Co. Ltd., Tokyo, 144 pp. [in Japanese]","Anker, A., Pratama, I. S., Firdaus, M. & Rahayu, D. L. (2015) On some interesting marine decapod crustaceans (Alpheidae, Laomediidae, Strahlaxiidae) from Lombok, Indonesia. Zootaxa, 3911 (3), 301 - 342. https: // doi. org / 10.11646 / zootaxa. 3911.3.1","Anker, A. (2022) Description of two new species of goby-associated snapping shrimps from the tropical western Pacific (Decapoda: Alpheidae: Alpheus). Zootaxa, 5092 (3), 273 - 290. https: // doi. org / 10.11646 / zootaxa. 5092.3.2","De Man, J. G. (1909) Diagnoses of new species of macrurous decapod Crustacea from the \" Siboga-Expedition \". Tijdschrift der Nederlandse Dierkundige Vereeniging, 2, 99 - 125.","Anker, A. & De Grave, S. (2016) An updated and annotated checklist of marine and brackish caridean shrimps of Singapore (Crustacea, Decapoda). Raffles Bulletin of Zoology, 34, 343 - 454.","Coutiere, H. (1899) Les \" Alpheidae \", morphologie externe et interne, formes larvaires, bionomie. Annales des Sciences Naturelles, Zoologie, 8, 91 - 560. Masson, Paris. https: // doi. org / 10.5962 / bhl. title. 13143","Chace, F. A. Jr. (1988) The caridean shrimps (Crustacea: Decapoda) of the Albatross Philippine Expedition, 1907 - 1910, Part 5: Family Alpheidae. Smithsonian Contributions to Zoology, 466, 1 - 99. https: // doi. org / 10.5479 / si. 19436696.391.1","Purushothaman, P., Abhilash, C. P., Ajith Kumar, T. T. & Lal, K. K. (2021) A new alpheid shrimp, Alpheus mannarensis sp. nov. (Crustacea; Decapoda; Alpheidae) from Gulf of Mannar, southern India. Zootaxa, 5026, 127 - 135. https: // doi. org / 10.11646 / zootaxa. 5026.1.5","Banner, D. M. & Banner, A. H. (1981) Annotated checklist of the alpheid shrimp of the Red Sea and Gulf of Aden. Zoologische Verhandelingen, Leiden, 190, l - 99.","Banner, D. M. & Banner, A. H. (1982) The alpheid shrimp of Australia, part III: the remaining alpheids, principally the genus Alpheus, and the family Ogyrididae. Records of the Australian Museum, 34, 1 - 357. https: // doi. org / 10.3853 / j. 0067 - 1975.34.1982.434","Pinault, M., Fricke, R., Wickel, J., Peneau, C. & Quod, J. P. (2015) First record of the chocolate shrimp-goby (Gobiidae: Cryptocentrus malindiensis) from Reunion Island with a brief description of its natural habitat. Cybium, 39, 237 - 239."]}

Alpheus sciolii sp. nov. (Figs. 4���7, 8C) Alpheus cf. bellulus ��� Marin 2008: 382, fig. 9b (as A. bellulus in the figure legend) [not A. bellulus Miya & Miyake, 1969]. Alpheus sp. aff. bellulus ��� Anker & De Grave 2019: 148, fig. 4C. Alpheus aff. djiboutensis ��� Hurt et al. 2021: 2, fig. 1D. Alpheus sp. ��� Debelius 2001: 153, colour photograph. Alpheus sp. 2 ��� Minemizu 2013: 101, colour photograph. Alpheus sp. 6 ��� Kuiter & Debelius 2009: 152, 4 colour photographs. Type material. Holotype: male (cl 8.6 mm), MNHN-IU-2018-5667, Solomon Islands, New Georgia, Munda, Hopei Island, shallow sand flat, suction (yabby) pump, depth 0.5���1 m, leg. A. Anker, 14.09.2016 [fcn SOL-106]. Paratypes: 1 male (cl 9.3 mm), MNHN-IU-2018-5669, Solomon Islands, New Georgia, Munda, Sosuhite Island, shallow sand flat with coral rubble, depth 0.5���2 m, in burrow under large piece of coral rubble, leg. A. Anker, 19.09.2016 [fcn SOL-047]; 1 ov. female (cl 10.7 mm), MNHN-IU-2018-5668, Australia, Queensland, Heron Island, southern side, shallow reef flat, coarse sand, suction (yabby) pump, 0.5���1 m, leg. A. Anker, 28.09.2016 [fcn HE-011]; 1 male (cl 6.6 mm), 1 female (cl 8.4 mm), MNHN-IU-2018-5670, Australia, Queensland, Heron Island, southern side, shallow reef flat, coarse sand, suction (yabby) pump, 0���0.5 m, leg. A. Anker, 24.09.2016 [fcn HE-120]; 1 male (cl 16.0 mm, missing both chelipeds), MNHN-IU-2018-5525, Australia, Queensland, Heron Island, southern side, shallow reef flat, coarse sand, suction (yabby) pump, 0���0.5 m, leg. A. Anker, 25.09.2016 [fcn HE-026]. Additional material. 1 male (cl 14.5 mm), MNHN-IU-2018-5671, Vietnam, Nha Trang Bay, Tre Island, Tre Bay, intertidal area near mangrove, suction (yabby) pump, with goby, leg. I.N. Marin, 12.07.2006. Description. Carapace not setose, not pubescent, pitted (Fig. 4A, B). Rostrum well developed, wide, about 1.7 times as long as wide at base, subacute distally, just reaching distal third of first article of antennular peduncle, pointing straight-forward in lateral view; rostral carina well marked, rounded dorsally, gently sloping into welldemarcated adrostral furrows, continuing well beyond base of orbital hoods, gradually flattening, not reaching midlength of carapace (Fig. 4A, B). Orbital hoods swollen, feebly projecting anteriorly in lateral view, unarmed; frontal margin between rostrum and orbital hood shallowly concave (Fig. 4A, B). Pterygostomial angle rounded; cardiac notch well developed, deep (Figs. 4B, 7B). Telson very broad, subrectangular, gently tapering distally, about 1.8 times as long as maximal width, with lateral margins not noticeably constricted; dorsal surface with two pairs of stout cuspidate setae both inserted far from lateral margin, first pair at about 0.5 telson length, second pair at about 0.7 of telson length; posterior margin broadly rounded, with row of short spiniform setae above plumose setae (less conspicuous or with slender spiniform setae in some specimens); posterolateral angles each with one pair of spiniform setae (typically, sometimes with only one spiniform seta), mesial ones stouter and two to three times as long as lateral ones [posterolateral spiniform setae somewhat aberrant in the illustrated paratype (Fig. 4C), see Fig. 6C for more typical condition]. Eyes well developed, with large, normally pigmented corneas (Figs. 1A, B, 7). Antennular peduncle moderately elongate and stout; stylocerite feebly swollen laterally, distal part slenderer, ending in sharp point, latter not reaching distal margin of first article; ventromesial carina with large subtriangular tooth; second article about 2.7 times as long as wide; lateral antennular flagellum with secondary ramus fused to main ramus over most of its length, with numerous groups of aesthetascs distally, starting from about 14th subdivision (Fig. 4A, B, D). Antenna with basicerite fairly stout, its distoventral margin armed with large, anteriorly projecting, sharp tooth; scaphocerite with moderately broad blade distinctly exceeded by sharp distolateral tooth, lateral margin shallowly concave; scaphocerite reaching distinctly beyond end of antennular peduncle and slightly beyond carpocerite (Fig. 4A, B). Mouthparts not dissected, generally typical for genus in external observation. Third maxilliped relatively stout; coxa with bluntly projecting lateral plate; antepenultimate article flattened ventrolaterally, with distinct ridge running parallel to dorsal margin on lateral surface and rugose mesial margin, about 3.9 times as long as high; penultimate article very short, cup-shaped, bulging ventrally, its distoventral margin with few moderately long setae, its dorsal margin with row of elongate setae; ultimate article densely furnished with rows of very long, stiff setae, dorsal and apical setae longest; arthrobranch well developed (Fig. 4E). Male major cheliped robust; ischium short, stout, smooth; merus very stout, trigonal in cross-section, about twice as long as maximal width, distodorsal margin ending bluntly, ventromesial margin somewhat rugose, with more or less pronounced, blunt, distal tooth and row of five or six spiniform setae; carpus short, distally widening, cup-shaped; chela not particularly elongate, fringed with long setae along dorsal and ventral surfaces; palm strongly compressed, stout, subrectangular in cross-section, length / maximal height ratio 1.8���2.1 (lower in larger males); surfaces relatively smooth, without granules; dorsal margin with well-marked transverse groove subdistally, without marked longitudinal ridges; ventral margin almost straight; lateral surface with shallow depression distally, near base of pollex; fingers subequal in length, about half-length of palm, not twisted or significantly deviating from chela axis; dactylus distally rounded, plunger somewhat reduced, poorly demarcated from anterior cutting edge, latter slightly bulging; adhesive disks very small (Fig. 5A���D; see also Fig. 6A). Female major cheliped generally similar to male major cheliped, smaller, weaker; chela with somewhat different proportions (see Fig. 7C). Male minor cheliped with ischium short, smooth; merus similar to that of major cheliped in size and stoutness, trigonal in cross-section, distodorsal angle blunt, ventromesial margin slightly rugose distally, usually without distal tooth, armed with spiniform setae; carpus distinctly longer than that of major cheliped, distally widening, cupshaped; chela not elongate, not swollen, fringed with long setae along dorsal and ventral surfaces; palm compressed, subrectangular in cross-section, length / maximal height ratio 1.3���1.5 (lower in larger males); surfaces smooth, not granulated; dorsal margin without groove; ventral margin broadly convex; fingers subequal in length, about 1.3 times length of palm, not twisted, slightly gaping when closed, with balaeniceps crests lined with setae (better developed in larger males); adhesive disks greatly reduced (Fig. 5E, F; see also Fig. 6B). Female minor cheliped generally similar, but with weaker chela, latter with balaeniceps ridges and setae on fingers poorly developed or lacking (Fig. 7C). Second pereiopod slender; ischium and merus subequal in length; carpus with five subarticles, first noticeably longer than second, ratio of carpal subarticles approximately equal to 3.0/2.5/1.0/1.0/1.7; chela longer than distalmost carpal subarticle (Fig. 4F). Third pereiopod fairly robust; ischium with stout spiniform seta on ventrolateral surface; merus about 4.1 times as long as maximal width, ventromesial margin unarmed distally; carpus about 0.6 length of merus, much more slender than merus, unarmed; propodus subequal to carpus, setose, with at least 10 spiniform setae along ventral margin, including distal pair adjacent to dactylar base; dactylus slightly more than halflength of propodus, gradually curving distally, subspatulate, moderately broadened, flattened ventrally, with distinct longitudinal keel separating ventral and dorsal surfaces, with scarce setae (Fig. 4G). Fourth pereiopod generally similar to third pereiopod, more slender. Fifth pereiopod much more slender than third and fourth pereiopods; ischium with small spiniform seta on ventrolateral surface; merus about 5.7 times as long as wide; carpus slightly more slender than merus, about 0.9 length of merus; propodus subequal to carpus, with rows of serrulate setae forming cleaning brush on distal third of ventrolateral surface, and at least eight spiniform setae along ventromesial margin, including distal pair adjacent to dactylar base (Fig. 4H, I). Second pleopod with appendices masculina and interna equal in length, former with long stiff setae (Fig. 4J). Uropod with mesial and lateral lobes of protopod each ending in sharp (lateral) or blunt (mesial) distal tooth; exopod broad, broadly rounded distally, with short triangular distolateral tooth; diaeresis straight for most part, except for rounded lobe adjacent to slender spiniform seta, latter not reaching level of distal margin of exopod; endopod much narrower than exopod, with row of spiniform setae on distal margin (Fig. 4K). Colour pattern. General background colour white with pale yellowish tinge; carapace with several (three to four) broad bands, running transversely dorsally and more obliquely laterally, some broadly croissant-shaped; carapace flanks with very large brown to almost black spot (quickly fading in captured individuals); pleon with narrower, transversely-obliquely running, brown bands, usually two per pleonite; posterior end of carapace and first pleonite with conspicuous, white, dorsal ���saddle���; fourth and fifth pleonites with transverse, whitish, dorsal patches; chelipeds whitish or greyish, with irregular, brown, green-brown or green-bluish patches; distal half of fingers hyaline-white; antennules and antennae whitish with brownish spots and streaks; antennular flagella pale yellow proximally, more intense yellow distally; antennal flagella pale yellow proximally, uncoloured distally; second pereiopods and walking legs greyish-blue with yellow markings around articulations; pleopods brownish yellow; uropods whitish with some brown-red bands or spots, mainly on endopod; telson yellow whitish proximally, with brown-red pattern distally (Figs. 7, 8C; see also Anker & De Grave 2019: fig. 4C). Etymology. This new species is named after Justin Scioli, PhD cand., the author���s great friend and collaborator in projects involving taxonomy and phylogenetics of alpheid shrimps. Distribution. Western Pacific: confirmed records from Solomon Islands, Australia (Great Barrier Reef) and Vietnam (present study); photographic records from the Philippines (Palawan), Indonesia (Java, Flores, Bali), Papua New Guinea and Palau (Debelius 2001; Randall et al. 2007; Kuiter & Debelius 2009; Hurt et al. 2021). Ecology. Shallow coral reefs and adjacent reef flats and lagoons; associated with gobies, typically Cryptocentrus cinctus (Herre), C. fasciatus (Playfair), C. leptocephalus Bleeker and C. inexplicatus (Herre), but also Ctenogobiops pomastictus (Fig. 8C; see also Debelius 2001; Randall et al. 2007; Kuiter & Debelius 2009; Hurt et al. 2021); may also harbour the ���commensal��� palaemonid shrimp, Palaemonella aliska Marin, 2008 (Marin 2008; Anker & De Grave 2019). Remarks. Alpheus sciolii sp. nov. has a unique and unmistakable colour pattern (Figs. 7, 8C), characterised by the dark-brown or olive-brown transverse banding and the presence of a very large dark spot on the flanks of the carapace. As in the previous species, the white ���saddle��� marking the limit between the carapace and the pleon is quite conspicuous. This colour pattern (Fig. 7; see also Kuiter & Debelius 2009, as Alpheus sp. 6, and Minemizu 2013, as Alpheus sp. 2) is very different from the colour patterns of A. bellulus, A. macellarius, A. fenneri, A. mannarensis (see Miya & Miyake 1969; Debelius 2001; Kuiter & Debelius 2009; Minemizu 2013; Anker et al. 2015; Purushothaman et al. 2021; for A. bellulus see also Fig. 8F) and A. thompsoni sp. nov. (Figs. 3, 8A, B). The non-type male specimen of A. sciolii sp. nov. from Nha Trang, Vietnam, has noticeably larger and stouter chelipeds, with a markedly stronger balaeniceps condition on the minor chela fingers, compared to that of the complete male type specimens from the Solomon Islands and Australia, (Fig. 6A, B), which may be simply due to its larger size (cl 14.5 mm vs. 6.6���9.3 mm). A large paratype male of similar size (cl 16.0 mm) from Heron Island is unfortunately lacking both of its chelipeds (see Anker & De Grave 2019: fig. 4C). The Nha Trang specimen also seems to have a darker colour (see Marin 2008: fig. 9b), although it is partly covered by detritus and sediment. In addition, a colour pattern very similar to that of A. sciolii exists in shrimps associated with different gobies, such as Amblyeleotris steinitzi (Klausewitz) (see Fig. 8D). Therefore, collection of additional material and a molecular comparison between the specimens of A. sciolii sp. nov. from Solomon Islands / Australia and Vietnam (or other parts of South-East Asia) are highly desirable. The separation of A. sciolii sp. nov. from A. djeddensis is based on the same criteria as those used by De Man (1909) to distinguish A. djiboutensis from A. djeddensis, viz. the scaphocerite blade reaching well beyond the adjacent distolateral tooth in both A. sciolii sp. nov. and A. djiboutensis (vs. not reaching beyond it in A. djeddensis), and the first subarticle of the second pereiopod carpus noticeably longer than the second in A. sciolii sp. nov. and A. djiboutensis (vs. as long as the second in A. djeddensis). On the other hand, A. sciolii sp. nov. may be separated from A. djiboutensis by the much broader telson, which is around 1.8 times as long as maximal width in A. sciolii sp. nov. (Figs. 4C, 6C) vs. 2.1 times in A. djiboutensis (see De Man 1909: fig. 18). In fact, the telson of A. sciolii sp. nov. appears to be the broadest among the presently known species of the A. djeddensis ��� A. djiboutensis complex. Also noteworthy is that no snapping shrimp with the colour pattern of A. sciolii sp. nov. is known from the Red Sea (Karplus et al. 1981) or western Indian Ocean (Polunin & Lubbock 1977), from where A. djeddensis and A. djiboutensis were origially described. The Australian material reported as A. djiboutensis by Banner & Banner (1982) appears to contain more than one species (possibly including the true A. djiboutensis and A. sciolii sp. nov.) and will need to be carefully re-examined. Alpheus sciolii sp. nov. can be distinguished morphologically from A. bellulus by the rostrum much narrower and longer (vs. shorter and subtriangular in A. bellulus), the ventral margin of the major chela almost straight to slightly convex (vs. somewhat concave in A. bellulus), and the telson 1.8 times as long as broad (vs. 2.5 times as long as broad) (see Miya & Miyake 1969: figs. 1, 2B, C); from A. macellarius by the distolateral tooth of the antennal scaphocerite pointing straight-forward (vs. curved mesially in A. macellarius), the much stouter minor chela (which is very slender in A. macellarius), and the much stouter third pereiopod merus, about four times as long as wide (vs. almost six times in A. macellarius) (see Chace 1988: fig. 6e, f, i); from A. fenneri by the distolateral tooth of the scaphocerite reaching far beyond the blade (vs. not or very slightly overreaching the blade in A. fenneri), the mesial surface of the major and minor chelae not granulated (vs. distinctly granulated in A. fenneri), and the posterior margin of the telson unarmed between the mesial pair of the posterolateral spiniform setae (vs. with a row of spiniform setae in A. fenneri) (see Bruce 1994: figs. 1C, M, N, 2C); from A. mannarensis by the distolateral tooth of the scaphocerite reaching far beyond the blade (vs. not reaching the anterior margin of the blade in A. mannarensis), the second pereiopod carpus with the first subarticle slightly longer than the second (vs. with the second subarticle noticeably longer than the first in A. mannarensis), the posterior margin of the telson unarmed between the mesial pair of the posterolateral spiniform setae (vs. with a row of spiniform setae in A. mannarensis) (see Purushothaman et al. 2021: figs. 2B, E, 3G); and from the above-described A. thompsoni sp. nov. by the relative development of the distolateral tooth of the scaphocerite (cf. Figs. 1A, 4A), the length-width proportions of the telson (cf. Figs. 1C, 4C), the much stouter third maxilliped, without extremely elongate setae on the pendultimate article (cf. Figs. 1E, 4E), and the much stouter walking legs (cf. Figs. 1G, I, 4G, H)., Published as part of Anker, Arthur, 2022, Description of two new species of goby-associated snapping shrimps from the tropical western Pacific (Decapoda: Alpheidae: Alpheus), pp. 273-290 in Zootaxa 5092 (3) on pages 281-288, DOI: 10.11646/zootaxa.5092.3.2, http://zenodo.org/record/5881095, {"references":["Marin, I. (2008) Description of two new species from the genera Palaemonella Dana, 1852 and Vir Holthuis, 1952 (Crustacea: Caridea: Palaemonidae: Pontoniinae). Zoologische Mededelingen, 82, 375 - 390.","Miya, Y. & Miyake, S. (1969) Description of Alpheus bellulus sp. nov. associated with gobies from Japan (Crustacea, Decapoda, Alpheidae). Publications of the Seto Marine Biological Laboratory, 16, 307 - 314. https: // doi. org / 10.5134 / 175556","Anker, A. & De Grave, S. (2019) Further records of burrow-associated palaemonid shrimps (Decapoda: Palaemonidae). Zootaxa, 4612 (1), 145 - 150. https: // doi. org / 10.11646 / zootaxa. 4612.1.13","Hurt, C., Hultgren, K. M., Anker, A., Lemmon, A. R., Moriarty Lemmon, E. & Bracken-Grissom, H. (2021) First worldwide molecular phylogeny of the morphologically and ecologically hyperdiversified snapping shrimp genus Alpheus (Malacostraca: Decapoda). Molecular Phylogenetics and Evolution, 158 (197080), 1 - 11. https: // doi. org / 10.1016 / j. ympev. 2021.107080","Debelius, H. (2001) Crustacea guide of the world. IKAN, Frankfurt, 321 pp.","Minemizu, R. (2013) Coral reef shrimps of Indo-West Pacific. Bun-ichi Sogo Shuppan, Co. Ltd., Tokyo, 144 pp. [in Japanese]","Kuiter, R. H. & Debelius, H. (2009) World atlas of marine fauna. IKAN, Frankfurt, 725 pp.","Randall, J. E., Shao, K. T. & Chen, J. P. (2007) Two new shrimp gobies of the genus Ctenogobiops (Perciformes: Gobiidae), from the Western Pacific. Zoological Studies, 46, 26 - 34.","Anker, A., Pratama, I. S., Firdaus, M. & Rahayu, D. L. (2015) On some interesting marine decapod crustaceans (Alpheidae, Laomediidae, Strahlaxiidae) from Lombok, Indonesia. Zootaxa, 3911, 301 - 342. https: // doi. org / 10.11646 / zootaxa. 3911.3.1","Purushothaman, P., Abhilash, C. P., Ajith Kumar, T. T. & Lal, K. K. (2021) A new alpheid shrimp, Alpheus mannarensis sp. nov. (Crustacea; Decapoda; Alpheidae) from Gulf of Mannar, southern India. Zootaxa, 5026 (1), 127 - 135. https: // doi. org / 10.11646 / zootaxa. 5026.1.5","De Man, J. G. (1909) Diagnoses of new species of macrurous decapod Crustacea from the \" Siboga-Expedition \". Tijdschrift der Nederlandse Dierkundige Vereeniging, 2, 99 - 125.","Karplus, I., Szlep, R. & Tsurnamal, M. (1981) Goby-shrimp partner specificity. I. Distribution in the northern Red Sea and partner specificity. Journal of Experimental Marine Biology and Ecology, 51, 1 - 19. https: // doi. org / 10.1016 / 0022 - 0981 (81) 90151 - 9","Polunin, N. V. C. & Lubbock, R. (1977) Prawn-associated gobies (Teleostei: Gobiidae) from the Seychelles, western Indian Ocean: systematics and ecology. Journal of Zoology, London, 183, 63 - 101. https: // doi. org / 10.1111 / j. 1469 - 7998.1977. tb 04174. x","Banner, D. M. & Banner, A. H. (1982) The alpheid shrimp of Australia, part III: the remaining alpheids, principally the genus Alpheus, and the family Ogyrididae. Records of the Australian Museum, 34, 1 - 357. https: // doi. org / 10.3853 / j. 0067 - 1975.34.1982.434","Chace, F. A. Jr. (1988) The caridean shrimps (Crustacea: Decapoda) of the Albatross Philippine Expedition, 1907 - 1910, Part 5: Family Alpheidae. Smithsonian Contributions to Zoology, 466, 1 - 99. https: // doi. org / 10.5479 / si. 19436696.391.1","Bruce, A. J. (1994) Alpheus fenneri sp. nov. and A. williamsi sp. nov., two new Indo-West Pacific alpheid shrimps of the brevirostris species group. The Beagle, Records of the Museums and Art Galleries of the Northern Territory, 11, 15 - 28. https: // doi. org / 10.5962 / p. 264023"]}

Alpheus thompsoni sp. nov. (Figs. 1���3, 8A, B) (?) Alpheus ochrostriatus ��� Karplus et al. 1981: 6, fig. 2F (black-and-white photograph); Debelius 2001: 150 (part.?), 2 colour photographs; Randall et al. 2003: 512, fig. 8 (colour photograph); Poupin 2010: 33; Karplus & Thompson 2011: 591, fig. 4.4.10-B (black-and-white photograph) [nomen nudum]. (?) Alpheus ��� ochrostriatus ��� ��� Anker 2000: 3, fig. 2; Jaafar & Randall 2009: 29, pl. 2A (colour photograph); Kuiter & Debelius 2009: 151, 3 colour photographs (as Alpheus sp. ��� ochrostriata ���) [nomen nudum]. (?) Alpheus sp. 4 ��� Minemizu 2013: 102, colour photograph. Type material. Holotype: ov. female (cl 13.3 mm), MNHN-IU-2013-16813, Papua New Guinea, Madang, sta. PR242, 5��10.2���S, 145��50.4���E, depth 4���24 m, leg. A. Anker et al., 12.12.2012 [fcn PR242-PZD-660D]. Description. Carapace sparsely setose, not pubescent, pitted (Fig. 1A, B). Rostrum well developed, slender, about twice as long as wide at base, subacute distally, reaching distal third of first article of antennular peduncle, somewhat descendent in lateral view; rostral carina well marked, rounded dorsally, gently sloping into shallow adrostral furrows, continuing well beyond base of orbital hoods, gradually flattening, not reaching mid-length of carapace (Fig. 1A, B). Orbital hoods swollen, somewhat projecting anteriorly in lateral view, unarmed; frontal margin between rostrum and orbital hood shallowly concave (Fig. 1A, B). Pterygostomial angle rounded; cardiac notch well developed, deep (Figs. 1B, 3B). Telson moderately broad, subrectangular, gently tapering distally, about 2.5 times as long as maximal width, with lateral margins somewhat constricted and tapering at about 0.7 times telson length; dorsal surface with two pairs of stout cuspidate setae both inserted far from lateral margin, first pair at about 0.4 telson length, second pair at about 0.7 times telson length; posterior margin broadly rounded, with row of slender spiniform setae above plumose setae; posterolateral angles each with one pair of spiniform setae, mesial ones stouter and almost three times as long as lateral ones (Fig. 1C). Eyes well developed, with large, normally pigmented corneas (Figs. 1A, B, 3). Antennular peduncle moderately elongate and stout; stylocerite slightly swollen laterally, distal part more slender, ending in sharp point, latter slightly overreaching distal margin of first article; ventromesial carina with large, subacute tooth; second article about 2.7 times as long as wide; lateral antennular flagellum with secondary ramus fused to main ramus over most of its length, with numerous groups of aesthetascs distally, starting from about 13th subdivision (Fig. 1A, B, D). Antenna with basicerite fairly stout, its distoventral margin armed with large, anteriorly projecting, sharp tooth; scaphocerite with broad blade reaching slightly beyond stout, subacute distolateral tooth, lateral margin very shallowly concave; scaphocerite reaching slightly beyond end of antennular peduncle and far beyond carpocerite (Fig. 1A, B). Mouthparts not dissected, generally typical for genus in external observation. Third maxilliped relatively slender, elongate; coxa with bluntly projecting lateral plate; antepenultimate article flattened ventrolaterally, with distinct ridge running parallel to dorsal margin on lateral surface and rugose mesial margin, about 5.3 times as long as high; penultimate article short, cup-shaped, bulging ventrally, its distoventral margin with tuft of very long setae; ultimate article densely furnished with rows of very long, stiff setae, apical setae longest; arthrobranch well developed (Fig. 1E). Major cheliped (female) moderately robust; ischium short, stout, smooth; merus moderately stout, somewhat elongate, trigonal in cross-section, at most four times as long as maximal width, distodorsal margin ending bluntly, ventromesial margin somewhat rugose, with weak, blunt, distal tooth, without spiniform setae; carpus short, distally widening, cup-shaped; chela not particularly elongate, fringed with long setae along dorsal and ventral surfaces; palm strongly compressed, subrectangular in cross-section, length / maximal height ratio about 1.9; surfaces relatively smooth, not markedly granulated; dorsal margin with well-marked transverse groove subdistally, without marked longitudinal ridges; ventral margin almost straight; lateral surface with distinct depression distally, near base of pollex; fingers slightly unequal in length, with dactylus shorter than pollex, about half-length of palm, not twisted or significantly deviating from chela axis; dactylus distally rounded, with plunger greatly reduced, poorly demarcated from anterior cutting edge, latter feebly bulging; adhesive disks very small (Fig. 2A���D). Minor cheliped (female) with ischium short, smooth; merus slightly stouter than that of major cheliped, trigonal in cross-section, about 3.5 times as long as maximal width, distodorsal angle blunt, ventromesial margin slightly rugose distally, without pronounced distal tooth, armed with one spiniform seta; carpus somewhat more slender and longer than that of major cheliped, distally widening, cup-shaped; chela not elongate, not swollen, fringed with long setae along dorsal and ventral surfaces; palm compressed, subrectangular in cross-section, length / maximal height ratio about 2.0; surfaces smooth, not granulated; dorsal margin without groove; ventral margin straight; fingers slightly unequal in length, with dactylus shorter than pollex, about 1.2 times length of palm, not twisted, slightly gaping when closed, without balaeniceps crests and setae; adhesive disks greatly reduced (Fig. 2E���G). Second pereiopod slender; ischium and merus subequal in length; carpus with five subarticles, first and second subequal, ratio of carpal subarticles approximately equal to 2.3/2.2/1.0/0.9/1.5; chela longer than distal-most carpal subarticle (Fig. 1F). Third pereiopod moderately stout; ischium with small spiniform seta on ventrolateral surface; merus about 4.5 times as long as maximal width, ventromesial margin unarmed distally; carpus about half-length of merus, much more slender than merus, unarmed; propodus noticeably longer than carpus, moderately setose, with seven or so spiniform setae along ventral margin, including distal pair adjacent to dactylar base; dactylus about half-length of propodus, gradually curving distally, subspatulate, moderately broadened, flattened ventrally, with distinct longitudinal keel separating ventral and dorsal surfaces, latter with several tufts of setae (Fig. 1G, H). Fourth pereiopod generally similar to third pereiopod, somewhat slenderer. Fifth pereiopod much slenderer than third and fourth pereiopods; ischium with small spiniform seta on ventrolateral surface; merus about nine times as long as wide; carpus slightly more slender than merus, about 0.9 length of merus; propodus subequal to carpus, with rows of serrulate setae forming cleaning brush on distal third of ventrolateral surface, and about 10 spiniform setae along ventromesial margin, including distal pair adjacent to dactylar base (Fig. 1I, J). Uropod with mesial and lateral lobes of protopod each ending in sharp distal tooth; exopod broad, somewhat truncate distally, with short triangular distolateral tooth; diaeresis straight for most part, except for blunt lobe adjacent to slender spiniform seta, latter not reaching level of distal margin of exopod; endopod much narrower than exopod, with row of spiniform setae on distal margin (Fig. 1K). Colour pattern. General background colour pale pink; carapace and pleon with broad, parallel-running, longitudinal bands each composed of narrower, ochre-red to brick-red lines and streaks, many anastomosing, forming net-like pattern; carapace flanks with broader, more diffuse and more obliquely running bands; posterior end of carapace and first pleonite with very conspicuous, transverse, white, dorsal ���saddle���; fourth and fifth pleonites with smaller dorsal ���saddle���; fifth pleonite with small, whitish, dorsal patch; chelipeds with yellow tinge, especially on merus and carpus, and ochre-red stripes and patches on palms and fingers; distal half of fingers hyaline-white with pinkish tinge; antennules and antennae bright yellow with ochre-orange spots and streaks, especially on antennular peduncles and scaphocerite; antennular and antennal flagella almost colourless, with yellowish or bluish tinge, respectively; second pereiopods and walking legs uniform yellow; pleopods yellowish; uropods mainly yellow with whitish areas and few ochre-orange spots on endopod; telson yellow with two ochre-red longitudinal bands flanking white mid-dorsal band (Fig. 3; see also Fig. 8A, B). Etymology. This new species is named after the author���s friend and colleague, Dr. Andrew Thompson, an expert of ecology and evolution of goby-shrimp symbioses (Thompson 2004, 2005; Karplus & Thompson 2011). Distribution. Indo-West Pacific: so far confirmed only from Papua New Guinea (Madang); numerous photographic records (as ��� A. ochrostriatus ��� or Alpheus sp.) ranging from the Red Sea (Karplus et al. 1981) to Indonesia (Randall et al. 2007; Kuiter & Debelius 2009) and Japan (Anker 2000; Karplus & Thompson 2011); also recorded, possibly based on colour photographs, from New Caledonia (Poupin 2010, as ��� A. ochrostriatus ���); however, all records from outside of the type locality require confirmation as more than one species might be involved (see below). Ecology. Coral reefs and adjacent reef flats; associated with numerous gobies, including Amblyeleotris wheeleri (Polunin & Lubbock), and possibly also A. steinitzi (Klausewitz), A. guttata (Fowler), Ctenogobiops pomastictus Polunin & Lubbock, and C. tangaroai Lubbock & Polunin (Fig. 8A, B; see also Karplus et al. 1981; Anker 2000; Debelius 2001; Randall et al. 2003, 2007; Kuiter & Debelius 2009; Jaafar & Randall 2009; Karplus & Thompson 2011; however, see discussion below). Remarks. Alpheus thompsoni sp. nov. has a very striking colour pattern (Figs. 3, 8A, B), which is characterised by the long, narrow, longitudinal bands of ochre-red to brick-red colour on a generally pale-yellow body and chelipeds, sharply contrasting with the bright yellow colour of the remaining appendages (antennules, antennae, walking legs and uropods). The white ���saddle��� between the carapace and pleon, which is also present in several other goby-associated species (e.g. Karplus et al. 1981; Kuiter & Debelius 2009; Minemizu 2013) is very conspicuous. This colour pattern separates the new species from A. bellulus (see Miya & Miyake 1969; Debelius 2001; Kuiter & Debelius 2009; Minemizu 2013; see also Fig. 8F), A. macellarius (see Anker et al. 2015; Anker & De Grave 2016), A. fenneri (see Bruce 1994; Kuiter & Debelius 2009), and A. mannarensis (see Purushothaman et al. 2021), as well as the strikingly coloured species reported as A. djiboutensis by Holthuis (1958), A. djeddensis s. lat. by Anker & De Grave (2016) and A. djeddensis by Dehghani et al. (2019). The above described colour pattern of A. thompsoni sp. nov. is identical or at least very similar to that of many snapping shrimps associated with gobies across the Indo-West Pacific. This species was sometimes identified as ��� A. ochrostriatus ���, a nomen nudum first used by Karplus et al. (1981), or as ��� Alpheus sp. ��� Upon the author���s request, Dr. Ilan Karplus provided scans of two colour slides of a snapping shrimp identified as ��� A. ochrostriatus ��� from Eilat, Israel, which were taken during his ecological studies in the 1980s. Although the photographed specimen was not deposited (or at least is untraceable), the shrimp on these slides is indeed similar to the holotype of A. thompsoni sp. nov. both in general morphology and colour pattern. This fact combined with numerous in situ colour photographs (e.g. Randall et al. 2003; Kuiter & Debelius 2009; Jaafar & Randall 2009; Karplus & Thompson 2011; Minemizu 2013) suggest that A. thompsoni sp. nov. may be distributed from the northern Red Sea to southern Japan and New Caledonia. However, the taxonomic status of the Red Sea populations originally reported as ��� A. ochrostriatus ��� (Karplus et al. 1981) will need to be confirmed morphologically and genetically, in view of high level of endemism of the area and differences in some colour pattern details (A. Anker, pers. obs.). Therefore, A. thompsoni sp. nov. remains known with certainty only from the type locality in Papua New Guinea, until more material is collected and analysed morphologically and genetically. More generally, the name ��� A. ochrostriatus ��� should not be used in the future, even if the Red Sea material is confirmed to be a distinct, undescribed species, since (1) it represents a nomen nudum (De Grave & Fransen 2011) and (2) was apparently used for at least two colour morphs (e.g. Debelius 2001; Randall 2007; see also discussion in Anker 2000). This is also true for the two other unavailable (and therefore invalid) names in Karplus et al. (1981). The second colour morph of ��� A. ochrostriatus ��� (shown in Fig. 8E) could represent the recently described A. mannarensis, despite some differences in the shape and of the reddish lines on the pleon and the much less conspicuous white ���saddles��� in A. mannarensis. Using the key to the A. brevirostris group in Bruce (1994), A. thompsoni sp. nov. would key out to A. djeddensis, a species described based on three male and two female specimens from Djeddah, Saudi Arabia (Couti��re 1897). However, several elements of the morphology of A. djeddensis mentioned and/or illustrated by Couti��re (1897, 1899) and De Man (1909) separate A. thompsoni sp. nov. from A. djeddensis. In the new species, the scaphocerite blade exceeds the adjacent distolateral tooth (Fig. 1A), whereas in A. djeddensis, the blade has the same length as the distolateral tooth (De Man 1909: fig. 26). In A. thompsoni sp. nov., the fingers of the female minor chela are not balaeniceps, whereas the dorsal surface of the palm is smooth, without discernable grooves. In contrast, in A. djeddensis, the minor chela fingers are strongly balaeniceps in both sexes, whereas the palm has a longitudinal groove departing from a distinct transverse groove and running parallel to the palm���s dorsal margin, as seen in the very detailed illustration of the minor chela provided by Couti��re���s (1899: fig. 232, as A. rapax var. Djeddensis). Both characters were verified in the male and female syntypes of A. djeddensis (RMNH.CRUS.D.1781), although the palmar grooves seem to be less developed in the female minor chela. However, some other distinguishing features were found during the examination of the type material of A. djeddensis. The stylocerite of A. thompsoni sp. nov. appears to be somewhat longer and less swollen than in A. djeddensis (reaching or slightly overreaching the distal margin of the antennular peduncle in the new species vs. not reaching it in A. djeddensis), whereas the antennal carpocerite is shorter (not reaching the distal margin of the scaphocerite in the new species vs. overreaching it in A. djeddensis). Their major chelae are also somewhat different, for instance, in the proportion of the fingers to the palm (0.4 in A. thompsoni sp. nov. vs. 0.6 in A. djeddensis), and in the cutting edge of the dactylus (with a less developed plunger and feebly bulging in A. thompsoni sp. nov. vs. with a more pronounced plunger and with a strong anterior bulge in A. djeddensis). It must be noted, however, that in the A. brevirostris group, the major and minor chelipeds of females are generally smaller than their male counterparts and may exhibit slightly different proportions. Alpheus thompsoni sp. nov. can be separated from A. djiboutensis, as described by De Man (1909), by the shape of the antennal scaphocerite, more precisely, by the blade reaching slightly beyond the adjacent distolateral tooth in the new species (Fig. 1A) vs. by the distolateral tooth by far overreaching the blade in A. djiboutensis (De Man 1909: fig. 19; see also Banner & Banner 1982: fig. 55a, m). In addition, in A. thompsoni sp. nov., the first subarticle of the second pereiopod carpus is almost as long as the second (Fig. 1F), whereas in A. djiboutensis, the first subarticle is noticeably longer than the second (De Man 1909: fig. 23). The two species also seem to differ by the proportions of the major and minor chelae, which are much stouter in A. djiboutensis (cf. Fig. 2 and De Man 1909: 20, 21), although this character needs to be confirmed in the presence of adult male specimens in the future [same remark as above]. According to De Man (1909), the ventromesial margin of the major cheliped merus of A. djiboutensis is armed with a stout and sharp tooth, which is blunt and poorly developed in A. thompsoni sp. nov. (Fig. 2B). An ongoing study of the goby-associated snapping shrimps of the Red Sea, largely based on material collected off the Red Sea coast of Saudi Arabia in 2012 (FLMNH UF), enabled to distinguish five morphotypes, each with a different, highly diagnostic colour pattern. One of them corresponds to ��� A. ochrostriatus ��� (nomen nudum) of Karplus et al. (1981), as mentioned above, and may or may not be the same as A. thompsoni sp. nov. (Anker, in prep.). The second is a very distinctive morphotype with a unique colour pattern type, reported as ��� A. rubromaculatus ��� (nomen nudum) by Karplus et al. (1981); this clearly new species will be described elsewhere (Anker, in prep.). The remaining three morphotypes are morphologically in line with A. djeddensis and A. djiboutensis, although one of them represents yet another, possibly undescribed species (Anker, in prep.). Although the latter three taxa are still being evaluated taxonomically, their colour patterns are very different from that of A. thompsoni sp. nov., which strengthens its separation from A. djeddensis and A. djiboutensis. As mentioned above, A. thompsoni sp. nov. can be easily separated from A. bellulus, A. macellarius, A. fenneri and A. mannarensis by its diagnostic colour pattern and also, from each of them, by at least two morphological characters. For instance, A. thompsoni sp. nov. differs from A. bellulus by the rostrum much narrower and longer (vs. shorter and subtriangular in A. bellulus), and by the different proportions of the female major and minor chelipeds, especially by the more slender meri, the more sle, Published as part of Anker, Arthur, 2022, Description of two new species of goby-associated snapping shrimps from the tropical western Pacific (Decapoda: Alpheidae: Alpheus), pp. 273-290 in Zootaxa 5092 (3) on pages 274-280, DOI: 10.11646/zootaxa.5092.3.2, http://zenodo.org/record/5881095, {"references":["Karplus, I., Szlep, R. & Tsurnamal, M. (1981) Goby-shrimp partner specificity. I. Distribution in the northern Red Sea and partner specificity. Journal of Experimental Marine Biology and Ecology, 51, 1 - 19. https: // doi. org / 10.1016 / 0022 - 0981 (81) 90151 - 9","Debelius, H. (2001) Crustacea guide of the world. IKAN, Frankfurt, 321 pp.","Randall, J. E., Shao, K. T. & Chen, J. P. (2003) A review of the Indo-Pacific gobiid fish genus Ctenogobiops, with descriptions of two new species. Zoological Studies, 42, 506 - 515.","Poupin, J. (2010) Biodiversite de l'Indo-Pacifique tropical francais. 2514 especes de crustaces decapodes et stomatopodes. Rapport Scientifique, Institut de Recherche de l'Ecole Navale, Brest, 76 pp.","Karplus, I. & Thompson, A. R. (2011) The partnership between gobiid fishes and burrowing alpheid shrimps. In: Patzner, R. A., Van Tassell, J. L., Kovacic, M. & Kapoor, B. G. (Eds.), The biology of gobies. Science Publishers, Enfield, New Hampshire, pp. 559 - 608.","Anker, A. (2000) Taxonomical problems of the goby-associated species of Alpheus (Decapoda, Alpheidae). IOP Diving News, 11 (8), 1 - 7.","Jaafar, Z. & Randall, J. E. (2009) A pictorial review and key to the shrimp gobies of the genus Amblyeleotris of the Red Sea, with description of a new species. Smithiana Bulletin, 10, 23 - 29.","Kuiter, R. H. & Debelius, H. (2009) World atlas of marine fauna. IKAN, Frankfurt, 725 pp.","Minemizu, R. (2013) Coral reef shrimps of Indo-West Pacific. Bun-ichi Sogo Shuppan, Co. Ltd., Tokyo, 144 pp. [in Japanese]","Randall, J. E., Shao, K. T. & Chen, J. P. (2007) Two new shrimp gobies of the genus Ctenogobiops (Perciformes: Gobiidae), from the Western Pacific. Zoological Studies, 46, 26 - 34.","Miya, Y. & Miyake, S. (1969) Description of Alpheus bellulus sp. nov. associated with gobies from Japan (Crustacea, Decapoda, Alpheidae). Publications of the Seto Marine Biological Laboratory, 16, 307 - 314. https: // doi. org / 10.5134 / 175556","Anker, A., Pratama, I. S., Firdaus, M. & Rahayu, D. L. (2015) On some interesting marine decapod crustaceans (Alpheidae, Laomediidae, Strahlaxiidae) from Lombok, Indonesia. Zootaxa, 3911, 301 - 342. https: // doi. org / 10.11646 / zootaxa. 3911.3.1","Anker, A. & De Grave, S. (2016) An updated and annotated checklist of marine and brackish caridean shrimps of Singapore (Crustacea, Decapoda). Raffles Bulletin of Zoology, 34, 343 - 454.","Bruce, A. J. (1994) Alpheus fenneri sp. nov. and A. williamsi sp. nov., two new Indo-West Pacific alpheid shrimps of the brevirostris species group. The Beagle, Records of the Museums and Art Galleries of the Northern Territory, 11, 15 - 28. https: // doi. org / 10.5962 / p. 264023","Purushothaman, P., Abhilash, C. P., Ajith Kumar, T. T. & Lal, K. K. (2021) A new alpheid shrimp, Alpheus mannarensis sp. nov. (Crustacea; Decapoda; Alpheidae) from Gulf of Mannar, southern India. Zootaxa, 5026 (1), 127 - 135. https: // doi. org / 10.11646 / zootaxa. 5026.1.5","Holthuis, L. B. (1958) Contributions to the knowledge of the Red Sea. No. 8. Crustacea Decapoda from the northern Red Sea (Gulf of Aqaba and Sinai Peninsula). I. Macrura. Bulletin of Sea Fisheries Research Station, Haifa, 17, l - 40.","Dehghani, A., Sari, A. & Naderloo, R. (2019) Annotated checklist of the snapping shrimps of the genus Alpheus Fabricius, 1798 (Crustacea: Decapoda: Alpheidae) from the Persian Gulf and Gulf of Oman, Iran. Zootaxa, 4544 (4), 479 - 504. https: // doi. org / 10.11646 / zootaxa. 4544.4.2","De Grave S. & Fransen, C. H. J. M. (2011) Carideorum catalogus: the recent species of the dendrobranchiate, stenopodidean, procarididean and caridean shrimps (Crustacea: Decapoda). Zoologische Mededelingen, 85, 195 - 588.","Coutiere, H. (1897) Note sur quelques especes du genre Alpheus du Musee de Leyde. Notes from the Leyden Museum, 19, 195 - 207.","Coutiere, H. (1899) Les \" Alpheidae \", morphologie externe et interne, formes larvaires, bionomie. In: Annales des Sciences Naturelles. Zoologie. Vol. 8. Masson, Paris, pp. 91 - 560. https: // doi. org / 10.5962 / bhl. title. 13143","De Man, J. G. (1909) Diagnoses of new species of macrurous decapod Crustacea from the \" Siboga-Expedition \". Tijdschrift der Nederlandse Dierkundige Vereeniging, 2, 99 - 125.","Banner, D. M. & Banner, A. H. (1982) The alpheid shrimp of Australia, part III: the remaining alpheids, principally the genus Alpheus, and the family Ogyrididae. Records of the Australian Museum, 34, 1 - 357. https: // doi. org / 10.3853 / j. 0067 - 1975.34.1982.434","Chace, F. A. Jr. (1988) The caridean shrimps (Crustacea: Decapoda) of the Albatross Philippine Expedition, 1907 - 1910, Part 5: Family Alpheidae. Smithsonian Contributions to Zoology, 466, 1 - 99. https: // doi. org / 10.5479 / si. 19436696.391.1"]}

Alpheus viserion sp. n. (Figures 1–6) Type material. Holotype: male (cl 10.9 mm), FLMNH UF 51915, Panama, Bocas del Toro, Isla Colón, Boca del Drago, 9°24’54.9”N 82°19’52.5”W, seagrass meadows, depth 2–3 m, under rock on sand, coll. M. Leray, 1 April 2019. Paratypes: 1 ovigerous female (cl 11.6 mm), FLMNH UF 51916, same collection data as for the holotype; 1 male (cl 7.5 mm), FLMNH UF 51919, same collection data as for the holotype. Description. Medium-sized species of Alpheus (cl range of type material: 7.5–11.6 mm). Carapace smooth, glabrous, not setose, not pubescent. Rostrum short, subtriangular, about twice as long as wide at base, subacute distally, almost reaching distal margin of first article of antennular peduncle; rostral carina feebly developed, rounded dorsally, flattening and broadening posteriorly between orbital hoods (Fig. 1A, B). Orbital hoods well developed, moderately swollen, semi-open frontally, with anterior margin broadly rounded; margin between rostrum and orbital hoods slightly concave; adrostral furrows relatively shallow, margin between post-rostral carina and adrostral furrows relatively smooth, not abrupt (Fig. 1A, B). Pterygostomial angle broad, rounded, not protruding anteriorly (Fig. 1B); cardiac notch well developed. Male first and second pleonal sternites each with sharp median process (Fig. 1D). Female first and second pleonal sternites smooth (Fig. 1I). Telson broad, about 1.8 times as long as maximal width, somewhat tapering distally; lateral margins sinuose, without marked constriction; dorsal surface with two pairs of sharp stout cuspidate setae, both inserted at significant distance from lateral margin; proximal pair inserted at about half-length of telson, with cuspidate setae more widely spaced; distal pair inserted at about 0.7 length of telson; distal margin broadly rounded, fringed with long plumose setae and series of small spiniform setae of somewhat variable length; distolateral angles each with one pair of slender spiniform setae, lateral more slender and half as long as mesial (Fig. 1E, F). Antennular peduncle moderately stout; stylocerite slightly overreaching distal margin of first article of peduncle, with acute point; ventromesial carina of first article with small, triangular, distally subacute, anteriorly directed tooth; second article almost twice as long as wide; third article slightly longer than wide; antennular flagella noticeably uneven in length; mesial flagellum longest, diminishing in thickness towards terminal units; lateral flagellum thickened in its basal half, groups of aesthetascs starting at about eighth or ninth unit (Fig. 1 A–C). Antenna with basicerite rather small, distolateral margin armed with slender, distally sharp tooth; scaphocerite subovate, tapering distally, about twice as long as its greatest width, lateral margin almost straight, faintly concave proximally, distolateral tooth very stout, sharp, clearly overreaching blade, reaching to or slightly beyond third article of antennular peduncle; carpocerite long, stout, overreaching both scaphocerite and antennular peduncle; flagellum stout, noticeably thicker than antennular flagella (Fig. 1A, B). Mouthparts typical for genus in external view. Third maxilliped relatively stout, pediform; coxa with lateral plate distally produced into small subacute tooth; antepenultimate article longest, lateral margin with longitudinal ridge delimiting shallowly depressed ventrolateral surface, mesial margin somewhat rugose; penultimate article about 0.3 times as long as antepenultimate article, about 2.5 times as long as wide proximally, widening distally; ultimate article about 3.5 times as long as wide, tapering distally, mesial surface furnished with several rows of thick serrulate setae, tip unarmed; much longer setae present on distodorsal margin of antepenultimate article, ventral surface of penultimate article and dorsal surface and apex of ultimate article; exopod long, reaching distal margin of antepenultimate article; arthrobranch very large (Fig. 2A, B). Major cheliped (= first pereiopod) with chela of edwardsii - type, with both dorsal and ventral notches of palm well developed and lateral grooves extending from dorsal notch on both mesial and lateral surfaces of palm; ischium very short; merus stout, about twice as long as wide; distodorsal angle blunt; mesial margin smooth, with short, simple, non-spiniform setae; distomesial angle with stout sharp tooth; distolateral angle smooth; carpus very short, cup-shaped, distodorsal margin with setae; chela slightly compressed, relatively slender, about 1.8 times as high (at maximal height) as long; surface smooth, not granulate, with numerous setae, especially on distal half of palm and fingers; dorsal shoulder rounded, not overhanging adjacent dorsal notch, latter deep, moderately broad; dorsomesial and dorsolateral surface of palm each with shallow longitudinal groove extending from dorsal notch posteriorly, reaching linea impressa on lateral surface; ventral shoulder rounded, not protruding forward, adjacent ventral notch deep; ventromesial and ventrolateral surface of palm each with broad, fairly deep oblique groove; fingers slightly longer than half-length of palm; distomesial margin of palm forming almost 90° angle, without tooth; distolateral margin smooth, without tooth; pollex shorter than dactylus, without oblique ridge on mesial surface; dactylus with distal margin broadly rounded; plunger stout, rounded anteriorly feebly bulging distally, with tuft of stamen-shaped sensillae; adhesive disks on dactylus and palm very small (Fig. 3 A–E). Male and female major chelipeds generally very similar, that of female slightly less robust and smaller (cf. Figs. 5A, 6A). Minor cheliped with chela simple, non-balaeniceps in both sexes. Male minor cheliped much smaller and weaker than major cheliped; ischium short; merus stout, about twice as long as wide; distodorsal angle blunt; mesial margin smoth, with simple, non-spiniform setae, distomesial angle with stout sharp tooth; distolateral angle smooth; carpus cup-shaped, more elongate than that of major cheliped; distodorsal margin setose; chela moderately stout; surface smooth, not granulate, very setose, especially on distal half of palm and fingers; palm about twice as long as high, devoid of notches or grooves, distodorsal angle slightly elevated; fingers subequal in length, approximately as long as palm, without balaeniceps ridges, not particularly expanded or modified otherwise; cutting edges bladelike, simple; adhesive disks on dactylus and palm very small (Fig. 3 F–H; see also Fig. 5A). Female minor cheliped similar to that of male, somewhat more slender (cf. Figs. 5A, 6A). Second pereiopod slender; ischium long, about six times as long as wide; merus slightly longer than ischium; carpus subdivided into five units, first longest, almost seven times as long as wide; ratio of carpal units approximately equal to 3.5 – 2.5 – 1 – 1 – 1.7; chela slightly longer than distal-most carpal unit, with fingers slightly longer than palm, furnished with several clusters of simple setae (Fig. 2C). Third pereiopod moderately slender; ischium about 1.6 times as long as wide, ventrolateral surface with stout cuspidate seta; merus about 4.2 times as long as wide; distomesial margin unarmed; distodorsal margin furnished with long setae; carpus about five times as long as wide, about half as long as merus, much slenderer than merus; propodus about six times as long as wide; ventral margin with row of eight or so spiniform setae in addition to one pair of stouter spiniform near articulation with dactylus; dactylus simple, conical, somewhat elongate, with gentle curvature, about half-length of propodus (Fig. 4A, B). Fourth pereiopod similar to third, noticeably slenderer; ischium about 2.5 times as long as wide, ventrolateral surface with stout cuspidate seta; propodus with ventral margin armed with row of eight or so spiniform setae in addition to one distal pair of spiniform setae near dactylar base; dactylus somewhat shorter and stouter than than of third pereiopod, otherwise similar (Fig. 4C, D). Fifth pereiopod slenderest of all; ischium with ventrolateral surface armed with stout spiniform seta; merus almost six times as long as wide; carpus slightly shorter than merus, about 5.5 times as long as wide; propodus with row of five spiniform setae in addition to one distal spiniform seta near dactylar base, on ventromesial side; distal half of propodus with about eight transverse rows of serrulate setae forming cleaning brush; dactylus almost 0.6 times as long as propodus, otherwise similar to that of third pereiopod (Fig. 4 E–G). First to fifth pleopods without spiniform setae on ventrolateral margin of protopod in both sexes (Fig. 2D, E, G, H). Male first pleopod with endopod reduced, much smaller than exopod (Fig. 2D). Male second pleopod with endopod and exopod similar in size; appendix masculina small, about 0.6 times as long as appendix interna, with stiff setae along most of its surface (Fig. 2E, F). Female first and second pleopods similar to those of male; protopod with long flexible setae for egg retention (Fig. 2G, H). Developing embryos moderately large, about 0.8 mm long and 0.6 mm wide (Fig. 1J). Uropod with endopod and exopod broadly rounded; exopod broader and slightly longer than endopod, both fringed with plumose setae; protopod with distolateral and distomedial lobes each produced into strong triangular tooth; exopod with sharp distolateral tooth and stout distolateral spiniform seta; diaeresis sinuous, with broadly rounded tooth laterally, near base of distolateral spiniform seta; endopod with row of small spiniform setae of variable length on distal margin, interrupted by cluster of few long setae (Fig. 1G, H). Colour pattern. Body and appendages generally appearing as olive-green; carapace, pleon, telson and uropods with olive-green to brown-greenish background, speckled with pale greenish-yellow chromatophores, many interconnecting and forming nets of greenish-yellow spots and chains; antennular peduncles dark brown-green, stylocerite with pale green spots; antennal scaphocerite greenish, with some larger pale green blotches; antennular and antennal flagella pale olive-green; major chela almost uniformly olive-green on mesial surface, except for small cluster of paler blotches between linea impressa and dorsal notch, one large pale green area on proximal half of palm, and contrasting pinkish fingertips; minor cheliped uniformly olive green on mesial surface, with paler palm; second to fifth pereiopods pale olive-green to olive-yellow (Figs. 5, 6). Etymology. The new species’ name refers to Viserion, one of Daenerys Targaryen’s three dragons of the popular TV series “Game of Thrones”. The dragon-related name was originally inspired from the Spanish name of the collection site, Boca del Drago (= dragon’s mouth). Used as a noun in apposition. Distribution. West Atlantic: presently known only from the type locality on the Caribbean coast of Panama. Ecology. All three specimens were collected under large rocks on fine white sand approximately 1–2 m away from a dense meadow of turtlegrass (Thalassia testudinum Koenig), at a depth of about 2– 3 m. Remarks. Alpheus viserion sp. n. is morphologically closest to several species of the transisthmian, taxonomically challenging A. armillatus H. Milne-Edwards, 1837 species complex (see below), which was studied with three molecular markers by Mathews & Anker (2009). The western Atlantic members of the A. armillatus complex were subsequently revised by Anker (2012). The eastern Pacific species of the A. armillatus complex, among them two taxonomically problematic species and at least two undescribed ones (Mathews & Anker 2009), will be revised in the near future (A. Anker, in prep.). All members of the A. armillatus species complex are characterised by the rostral carina conspicuously flattened between the orbital hoods, forming a more or less well-defined, V-shaped or U-shaped post-rostral plate, rising above the usually deep adrostral furrows; the major chela of a basic edwardsii - type (see above); the merus of the cheliped armed with a strong distomesial tooth; the minor chela non-balaeniceps in both sexes; the ambulatory pereiopods with simple, conical dactyli; and the third and fourth pereiopods with distomesially unarmed meri (modified from Anker 2012). Additionally, in several species, the pleon is armed with sternal processes, usually on the first two sternites, sometimes on the first four sternites (Kim & Abele 1988; McClure 2002; Anker 2012). The morphological features of A. viserion sp. n. are generally consistent with the characteristics of the A. armillatus complex, although the post-rostral plate of the new species is less obvious and not abruptly delimited from the adjacent adrostral furrows as in most species of the A. armillatus complex (Fig. 1A). Among the western Atlantic members of the A. armillatus complex, only two species are characterised by the feebly demarcated, roughly V-shaped post-rostral plate, viz. A. carlae Anker, 2012 and A. angulosus McClure, 2002 (McClure 1995, 2002; Anker 2012), both of them also occurring in Bocas del Toro (Anker 2012). All other western Atlantic species of the A. armillatus complex have abruptly delimited, V- or U-shaped post-rostral plates and present additionally a series of other morphological differences and/or have different colour patterns (Anker 2012). Alpheus carlae has a wide distribution in the western Atlantic, ranging from Florida through the Caribbean Sea to southern Brazil (Anker 2012). The species seems to be genetically strongly structured (Mathews & Anker 2009) and exhibits a great deal of variation in both morphology and colouration, although the colour pattern type is essentially the same (Anker 2012). Alpheus viserion sp. n. seems to share more morphological characters with A. carlae than with any of the other species of the A. armillatus complex. These include the identical armature of the pleonal sternum (with a median process on each of the first two pleonites, although with some variation); the similar proportions of the major chela; the presence of a cuspidate seta on the ischium of the fifth pereiopod; and the appendix masculina being distinctly shorter than the appendix interna (Figs. 1D, 2F, 3A, D, 4E; cf. Anker 2012: figs. 41H–J, 42A, B, 43O). The new species can be separated morphologically from A. carlae by at least three relatively subtle characters, viz. the shape of the post-rostral area (with an anteriorly wide post-rostral carina, broadening and flattening between the eyes in A. viserion sp. n. vs. with an anteriorly narrow post-rostral carina, widening and flattening only posterior to the level of the eyes in A. carlae); the major chela dactylus longer and with a much less bulging plunger; and the male minor chela without a trace of a ventral notch on the palm (vs. with a slight sinus in A. carlae) (Figs. 1A, 3E, H; cf. Anker 2012: figs. 41A, 42D–F, 43F, K). In addition, the colour patterns of A. viserion n. sp. and A. carlae are radically different. The new species has uniformly olive-green background colour, speckled with pale green-yellow spots, and a homogenously coloured major and minor chelae (Figs. 5, 6). In contrast, in A. carlae, the pleon displays transverse bands of red-brown, green-brown or dark brown colour, with at least two pairs of large black lateral spots, and the chelae are brownish speckled with white blotches or dots (Anker 2012: figs. 44–48). Alpheus angulosus is another genetically highly structured western Atlantic snapping shrimp, occurring from South Carolina to southern Brazil and throughout the Caribbean Sea and Gulf of Mexico (Mathews & Anker 2009; Anker 2012; Heuring et al. 2017). As in the previous species, the populations of A. angulosus exhibit some variation in the colouration of the body and chelipeds (Anker 2012), which apparently is not directly linked to the genetic structure of the species (Heuring et al. 2017). Alpheus viserion sp. n. differs from A. angulosus by the presence of a strong median sternal process on each of the first two pleonites in males (vs. with a weak median sternal process on the first pleonite and without such a process on the second pleonite in males of A. angulosus); the telson noticeably narrower, about 1.8 times as long as maximally wide (vs. broader, about 1.5 times as long as wide, in A. angulosus); the blade of the antennal scaphocerite distally wider and approaching more the point of the distolateral tooth (vs. distally narrower and with the distal margin greatly exceeded by the distolateral tooth in A. angulosus); the male major chela with the fingers longer relative to the palm and with the dactylar plunger shorter and more truncate (i.e. less bulging distally); the male minor chela without trace of a ventral notch (typically present in larger specimens of A. angulosus); the ischium of the fifth pereiopod armed with a stout cuspidate seta (vs. unarmed in A. angulosus); and the appendix masculina much shorter than the appendix interna (vs. the two being subequal in length in A. angulosus) (Figs. 1A, D, E, 2F, 3A, H, 4E; cf. Anker 2012: figs. 19D, H–K, 20A, D, E). In addition, the two species present clear differences in their colour patterns, despite a generally similar, uniform colouration of the carapace and pleon. In A. viserion sp. n., the background of the body is olive-green to olive-brown, speckled with pale green-yellow dots (Figs. 5, 6), whilst in A. angulosus, in contrast to the new species, the background varies from pale greenish-grey to bluish-grey and is speckled with darker brownish or dark greenish-red dots (Anker 2012: figs. 22–26). Furthermore, in A. viserion sp. n., the palm of the major chela is uniformly olive-green in dorsal view (= mesial side of the chela), except for a paler patch on the proximal half of the palm (Figs. 5A, 6A), whereas in A. angulosus, the palm of the major chela is generally more patterned, for instance, the dorsal and ventral notches and adjacent oblique grooves are always conspicuously marked with white or pale yellow (Anker 2012: figs. 22–26). The only eastern Pacific species of the A. armillatus complex, which has a smoothly delimited post-rostral plate, is A. tenuis Kim & Abele, 1988, known from Panama to Mexico (Kim & Abele 1988; Villalobos Hiriart et al. 1989; Mathews & Anker 2009). The taxonomic identity of A. tenuis is presently being investigated due to the presence of two genetically distinct and apparently geographically at least partly overlapping clades in the eastern Pacific (Mathews & Anker 2009). Nevertheless, A. viserion sp. n. can be easily separated from A. tenuis sensu Kim & Abele (1988) by the major chela palm with the dorsal shoulder not protruding and not overhanging the dorsal notch (vs. clearly overhanging it in A. tenuis); the ventromesial carina of the first article of the antennular peduncle b, Published as part of Anker, Arthur, Leray, Matthieu & Pachelle, Paulo P. G., 2021, A new species of Alpheus Fabricius, 1798 (Decapoda: Caridea: Alpheidae) from the Caribbean coast of Panama, pp. 379-390 in Zootaxa 4933 (3) on pages 380-389, DOI: 10.11646/zootaxa.4933.3.5, http://zenodo.org/record/4554997, {"references":["Mathews, L. M. & Anker, A. (2009) Molecular phylogeny reveals extensive ancient and ongoing radiations in a snapping shrimp species complex (Crustacea, Alpheidae, Alpheus armillatus). Molecular Phylogenetics and Evolution, 50, 268 - 281. https: // doi. org / 10.1016 / j. ympev. 2008.10.026","Kim, W. & Abele, L. G. (1988) The snapping shrimp genus Alpheus from the eastern Pacific (Decapoda: Caridea: Alpheidae). Smithsonian Contributions to Zoology, 454, i-iv + 1 - 119. https: // doi. org / 10.5479 / si. 00810282.454","McClure, M. R. (2002) Revised nomenclature of Alpheus angulatus McClure, 1995 (Decapoda: Caridea: Alpheidae). Proceedings of the Biological Society of Washington, 115, 368 - 370.","McClure, M. R. (1995) Alpheus angulatus, a new species of snapping shrimp from the Gulf of Mexico and northwestern Atlantic, with a redescription of A. heterochaelis Say, 1818 (Decapoda: Caridea: Alpheidae). Proceedings of the Biological Society of Washington, 108, 84 - 97.","Heuring, W. L., Hill-Spanik, K. M. & Sotka, E. E. (2017) Identifying lineages of Alpheus angulosus McClure, 2002 (Caridea: Alpheidae) in South Carolina, USA. Journal of Crustacean Biology, 37, 499 - 502. https: // doi. org / 10.1093 / jcbiol / rux 056"]}

Alpheus inopinatus Holthuis & Gottlieb, 1958 (Figs. 1, 3 A���C) Alpheus inopinatus Holthuis & Gottlieb 1958: 42, figs. 8, 9; Lewinsohn & Holthuis 1964: 51; Tirmizi & Kazmi 1969: 99 (379), fig. 1a; Karim & Hafizur 1974: 26, pl. 4, figs. 1���4b; Banner & Banner 1978: 223; Kazmi & Kazmi 1979: 153; Barkati 1980: 355; Kocataş 1981: 162; Banner & Banner 1982: 241, fig. 75; Lewinsohn & Galil 1982: 208; Afzal et al. 1986: 339; Jeng 1997: 13; Cunha et al. 2020: 343, 351, fig 5C���D, 6C���D, 7C���D. Alpheus lobidens. ��� Jones 1986: 152, pl. 42 (partim ?); Geldiay 1969: 3, 5 (partim?); Kocataş & Kataĝan 2003: 65, 70 (partim ?); Dehghani et al. 2019a: 486, fig. 4A���D (partim ?) [not A. lobidens De Haan, 1849]. Alpheus sp. ��� Forest & Guinot 1958: 6, figs. 3���7 (see Banner & Banner 1982: 243). Material examined. Kuwait: 1 male, OUMNH. ZC.2020-01-001, Shaab, 29��21���16.78���N 48��01���47.38���E, mixed rocky-muddy intertidal, under rocks, leg. A. Anker, M. Al-Kandari, S. De Grave, Z. Sattari, 25 September 2019 [fcn KUW-011]; 1 male, OUMNH. ZC. 2020-01-002, same collection data as for previous specimen [fcn KUW-030]; 2 males, 1 female, OUMNH. ZC.2020-01-003, same collection data as for previous specimens [fcn KUW-034]; 1 male, OUMNH. ZC.2020-01-004, Failaka Island, eastern side, 29��23.409���N 48�� 23.927���E, mixed rocky-muddy intertidal, under rocks, leg. M. Al-Kandari, 3 October 2019 [fcn KUW-077]; 1 male, 1 female, KISR, Al Julaia���a, Seashell Hotel, leg. M. Al-Kandari, 14 December 2019. Comparative material. Alpheus cf. lobidens De Haan, 1849. Kuwait: 1 male, 1 ovigerous female, OUMNH. ZC.2020-01-005, Ras Ajuza, 29��23.656���N 47��59.840���E, mixed rocky-muddy intertidal, under rocks, leg. M. Al-Kandari, Z. Sattari, 25 September 2019 [fcn KUW-027]; 1 female, OUMNH. ZC.2020-01-006, same collection data as for previous specimens [fcn KUW-029]; 1 male, OUMNH. ZC.2020-01-007, Shaab, 29��21���16.78���N 48��01���47.38���E, mixed rocky-muddy intertidal, under rocks, leg. A. Anker, M. Al-Kandari, S. De Grave, Z. Sattari, 25 September 2019 [fcn KUW-035]; 1 male, 1 ovigerous female, OUMNH. ZC.2020-01-008, same collection data as for previous specimen [fcn KUW-036]; 1 female, OUMNH. ZC.2020-01-009, Sulaibikhat Bay, 29��19.548���N 47��49.35���E, intertidal mud flat, in burrows, suction (yabby) pump, leg. A. Anker, M. Al-Kandari, Z. Sattari, 7 October 2019 [fcn KUW-107]; 1 male, 1 female, OUMNH. ZC.2020-01-010, same collection data as for previous specimen [fcn KUW-108]. Description. See Holthuis & Gottlieb (1958), Tirmizi & Kazmi (1969), Karim & Hafizur (1974) and Banner & Banner (1982). Colour pattern. Pale greenish or bluish grey to various tones of brown, often darker in females; pleon with three pairs of small dark spots distributed as following: one pair of small lateral spots on second pleuron, one pair of small dorsolateral spots on third pleuron (sometimes very faint and inconspicuous) and one pair of larger, more conspicuous lateral spots on fourth pleuron; mesial side of major and minor chelae greenish-brownish, with some olive or orange areas; dactylar plunger of major chela creamy white with some bluish tinge proximally; antennular and antennal flagella pale greenish-yellow; second pereiopods and walking legs (third to fifth pereiopods) pale reddish or pinkish with whitish articulations; tail fan brownish or greenish, with bluish area on uropodal exopod (Figs. 1, 3 A���C). General distribution. Possibly widely distributed in the Indo-West Pacific with records from the Red Sea, Djibouti, Arabian-Persian Gulf (Kuwait, Iran, see Remarks), Oman, Pakistan, Hong Kong, Taiwan, Australia (Tirmizi & Kazmi 1969; Banner & Banner 1978, 1981, 1982; Jeng 1997; present study); however, all records from outside of the western Indian Ocean require confirmation (see Remarks). Lessepsian migrant with a well-established population in the eastern Mediterranean Sea, with confirmed records from Israel and Turkey (Holthuis & Gottlieb 1958; Kocataş 1981; Lewinsohn & Galil 1982). Ecology. Largely intertidal, sometimes also in the shallow subtidal (to at least 6 m), rocky and mixed sandyrocky shores, often heavily silted or with a significant mud component; typically on sand/mud under large rocks and coral rubble (Lewinsohn & Galil 1982; present study). Remarks. Alpheus inopinatus belongs to the A. lobidens De Haan, 1849 species complex, together with A. crassimanus Heller, 1862 (currently considered to be a synonym of A. lobidens), A. lobidens polynesicus Banner & Banner, 1974 (originally described as A. lobidens polynesica), and the recently described A. mohammadpouri Dehghani, Sari & Naderloo, 2019 (Banner & Banner 1974, 1982; Dehghani et al. 2019b), together with a large number of undescribed species. The complicated taxonomic situation of A. lobidens and the presence of several cryptic or pseudocryptic species in this species complex make identifications of specimens extremely difficult (Anker & De Grave 2016). Alpheus inopinatus was originally described based on material from the Mediterranean coast of Israel, as an ���immigrant from the Red Sea��� (Holthuis & Gottlieb 1958). This species is morphologically very close to another species of the A. lobidens complex, which is present in the Gulf and possibly throughout the western and northern Indian Ocean (including the Red Sea), and identified either as A. lobidens (type locality: Japan) or A. crassimanus (type locality: Nicobar Islands, India). Like A. inopinatus, this species also penetrated into the Mediterranean Sea through the Suez Canal, spreading along the North African coasts from Egypt westwards to Tunisia, and northwards to Syria and Turkey (Hasan et al. 2008; Bakir et al. 2015). Older Mediterranean records of this species are under the name A. crassimanus (e.g., Balss 1936; Forest & Guinot 1956, 1958; Holthuis & Gottlieb 1958; Geldiay 1969), which was later placed in the synonymy of A. lobidens (see discussion in Banner & Banner 1974, 1981, 1982), with more recent records under the name A. lobidens (e.g., Bakir et al. 2015). In the present study, we tentatively continue to accept the synonymy of A. crassimanus with A. lobidens proposed by Banner & Banner (1974, 1982), who compared the Japanese specimens they identified as A. lobidens with the material from other areas of the Indo-West Pacific. However, we feel that a thorough morphological and molecular study is needed to establish the true identity of this taxon in light of the fact that their view of A. lobidens as a single widespread species has now been refuted (see also Cunha et al. 2020). The colour pattern of A. crassimanus is currently unknown, which is especially relevant in light of growing evidence of the presence of several cryptic or pseudocryptic species within the A. lobidens complex distinguishable mainly by colour patterns, for instance, in Singapore (Anker & De Grave 2016), Taiwan (A. Anker, pers. obs.), Japan (K. Nomura, pers. comm.), and Iran (Dehghani et al. 2019a). Forest & Guinot (1958) were aware of the presence of two morphologically distinct species from the A. lobidens complex on the Mediterranean coast of Israel, viz. A. inopinatus (reported as Alpheus sp., as it was already being studied by Holthuis & Gottlieb) and A. lobidens (reported by them as A. crassimanus). These authors also examined several specimens in Henri Couti��re���s alpheid collection in the Mus��um National d���Histoire Naturelle in Paris (MNHN), which morphologically corresponded well to A. inopinatus, viz. two specimens from the Gulf of Oman and a mixed lot from Djibouti, containing both A. inopinatus and A. lobidens. Interestingly, Holthuis & Gottlieb (1958) contrasted A. inopinatus with A. lobidens only superficially, comparing it with three more distantly related species of Alpheus, whereas Forest & Guinot (1958) provided a full account of the morphological differences between the two taxa (see below). Banner & Banner (1981, 1982) questioned the separation of A. inopinatus from A. lobidens and synonymised the former species. The Hawaiian subspecies, A. lobidens polynesicus, was also placed in the synonymy of A. lobidens, despite some rather striking colour pattern differences (Banner & Banner 1981). However, at least some of the synonymies proposed by Banner & Banner (1981, 1982) in the A. lobidens complex were premature (Anker 2001; Anker & De Grave 2016). New material collected throughout the Indo-West Pacific clearly shows that several features of the colour pattern, such as the presence or absence of dark transverse bands on the pleon, the presence or absence of dark spots on the sides of the pleon, the markings on the mesial side of the major and minor chelae and the colour of the dactylar plunger, may provide additional differentiating characters between the species of the A. lobidens complex (Anker & De Grave 2016; A. Anker, pers. obs., see also Fig. 3). De Grave & Fransen (2011) treated A. inopinatus as a valid species, but continued to list A. lobidens polynesicus as a junior synonym of A. lobidens. The present material confirms that A. inopinatus is clearly distinct from the west Indian Ocean taxon often recorded as A. lobidens and thus represents a valid species (see also Cunha et al. 2020 for molecular comparison). The common and widespread taxon from the western Indian Ocean (e.g., Red Sea, Gulf, Pakistan), which penetrated into the Mediterranean Sea most likely does not represent A. lobidens sensu De Haan (1849) from Japan and is herein referred to A. cf. lobidens (see Comparative material). Whether this taxon is identical to A. crassimanus sensu Heller (1862) remains to be shown. Alpheus inopinatus can be separated from A. cf. lobidens by at least three morphological characters, in addition to a small but consistent difference in their colour patterns. Both species can also be separated morphologically from A. lobidens polynesicus and A. mohammadpouri. The taxonomic status of these taxa will be discussed elsewhere, as part of a larger study of the A. lobidens complex. Many of the differentiating characters between A. inopinatus and A. cf. lobidens listed by Forest & Guinot (1958) were confirmed in the present material. The most important character separating the two species is the absence of a sharp distomesial tooth on the meri of the major and minor cheliped in the former species. In some specimens of A. inopinatus, the distomesial margin terminates in a subacute angle, sometimes protruding as a minute tooth, which is not the same structure as a much stronger acute tooth, situated slightly below the distomesial angle, in A. cf. lobidens (cf. Forest & Guinot 1958: figs. 1, 3, 4). The male minor cheliped has a much stouter merus and a stouter and more sculptured chela in A. inopinatus, compared to the noticeably slenderer merus and chela of A. cf. lobidens (cf. Forest & Guinot 1958: figs. 2, 5). The third and fourth pereiopods are also stouter in A. inopinatus, as expressed by the more inflated merus, with a length-width ratio of about 3.0 (cf. Forest & Guinot 1958: fig. 7) vs. more than 4.0 in A. cf. lobidens from Kuwait or the specimens from Australia identified as A. lobidens by Banner & Banner (1982: fig. 78f). In life, A. inopinatus and A. cf. lobidens can be separated by their colour patterns, although the difference is rather subtle. The pleon of A. inopinatus presents three pairs of small dark spots: one pair of lateral spots on the second pleuron, one pair of very small dorsolateral spots on the third pleuron and one pair of more conspicuous lateral spots on the fourth pleuron (Fig. 1); all these spots are absent in A. cf. lobidens (Fig. 2). The dactylar plunger of the major chela of both species is similar in colour, whitish with creamy or yellow tinge, although in A. inopinatus, the base of the plunger has a bluish tinge (Fig. 3C), whilst in A. cf. lobidens, the anterior surface has a more intense pale orange colour (Fig. 3F). The Iranian material of A. lobidens listed by Dehghani et al. (2019a) likely contains both A. inopinatus and A. cf. lobidens, based on an analysis of the colour patterns of the four illustrated individuals. Generally, many records under the name ��� A. lobidens ��� from the Gulf, Red Sea, Mediterranean Sea and other localities may refer to A. inopinatus as the two species have been commonly confused in the past, especially after the synonymisation of A. inopinatus by Banner & Banner (1981, 1982). Anker & De Grave (2016), while reporting A. lobidens (as A. lobidens sensu lato) from Singapore, provided colour photographs of three different colour morphs, possibly three different species, of the A. lobidens species complex. They can be briefly described as (1) pleon non-banded, with two conspicuous lateral black spots on the second and fourth pleura and two small medially fused black spots on the dorsal surface of the fifth pleonite (Anker & De Grave 2016: fig. 16A, E); (2) pleon banded, with two small dorsolateral black spots on the third pleuron, two very large and conspicuous black lateral spots on the fourth pleuron and two smaller black lateral patches on the fifth pleuron (idem.: fig. 16C, D); and (3) pleon non-banded, without spots (idem.: fig. 17). The arrangement of the spots of the Singaporean specimens appears to be different from that of A. inopinatus. In addition, the dactylar plunger of the major chela of the two black-spotted colour morphs from Singapore is bright orange, which makes them clearly different from both A. inopinatus and the specimens herein identified as A. cf. lobidens, both with whitish-creamy plungers (Fig. 3C, F). Therefore, all material identified as A. inopinatus from outside of the western Indian Ocean (Hong Kong, Taiwan, Australia) needs to be carefully re-examined. These observations also contribute to the remaining uncertainty about the taxonomic status of A. crassimanus, currently regarded as a synonym of A. lobidens (see above). Recently, Cunha et al. (2020) redescribed A. lobidens from Nagasaki on the basis of what they considered to be topotypic specimens, collected from the vicinity of Nagasaki. However, the type locality of A. lobidens is Japan, i.e., the entire Japanese archipelago and not specifically Nagasaki (De Grave & Fransen 2011: 385). Only six species described in De Haan (1833 ���1850) actually have a type locality that is geographically more restricted than merely Japan (see Yamagushi & Baba 1993), so it cannot be assumed that the holotype of A. lobidens was indeed collected in the Nagasaki area. Furthermore, it is well known that the botanical and zoological collections gathered by Von Siebold and B��rger, a part of which was studied by De Haan, came from all over Japan. For example, the type series of the giant spider crab Macrocheira kaempferi (Temminck) was collected in Suruga Province, further east, whereas their collection also contained cold-water decapods from much further north, e.g., Lithodes japonicus Von Siebold, now known as Paralithodes camtschaticus (Tilesius). The authors simply ������assumed that the topotypes from Shimabara, Nagasaki, Japan (RMNH PEG 25134) correspond to A. lobidens sensu stricto, i.e., are conspecifics of the damaged holotype ������, without a direct comparison to the holotype. Although it is possible that Cunha et al. (2020) did redescribe what in future could be considered as true A. lobidens (i.e. A. lobidens sensu De Haan, 1849), it should be noted that numerous species of the A. lobidens complex occur in Japan, an observation based primarily on the diversity of colour patterns (K. Nomura, pers. comm.). As no colour pattern was documented for A. lobidens sensu Cunha et al. (20202), any further comparison with other taxa is hindered, as it is obvious now that colour patterns, along with molecular analyses and morphology, will be crucial to resolve this challenging species complex. For example, in Singapore (Anker & De Grave 2016), Kuwait (present study) and some localities in eastern Australia (A. Anker, pers. obs.), two or more species of the A. lobidens complex occur syntopically, with the easiest separation between them being consistent differences in the colour patterns. Al-Maliky et al. ( 2017) reported and illustrated Alpheus edwardsii (Audouin, 1826) from Basrah area in Iraq. Although their photographs of preserved specimens are in very low resolution, the dorsal shoulder of the major chela is not overhanging the adjacent transverse groove (idem: figs. 2, 3A), whilst the merus of the major cheliped seems to be armed with a strong distomesial tooth (idem: fig. 3B, D). Therefore, the Iraqi material can be tentatively referred to A. cf. lobidens, as suggested by Al-Khandari et al. (2020: p. 254; mind a small error therein, read correctly: ��� Al-Maliky et al. (2017), who erroneously reported A. lobidens as A. edwardsii from Shatt-Al-Basrah and Faw in Iraq.���)., Published as part of Anker, Arthur, Al-Kandari, Manal & Grave, Sammy De, 2020, Taxonomic notes on Alpheus inopinatus Holthuis & Gottlieb, 1958 and Alpheus cf lobidens De Haan, 1849 from Kuwait (Malacostraca: Decapoda: Alpheidae), pp. 189-197 in Zootaxa 4851 (1) on pages 190-195, DOI: 10.11646/zootaxa.4851.1.10, http://zenodo.org/record/4407192, {"references":["Holthuis, L. B. & Gottlieb, E. (1958) An annotated list of the Decapod Crustacea of the Mediterranean Coast of Israel, with an appendix listing the Decapoda of the eastern Mediterranean. Bulletin of the Research Council of Israel, 7 B, 1 - 126, pls. 1 - 3.","Lewinsohn, C. & Holthuis, L. B. (1964) New records of decapod Crustacea from the Mediterranean coast of Israel and the eastern Mediterranean. Zoologische Mededelingen, 40, 45 - 63.","Tirmizi, N. M. & Kazmi, M. A. (1969) The occurrence of Alpheus inopinatus Holthuis & Gottlieb, 1958 in the Arabian Sea (Decapoda, Alpheidae). Crustaceana, 16, 99 - 100. https: // doi. org / 10.1163 / 156854068 X 00250","Karim, S. I. & Hafizur, R. (1974) On some Carides of Karachi coasts. Records of Zoological Survey, 6, 25 - 28, pls. 1 - 4.","Banner, D. M. & Banner, A. H. (1978) Annotated checklist of alpheid and ogyridid shrimp from the Philippine Archipelago and the South China Sea. Micronesica, 14, 215 - 257.","Kazmi, M. A. & Kazmi, Q. B. (1979) A check-list of marine caridean prawns of Pakistan. Biologia, 25, 151 - 157.","Barkati, S. (1980) Sex ratio, fecundity and larval development of Alpheus inopinatus Holthuis and Gottlieb, 1958 (Crustacea, Decapoda, Caridea). Proceedings of the First Pakistan Congress of Zoology, B, 355 - 359.","Kocatas, A. (1981) Liste preliminaires et repartition des crustaces decapodes des eaux turques. Rapports de la Commission Internationale de la Mer Mediterranee, 27, 161 - 162.","Banner, D. M. & Banner, A. H. (1982) The Alpheid Shrimp of Australia, part III: the remaining alpheids, principally the genus Alpheus, and the family Ogyrididae. Records of the Australian Museum, 34, 1 - 357. https: // doi. org / 10.3853 / j. 0067 - 1975.34.1982.434","Lewinsohn, C. & Galil, B. (1982). Notes on species of Alpheus (Crustacea Decapoda) from the Mediterranean coast of Israel. Quaderni del Laboratorio di Tecnologia della Pesca, Ancona, 3, 207 - 210.","Afzal, M., Javed, A. & Barkati, S. (1986) Checklist, synonymy and key to the genera and species of alpheid shrimps of Karachi coast. Pakistan Journal of Agriculture Research, 7, 337 - 340.","Jeng, M. S. (1997) Studies on the land and aquatic decapod crustacean fauna of the Kenting National Park (II). Conservation Research Report, 96, 1 - 66.","Cunha, A. M., Terossi, M., Mantelatto, F. M. & Almeida, A. O. (2020) Delimiting the snapping shrimp Alpheus lobidens De Haan, 1849 (Caridea: Alpheidae) based on morphological and molecular data. Zootaxa, 4718 (3), 337 - 354. https: // doi. org / 10.11646 / zootaxa. 4718.3.3","Jones. D. A. (1986) A field guide to the sea shores of Kuwait and the Arabian Gulf. University of Kuwait, Kuwait City, 192 pp.","Geldiay, R. 1969. A report on the collection of Natantia (Crustacea, Decapoda) along the coast of Turkey from the eastern Mediterranean to the vicinity of Izmir. Scientific Reports of the Faculty of Science Ege University, 74, 1 - 17.","Kocatas, A. & Katagan, T. (2003) The decapod crustacean fauna of the Turkish seas. Zoology in the Middle East, 29, 63 - 74. https: // doi. org / 10.1080 / 09397140.2003.10637971","Dehghani, A., Sari A. & Naderloo, R. (2019 a) Annotated checklist of the snapping shrimps of the genus Alpheus Fabricius, 1798 (Crustacea: Decapoda: Alpheidae) from the Persian Gulf and Gulf of Oman, Iran. Zootaxa, 4544 (4), 479 - 504. https: // doi. org / 10.11646 / zootaxa. 4544.4.2","Forest, J. & Guinot, D. (1958) Sur une collection de crustaces decapodes des cotes d'Israel. Bulletin of the Sea Fisheries Research Station of Israel, 15, 4 - 16.","Banner, D. M. & Banner, A. H. (1981) Annotated checklist of the alpheid shrimp of the Red Sea and Gulf of Aden. Zoologische Verhandelingen, Leiden, 190, 1 - 99.","Heller, C. (1862) Neue Crustaceen gesammelt wahrend der Weltumsegelung der k. k. Fregatte Novara. Zweiter vorlaufiger Beri- cht. Verhandlungen der Koeniglich-Kaiserlichen Zoologisch-Botanischen Gesellschaft, Wien, 28, 519 - 528.","Banner, A. H. & Banner, D. M. (1974) Contributions to the knowledge of the alpheid shrimp of the Pacific Ocean, part XVII: Additional notes on the Hawaiian alpheids: new species, subspecies and some nomenclatorial changes. Pacific Science, 28, 423 - 437.","Dehghani, A., Sari, A. & Naderloo, R. (2019 b) Three new species of narrowly endemic snapping shrimp, genus Alpheus (Decapoda: Caridea: Alpheidae) from the Persian Gulf. Journal of the Marine Biological Association of the United Kingdom, 99, 911 - 920. https: // doi. org / 10.1017 / S 0025315418000693","Anker, A. & De Grave, S. (2016) An updated and annotated checklist of marine and brackish caridean shrimps of Singapore (Crustacea, Decapoda). Raffles Bulletin of Zoology, Supplement 34, 343 - 454.","Hasan, H., Zeini, A. & Noel, P. Y. (2008) The marine decapod Crustacea of the area of Lattakia, Syria. Crustaceana, 81, 513 - 536. https: // doi. org / 10.1163 / 156854008784092175","Bakir, K., Ilkyaz, A. T., Aydin, C. & Turkmen, G. (2015) The presence of Alpheus lobidens De Haan, 1849 (Decapoda, Alpheidae) on the Turkish Aegean Sea coast. Crustaceana, 88, 651 - 656. https: // doi. org / 10.1163 / 15685403 - 00003434","Forest, J. & Guinot, D. (1956) Sur une collection de crustaces decapodes et stomatopodes des mers tunisiennes. Bulletin de la Station Oceanographique Salammbo, 53, 24 - 43.","Anker, A. (2001) Two new species of snapping shrimps from the Indo-Pacific, with remarks on colour patterns and sibling species in Alpheidae (Crustacea: Caridea). Raffles Bulletin of Zoology, 49, 57 - 72.","De Grave, S. & Fransen, C. H. J. M. (2011) Carideorum catalogus: the recent species of the dendrobranchiate, stenopodidean, procarididean and caridean shrimps (Crustacea: Decapoda). Zoologische Mededelingen, Leiden, 85, 195 - 589.","De Haan, W. (1833 - 1850) Crustacea. In: von Siebold, P. F. (Ed.), Fauna Japonica sive descriptio animalium, quae in itinere per Japoniam, jussuet auspicilis superiorum, qui summum in India Batava Imperium tenent, suspecto, annis 1823 - 1830 collegit, notis, observationibus et dumbrationibus illustravit. Fasc. 1 - 8. Apud Auctorem, Lugduni Batavorum, xvii + xxxi + 244 pp., 74 pls. [pp. i-xvii + i-xxxi + 1 - 244, pls. 1 - 55 + A-Q + 1 + 2. pp. i-xxi + vii-xvii + ix-xvi + 1 - 243, pls. 1 - 55 + A-Q, Circ. pl. 2]","Al-Maliky, T. H. Y., Al-Khafaji, K. K. & Khalaf, T. A. (2017) New records of the snapping shrimp Alpheus edwardsii (Audouin, 1826) (Crustacea: Alpheoidea) in Basrah, Iraq. Arthropods, 6, 47 - 53.","Audouin, V. (1826) Explication sommaire des planches de crustaces de l'Egypte et de la Syrie, publiees par Jules-Cesar Savigny, Membre de l'Institut; offrant un expos des characteres naturels des genres avec la distinction des especes. Animaux invertebres. In: Description de l'Egypte ou receuil des observations et des recherches qui ont ete faites en Egypte pendant l'expedition de l'armee francaise, publie par les ordres de sa Majeste l'Empereur Napoleon le Grand. Imprimerie Imperiale, Paris, pp. 77 - 98."]}

Alpheus inopinatus Holthuis & Gottlieb, 1958 (Figs. 1, 3 A–C) Alpheus inopinatus Holthuis & Gottlieb 1958: 42, figs. 8, 9; Lewinsohn & Holthuis 1964: 51; Tirmizi & Kazmi 1969: 99 (379), fig. 1a; Karim & Hafizur 1974: 26, pl. 4, figs. 1–4b; Banner & Banner 1978: 223; Kazmi & Kazmi 1979: 153; Barkati 1980: 355; Kocataş 1981: 162; Banner & Banner 1982: 241, fig. 75; Lewinsohn & Galil 1982: 208; Afzal et al. 1986: 339; Jeng 1997: 13; Cunha et al. 2020: 343, 351, fig 5C–D, 6C–D, 7C–D. Alpheus lobidens. — Jones 1986: 152, pl. 42 (partim ?); Geldiay 1969: 3, 5 (partim?); Kocataş & Kataĝan 2003: 65, 70 (partim ?); Dehghani et al. 2019a: 486, fig. 4A–D (partim ?) [not A. lobidens De Haan, 1849]. Alpheus sp. — Forest & Guinot 1958: 6, figs. 3–7 (see Banner & Banner 1982: 243). Material examined. Kuwait: 1 male, OUMNH. ZC.2020-01-001, Shaab, 29°21’16.78”N 48°01’47.38”E, mixed rocky-muddy intertidal, under rocks, leg. A. Anker, M. Al-Kandari, S. De Grave, Z. Sattari, 25 September 2019 [fcn KUW-011]; 1 male, OUMNH. ZC. 2020-01-002, same collection data as for previous specimen [fcn KUW-030]; 2 males, 1 female, OUMNH. ZC.2020-01-003, same collection data as for previous specimens [fcn KUW-034]; 1 male, OUMNH. ZC.2020-01-004, Failaka Island, eastern side, 29°23.409’N 48° 23.927’E, mixed rocky-muddy intertidal, under rocks, leg. M. Al-Kandari, 3 October 2019 [fcn KUW-077]; 1 male, 1 female, KISR, Al Julaia’a, Seashell Hotel, leg. M. Al-Kandari, 14 December 2019. Comparative material. Alpheus cf. lobidens De Haan, 1849. Kuwait: 1 male, 1 ovigerous female, OUMNH. ZC.2020-01-005, Ras Ajuza, 29°23.656’N 47°59.840’E, mixed rocky-muddy intertidal, under rocks, leg. M. Al-Kandari, Z. Sattari, 25 September 2019 [fcn KUW-027]; 1 female, OUMNH. ZC.2020-01-006, same collection data as for previous specimens [fcn KUW-029]; 1 male, OUMNH. ZC.2020-01-007, Shaab, 29°21’16.78”N 48°01’47.38”E, mixed rocky-muddy intertidal, under rocks, leg. A. Anker, M. Al-Kandari, S. De Grave, Z. Sattari, 25 September 2019 [fcn KUW-035]; 1 male, 1 ovigerous female, OUMNH. ZC.2020-01-008, same collection data as for previous specimen [fcn KUW-036]; 1 female, OUMNH. ZC.2020-01-009, Sulaibikhat Bay, 29°19.548’N 47°49.35’E, intertidal mud flat, in burrows, suction (yabby) pump, leg. A. Anker, M. Al-Kandari, Z. Sattari, 7 October 2019 [fcn KUW-107]; 1 male, 1 female, OUMNH. ZC.2020-01-010, same collection data as for previous specimen [fcn KUW-108]. Description. See Holthuis & Gottlieb (1958), Tirmizi & Kazmi (1969), Karim & Hafizur (1974) and Banner & Banner (1982). Colour pattern. Pale greenish or bluish grey to various tones of brown, often darker in females; pleon with three pairs of small dark spots distributed as following: one pair of small lateral spots on second pleuron, one pair of small dorsolateral spots on third pleuron (sometimes very faint and inconspicuous) and one pair of larger, more conspicuous lateral spots on fourth pleuron; mesial side of major and minor chelae greenish-brownish, with some olive or orange areas; dactylar plunger of major chela creamy white with some bluish tinge proximally; antennular and antennal flagella pale greenish-yellow; second pereiopods and walking legs (third to fifth pereiopods) pale reddish or pinkish with whitish articulations; tail fan brownish or greenish, with bluish area on uropodal exopod (Figs. 1, 3 A–C). General distribution. Possibly widely distributed in the Indo-West Pacific with records from the Red Sea, Djibouti, Arabian-Persian Gulf (Kuwait, Iran, see Remarks), Oman, Pakistan, Hong Kong, Taiwan, Australia (Tirmizi & Kazmi 1969; Banner & Banner 1978, 1981, 1982; Jeng 1997; present study); however, all records from outside of the western Indian Ocean require confirmation (see Remarks). Lessepsian migrant with a well-established population in the eastern Mediterranean Sea, with confirmed records from Israel and Turkey (Holthuis & Gottlieb 1958; Kocataş 1981; Lewinsohn & Galil 1982). Ecology. Largely intertidal, sometimes also in the shallow subtidal (to at least 6 m), rocky and mixed sandyrocky shores, often heavily silted or with a significant mud component; typically on sand/mud under large rocks and coral rubble (Lewinsohn & Galil 1982; present study). Remarks. Alpheus inopinatus belongs to the A. lobidens De Haan, 1849 species complex, together with A. crassimanus Heller, 1862 (currently considered to be a synonym of A. lobidens), A. lobidens polynesicus Banner & Banner, 1974 (originally described as A. lobidens polynesica), and the recently described A. mohammadpouri Dehghani, Sari & Naderloo, 2019 (Banner & Banner 1974, 1982; Dehghani et al. 2019b), together with a large number of undescribed species. The complicated taxonomic situation of A. lobidens and the presence of several cryptic or pseudocryptic species in this species complex make identifications of specimens extremely difficult (Anker & De Grave 2016). Alpheus inopinatus was originally described based on material from the Mediterranean coast of Israel, as an “immigrant from the Red Sea” (Holthuis & Gottlieb 1958). This species is morphologically very close to another species of the A. lobidens complex, which is present in the Gulf and possibly throughout the western and northern Indian Ocean (including the Red Sea), and identified either as A. lobidens (type locality: Japan) or A. crassimanus (type locality: Nicobar Islands, India). Like A. inopinatus, this species also penetrated into the Mediterranean Sea through the Suez Canal, spreading along the North African coasts from Egypt westwards to Tunisia, and northwards to Syria and Turkey (Hasan et al. 2008; Bakir et al. 2015). Older Mediterranean records of this species are under the name A. crassimanus (e.g., Balss 1936; Forest & Guinot 1956, 1958; Holthuis & Gottlieb 1958; Geldiay 1969), which was later placed in the synonymy of A. lobidens (see discussion in Banner & Banner 1974, 1981, 1982), with more recent records under the name A. lobidens (e.g., Bakir et al. 2015). In the present study, we tentatively continue to accept the synonymy of A. crassimanus with A. lobidens proposed by Banner & Banner (1974, 1982), who compared the Japanese specimens they identified as A. lobidens with the material from other areas of the Indo-West Pacific. However, we feel that a thorough morphological and molecular study is needed to establish the true identity of this taxon in light of the fact that their view of A. lobidens as a single widespread species has now been refuted (see also Cunha et al. 2020). The colour pattern of A. crassimanus is currently unknown, which is especially relevant in light of growing evidence of the presence of several cryptic or pseudocryptic species within the A. lobidens complex distinguishable mainly by colour patterns, for instance, in Singapore (Anker & De Grave 2016), Taiwan (A. Anker, pers. obs.), Japan (K. Nomura, pers. comm.), and Iran (Dehghani et al. 2019a). Forest & Guinot (1958) were aware of the presence of two morphologically distinct species from the A. lobidens complex on the Mediterranean coast of Israel, viz. A. inopinatus (reported as Alpheus sp., as it was already being studied by Holthuis & Gottlieb) and A. lobidens (reported by them as A. crassimanus). These authors also examined several specimens in Henri Coutière’s alpheid collection in the Muséum National d’Histoire Naturelle in Paris (MNHN), which morphologically corresponded well to A. inopinatus, viz. two specimens from the Gulf of Oman and a mixed lot from Djibouti, containing both A. inopinatus and A. lobidens. Interestingly, Holthuis & Gottlieb (1958) contrasted A. inopinatus with A. lobidens only superficially, comparing it with three more distantly related species of Alpheus, whereas Forest & Guinot (1958) provided a full account of the morphological differences between the two taxa (see below). Banner & Banner (1981, 1982) questioned the separation of A. inopinatus from A. lobidens and synonymised the former species. The Hawaiian subspecies, A. lobidens polynesicus, was also placed in the synonymy of A. lobidens, despite some rather striking colour pattern differences (Banner & Banner 1981). However, at least some of the synonymies proposed by Banner & Banner (1981, 1982) in the A. lobidens complex were premature (Anker 2001; Anker & De Grave 2016). New material collected throughout the Indo-West Pacific clearly shows that several features of the colour pattern, such as the presence or absence of dark transverse bands on the pleon, the presence or absence of dark spots on the sides of the pleon, the markings on the mesial side of the major and minor chelae and the colour of the dactylar plunger, may provide additional differentiating characters between the species of the A. lobidens complex (Anker & De Grave 2016; A. Anker, pers. obs., see also Fig. 3). De Grave & Fransen (2011) treated A. inopinatus as a valid species, but continued to list A. lobidens polynesicus as a junior synonym of A. lobidens. The present material confirms that A. inopinatus is clearly distinct from the west Indian Ocean taxon often recorded as A. lobidens and thus represents a valid species (see also Cunha et al. 2020 for molecular comparison). The common and widespread taxon from the western Indian Ocean (e.g., Red Sea, Gulf, Pakistan), which penetrated into the Mediterranean Sea most likely does not represent A. lobidens sensu De Haan (1849) from Japan and is herein referred to A. cf. lobidens (see Comparative material). Whether this taxon is identical to A. crassimanus sensu Heller (1862) remains to be shown. Alpheus inopinatus can be separated from A. cf. lobidens by at least three morphological characters, in addition to a small but consistent difference in their colour patterns. Both species can also be separated morphologically from A. lobidens polynesicus and A. mohammadpouri. The taxonomic status of these taxa will be discussed elsewhere, as part of a larger study of the A. lobidens complex. Many of the differentiating characters between A. inopinatus and A. cf. lobidens listed by Forest & Guinot (1958) were confirmed in the present material. The most important character separating the two species is the absence of a sharp distomesial tooth on the meri of the major and minor cheliped in the former species. In some specimens of A. inopinatus, the distomesial margin terminates in a subacute angle, sometimes protruding as a minute tooth, which is not the same structure as a much stronger acute tooth, situated slightly below the distomesial angle, in A. cf. lobidens (cf. Forest & Guinot 1958: figs. 1, 3, 4). The male minor cheliped has a much stouter merus and a stouter and more sculptured chela in A. inopinatus, compared to the noticeably slenderer merus and chela of A. cf. lobidens (cf. Forest & Guinot 1958: figs. 2, 5). The third and fourth pereiopods are also stouter in A. inopinatus, as expressed by the more inflated merus, with a length-width ratio of about 3.0 (cf. Forest & Guinot 1958: fig. 7) vs. more than 4.0 in A. cf. lobidens from Kuwait or the specimens from Australia identified as A. lobidens by Banner & Banner (1982: fig. 78f). In life, A. inopinatus and A. cf. lobidens can be separated by their colour patterns, although the difference is rather subtle. The pleon of A. inopinatus presents three pairs of small dark spots: one pair of lateral spots on the second pleuron, one pair of very small dorsolateral spots on the third pleuron and one pair of more conspicuous lateral spots on the fourth pleuron (Fig. 1); all these spots are absent in A. cf. lobidens (Fig. 2). The dactylar plunger of the major chela of both species is similar in colour, whitish with creamy or yellow tinge, although in A. inopinatus, the base of the plunger has a bluish tinge (Fig. 3C), whilst in A. cf. lobidens, the anterior surface has a more intense pale orange colour (Fig. 3F). The Iranian material of A. lobidens listed by Dehghani et al. (2019a) likely contains both A. inopinatus and A. cf. lobidens, based on an analysis of the colour patterns of the four illustrated individuals. Generally, many records under the name “ A. lobidens ” from the Gulf, Red Sea, Mediterranean Sea and other localities may refer to A. inopinatus as the two species have been commonly confused in the past, especially after the synonymisation of A. inopinatus by Banner & Banner (1981, 1982). Anker & De Grave (2016), while reporting A. lobidens (as A. lobidens sensu lato) from Singapore, provided colour photographs of three different colour morphs, possibly three different species, of the A. lobidens species complex. They can be briefly described as (1) pleon non-banded, with two conspicuous lateral black spots on the second and fourth pleura and two small medially fused black spots on the dorsal surface of the fifth pleonite (Anker & De Grave 2016: fig. 16A, E); (2) pleon banded, with two small dorsolateral black spots on the third pleuron, two very large and conspicuous black lateral spots on the fourth pleuron and two smaller black lateral patches on the fifth pleuron (idem.: fig. 16C, D); and (3) pleon non-banded, without spots (idem.: fig. 17). The arrangement of the spots of the Singaporean specimens appears to be different from that of A. inopinatus. In addition, the dactylar plunger of the major chela of the two black-spotted colour morphs from Singapore is bright orange, which makes them clearly different from both A. inopinatus and the specimens herein identified as A. cf. lobidens, both with whitish-creamy plungers (Fig. 3C, F). Therefore, all material identified as A. inopinatus from outside of the western Indian Ocean (Hong Kong, Taiwan, Australia) needs to be carefully re-examined. These observations also contribute to the remaining uncertainty about the taxonomic status of A. crassimanus, currently regarded as a synonym of A. lobidens (see above). Recently, Cunha et al. (2020) redescribed A. lobidens from Nagasaki on the basis of what they considered to be topotypic specimens, collected from the vicinity of Nagasaki. However, the type locality of A. lobidens is Japan, i.e., the entire Japanese archipelago and not specifically Nagasaki (De Grave & Fransen 2011: 385). Only six species described in De Haan (1833 –1850) actually have a type locality that is geographically more restricted than merely Japan (see Yamagushi & Baba 1993), so it cannot be assumed that the holotype of A. lobidens was indeed collected in the Nagasaki area. Furthermore, it is well known that the botanical and zoological collections gathered by Von Siebold and Bürger, a part of which was studied by De Haan, came from all over Japan. For example, the type series of the giant spider crab Macrocheira kaempferi (Temminck) was collected in Suruga Province, further east, whereas their collection also contained cold-water decapods from much further north, e.g., Lithodes japonicus Von Siebold, now known as Paralithodes camtschaticus (Tilesius). The authors simply “…assumed that the topotypes from Shimabara, Nagasaki, Japan (RMNH PEG 25134) correspond to A. lobidens sensu stricto, i.e., are conspecifics of the damaged holotype …”, without a direct comparison to the holotype. Although it is possible that Cunha et al. (2020) did redescribe what in future could be considered as true A. lobidens (i.e. A. lobidens sensu De Haan, 1849), it should be noted that numerous species of the A. lobidens complex occur in Japan, an observation based primarily on the diversity of colour patterns (K. Nomura, pers. comm.). As no colour pattern was documented for A. lobidens sensu Cunha et al. (20202), any further comparison with other taxa is hindered, as it is obvious now that colour patterns, along with molecular analyses and morphology, will be crucial to resolve this challenging species complex. For example, in Singapore (Anker & De Grave 2016), Kuwait (present study) and some localities in eastern Australia (A. Anker, pers. obs.), two or more species of the A. lobidens complex occur syntopically, with the easiest separation between them being consistent differences in the colour patterns. Al-Maliky et al. ( 2017) reported and illustrated Alpheus edwardsii (Audouin, 1826) from Basrah area in Iraq. Although their photographs of preserved specimens are in very low resolution, the dorsal shoulder of the major chela is not overhanging the adjacent transverse groove (idem: figs. 2, 3A), whilst the merus of the major cheliped seems to be armed with a strong distomesial tooth (idem: fig. 3B, D). Therefore, the Iraqi material can be tentatively referred to A. cf. lobidens, as suggested by Al-Khandari et al. (2020: p. 254; mind a small error therein, read correctly: “ Al-Maliky et al. (2017), who erroneously reported A. lobidens as A. edwardsii from Shatt-Al-Basrah and Faw in Iraq.”).

Alpheus alaincrosnieri n. sp. Figs. 1���4 Alpheus acutocarinatus ��� Chace 1988: 15 (part., specimen from Masbate) [not A. acutocarinatus De Man 1909]. (?) Alpheus miyakei Miya 1974: 106; Bruce 1994: 23 (key); De Grave & Fransen 2011: 476 (listed as Nomen nudum). Type material. Holotype: male, cl 7.6 mm, MNHN-IU-2011-2230, Papua New Guinea, off Sepik River, Campaign BIOPAPUA, R / V ���Alis���, Sta. CP 3701, 03��57���S 144��41���E, depth 198���219 m, leg. S. Samadi & L. Corbari, 01.10.2010. Paratypes: 1 male, cl 6.5 mm, 2 females, cl 6.8, 7.2 mm (missing both P1 and major P1, respectively, both with bopyrid isopod parasites), MNHN-IU-2019-3168, same collection data as for holotype; male, cl 7.0 mm, MNHN-2011-3024, Papua New Guinea, off New Ireland, Campaign BIOPAPUA, R / V ���Alis���, Sta. DW 3764, 04��39���S 153��03���E, depth 220 m, leg. S. Samadi & L. Corbari, 15.10.2010. Additional material. 1 male, cl 7.4 mm (missing most legs, including both P1), MNHN-IU-2017-1947, Vanuatu, Espiritu Santo Island, inner Big Bay, Campaign BOA 1, R / V ���Alis���, Sta. CP 2448, 15��06���60���S 166��50���80���E, depth 297���387 m, leg. S. Samadi / IRD et al., 10.09.2005; 1 ovigerous female, cl 8.5 mm, MNHN-IU-2010-1278, Madagascar, northwest coast, Mozambique Channel off Narinda (���Nazendry���) Bay, Campaign MIRIKY, Sta. CP 3293, 14��30���S 47��26���E, depth 268���408 m, leg. P. Bouchet et al., 14.07.2009. Description. Medium-sized (present material: cl 6.5���8.5 mm) species of Alpheus brevirostris group. Carapace glabrous, neither setose nor pubescent. Rostrum well developed, slender, about twice as long as wide at base, subacute distally, reaching mid-length of first article of antennular peduncle, somewhat ascendant in lateral view; rostral carina strong, sharp, continuing on carapace well past orbital hoods, ending in conspicuous post-rostral tubercle in form of anteriorly elevated, gradually flattening tooth (Fig. 1a, b). Orbital hoods swollen, somewhat projecting anteriorly in lateral view; anterior margin of each orbital hood armed with small sharp tooth; frontal margin between rostrum and orbital hood shallowly concave; adrostral furrows rather shallow, distinct mainly in anterior portion of orbital hoods, between eyes (Fig. 1a, b). Pterygostomial angle protruding anteriorly as stout sharp tooth (Fig. 1b); cardiac notch well developed, deep. Eyes well developed, with large, normally pigmented corneas (Fig. 1a, b). Telson narrow, subrectangular, tapering distally, about 2.7 times as long as maximal width, with lateral margins distinctly constricted at about telson mid-length; dorsal surface with two pairs of stout cuspidate setae both inserted at some distance from lateral margin, first pair at about 0.4 telson length, second pair at about 0.7 of telson length; posterior margin broadly rounded, with row of shorter slender setae above long plumose setae; posterolateral angles each with one pair of spiniform setae, mesial ones stouter and more than twice as long as lateral ones (Fig. 1c, d). Antennular peduncle elongate, slender; stylocerite with basal part broad, swollen laterally, distal part slender, ending in sharp point, latter falling short of distal margin of first article; ventromesial carina with large tooth (not illustrated but visible in Fig. 1b); second article elongate, almost five times as long as wide; lateral antennular flagellum with secondary ramus fused to main ramus over most of its length, with numerous groups of aesthetascs distally, starting from about 14th subdivision (Fig. 1a, b, e). Antenna with basicerite very stout, its distoventral margin armed with very large, anteriorly projecting, sharp tooth; scaphocerite well developed, with straight lateral margin, moderately broad blade and strong distolateral tooth, latter reaching far beyond distal margin of blade, but not reaching end of antennular peduncle; carpocerite reaching slightly beyond scaphocerite, also not reaching end of antennular peduncle (Fig. 1a, b). Mouthparts generally typical for genus in external observation. Mandible with incisor process bearing nine teeth. First maxilliped with well-developed caridean lobe and endopod subdivided into two articles (Fig. 1f). Second maxilliped with third article of endopod armed with small sharp tooth on mesial (ventral) margin (Fig. 1g, h). Third maxilliped slender, elongate, very setose; coxa with bluntly projecting lateral plate; antepenultimate article flattened ventrolaterally, with distinct ridge running parallel to dorsal margin on lateral surface, mesial margin slightly rugose; penultimate article relatively long, about 3.5 times as long as wide proximally, distoventral surface noticeably bulging distally, strongly convex, furnished with very long stiff setae; ultimate article unarmed distally, with numerous rows of shorter serrulate setae and much longer stiff setae, some of more apically situated setae extremely elongate, much longer than ultimate article; arthrobranch moderately developed (Fig. 1i). Major cheliped not sexually dimorphic, slightly weaker in smaller females. Male major cheliped more robust, but only slightly longer than minor cheliped; ischium short, stout, ventrolateral surface with small granules, ventromesial margin with at least two elevations each carrying slender spiniform seta; merus elongate, trigonal in cross-section, about four times as long as wide at base, patchily covered by small granules, distodorsal margin blunt, without distal or subdistal tooth, ventromesial margin rugose, armed with two or three slender spiniform setae in proximal third, with stout sharp distal tooth; carpus subcylindrical, short, distally widening, cup-shaped in larger individuals, with more or less protruding blunt distodorsal process; chela extremely elongate, slender; palm strongly compressed, subrectangular in cross-section, length / height ratio ranging from about 4.2 in smaller individuals to over 5.0 in larger individuals, ventral margin broadly concave distally, forming shallow sinus, without deep constriction, dorsal margin without transverse groove or notch subdistally, large sections of palmar surface covered by small granules; fingers unequal in length, with dactylus noticeably shorter than pollex, about 0.25���0.3 times (dactylus) to 0.3���0.35 times (pollex) length of palm, not twisted or significantly deviating from chela axis, largely gaping proximally when closed; dactylus distally rounded, plunger highly reduced, continuous with dactylar cutting edge anteriorly, abruptly delimited posteriorly; adhesive disks small; distodorsal surface of merus, ventral and dorsal margins of palm, and ventral margin of pollex fringed with long setae, those of pollex projecting forward, often forming conspicuous tuft of several parallel setae (Fig. 2 a���f). Minor cheliped sexually dimorphic, balaeniceps in males, simple in females (Fig. 3). Male minor cheliped with ischium short, stout, its ventromesial margin rugose, with at least two slender spiniform setae; merus elongate, trigonal in cross-section, about 4.5 times as long as wide at base, patchily covered by small granules, some organised in row on ventral surface, especially in larger individuals, distodorsal margin blunt, without distal or subdistal tooth, ventromesial margin rugose, armed with four or so slender spiniform setae in proximal third, with stout sharp distal tooth; carpus subcylindrical, short, distally widening, cup-shaped in larger individuals, with protruding blunt distodorsal process, dorsal and ventral surfaces more or less rugose or covered with small granules; palm compressed, subrectangular in cross-section, without grooves or notches, length / height ratio ranging from about 5.5 in smaller individuals to over 6.5 in larger individuals, surface slightly rugose in smaller individuals, covered with small granules in addition to rugosities in larger individuals; fingers about 0.6���0.7 length of palm, not twisted, slightly gaping when closed, subequal in length, strongly crossing distally; dactylus more or less flattened dorsoventrally and with margins slightly broadened, strongly bent distally, with two balaeniceps ridges, one on mesial side and one on lateral side, each furnished with densely inserted, thickened, plumose setae; both balaeniceps ridges extending from about half-length of dactylus to point of its curvature; pollex more gradually curving distally, without balaeniceps setae on either side; proximal surface of both dactylus and pollex covered with small granules; adhesive disks reduced; dorsal margin of palm, and ventral margin of pollex fringed with long setae, those of pollex projecting forward, often as conspicuous tuft of several parallel setae (Fig. 3 a���d). Female minor cheliped more slender than male minor cheliped; carpus longer, vase-shaped, with rugosities or granules and elongate setae; chela with fingers about 0.8 length of palm, slightly gaping; dactylus not expanded, simple, without balaeniceps setae; ventral and dorsal margins of palm, ventral margin of pollex and dorsal margin of dactylus fringed with long setae (Fig. 3e). Second pereiopod very slender, especially in larger individuals; ischium and merus subequal in length; carpus with five subdivisions, first longest, ratio of carpal subdivisions approximately equal to 3.0/2.2/1/0.9/1.2; chela longer than distal-most carpal subdivision (Fig. 4a, b). Third pereiopod generally slender; ischium with stout cuspidate seta on ventrolateral surface; merus slender, about 10.5 times as long as maximal width, unarmed distoventrally; carpus about 0.6 length of merus, much slenderer than merus, unarmed; propodus noticeably longer than carpus, with numerous long stiff setae, ventral margin without spiniform setae; dactylus about half-length of propodus, gradually curving distally, spatulate, broadened, flattened ventrally, with distinct longitudinal keel (Fig. 4 c���e). Fourth pereiopod generally similar to third, but slenderer; ischium longer than in third pereiopod, with stout cuspidate seta on ventrolateral surface; merus about 11.4 times as long as maximal width; propodus and dactylus as in third pereiopod (Fig. 4f, g). Fifth pereiopod much slenderer than third pereiopod and slightly slenderer than fourth pereiopod; ischium longer than in third and fourth pereiopods, unarmed; merus about 10.3 times as long as wide; carpus slenderer than merus, about 0.8 length of merus; propodus longer than carpus, with numerous long stiff setae, ventral margin without spiniform setae, distal ventrolateral surface with rows of serrulate setae forming cleaning brush; dactylus similar to that of third and fourth pereiopods (Fig. 4h, i). Male second pleopod with appendix masculina slightly longer than appendix interna, with long stiff setae on apex and along margins (Fig. 1j). Uropod with both mesial and lateral lobes of protopod ending bluntly; exopod fairly broad, somewhat truncate distally, with short triangular distolateral tooth; diaeresis straight for most part, except for blunt lobe adjacent to slender distolateral spiniform seta, latter reaching level of distal margin of exopod; endopod much narrower than exopod; neither exopod nor endopod with spiniform setae on their distal margins (Fig. 1k). Colour pattern presently unknown. Etymology. This spectacular deep-water species is named after Dr. Alain Crosnier (formerly at ORSTOM / MNHN), in recognition of his invaluable contribution to taxonomy of various groups of Decapoda, including the presently most complete monograph of the Alpheidae of the tropical eastern Atlantic (Crosnier & Forest 1966). Distribution. Indo-West Pacific: Papua New Guinea (off Sepik River, New Ireland), Vanuatu (Espiritu Santo Island) and Madagascar (off Narinda Bay), at depths of 198��� 408 m. Remarks. Alpheus alaincrosnieri n. sp. clearly belongs to the A. brevirostris group based on the overall shape of the chelipeds. However, this new species is unique within the A. brevirostris group by the presenting two small, but well-developed, acute orbital teeth, projecting from the anterior margin of each orbital hood (Fig. 1a, b). In addition, the pterygostomial angle of A. alaincrosnieri n. sp. is strongly projecting anteriorly in form of a stout sharp tooth (Fig. 1b), a feature not commonly seen in the genus Alpheus. Within the A. brevirostris group, only A. kagoshimanus Hayashi & Nagata, 2000 has a pterygostomial tooth, which, however, is much smaller than in the present new species.All other species of the A. brevirostris group characterised by the significantly elongated chelipeds, viz. A. macroskeles Alcock & Anderson, 1894, A. talismani Couti��re, 1898, A. acutocarinatus De Man, 1909, A. notabilis Stebbing, 1915, A. explorator Boone, 1935, A. migrans Lewinsohn & Holthuis, 1978, A. nonalter Kensley, 1969, and A. longipalma Komai & Ohtomi, 2018, can be easily separated from A. alaincrosnieri n. sp. both by the rounded orbital hoods and rounded to slightly angular pterygostomial angle. Alpheus alaincrosnieri n. sp. can be separated from each of these species by at least one additional, taxonomically significant character, for instance, from A. kagoshimanus by the much stronger distolateral tooth of the scaphocerite, distinctly overreaching the distal margin of the blade (cf. Hayashi & Nagata 2000); from A. kagoshimanus, A. notabilis and A. migrans (and possibly from most if not all other species) by the presence of a sharp tooth on the third article of the endopod of the second maxilliped (cf. Lewinsohn & Holthuis 1978; Hayashi & Nagata 2000); from A. macroskeles, A. talismani and A. longipalma by the cheliped merus without subdistal dorsal tooth (cf. Alcock & Anderson 1894; Couti��re 1898; Alcock 1901; Crosnier & Forest 1966; Komai & Ohtomi 2018); from A. macroskeles and A. talismani by the minor chela strongly balaeniceps in males (cf. Alcock & Anderson 1894; Couti��re 1898; Alcock 1901; Crosnier & Forest 1966); from A. macroskeles by the better developed cornea of eyes (cf. Alcock & Anderson 1894; Alcock 1901; Komai & Ohtomi 2018); from A. talismani by the presence of an elevated rostral carina armed with a post-rostral tubercle (cf. Couti��re 1898; Crosnier & Forest 1966); from A. acutocarinatus by the third pereiopod ischium armed with a stout cuspidate seta (cf. De Man 1911); from A. acutocarinatus and A. migrans by the absence of small tubercle on the mid-dorsal line of the posterior region of the carapace and the second pereiopod with the first carpal subdivision longer than the second (cf. De Man 1911; Lewinsohn & Holthuis 1978); from A. notabilis by the broader and less constricted telson and the scaphocerite with a straight (not noticeably concave, as in A. notabilis) lateral margin (cf. Stebbing 1915); from both A. explorator and A. nonalter by the major chela palm without a deep constriction near the base of the pollex (Boone 1935; Kensley 1969; Chace 1988; Komai 2011); from A. explorator by the position of the ventromesial tooth on the cheliped merus (distal in the new species vs. clearly subdistal in A. explorator) (Boone 1935; A. Anker, in prep.); and from A. migrans by the much longer penultimate article of the third maxilliped and the lower, more reduced plunger of the major chela (cf. Lewinsohn & Holthuis 1978; Dworschak & Pervesler 2002). It must be noted that A. notabilis, described by Stebbing (1915) on the basis of a heavily damaged specimen missing its major cheliped, with somewhat diagrammatic and possibly not very accurate illustrations, has not been reported since the original description (Banner & Banner 1983). Chace (1988) listed a mutilated ovigerous female (cl 8.3 mm) collected at a depth of 245 m near Masbate, Philippines, in the remarks under A. acutocarinatus. However, the author���s notes show that he was well aware that it did not belong to that species. Based on Chace���s (1988) observations, the specimen from Masbate had ���the remnant of what must have been a larger mesial gastric tooth [= post-rostral tubercle], by a small spine on the frontal margin either side of the rostrum [= orbital teeth], and by a rather distinct branchiostegal spine [= pterygostomial tooth]���, which are the most diagnostic features of the carapace of A. alaincrosnieri n. sp. Therefore, Chace���s (1988) specimen is here tentatively referred to the new species, extending its distribution to the Philippines. Miya (1974), while providing short diagnoses of the seven species groups of Alpheus, mentioned under the A. brevirostris group ��� A. miyakei sp. nov. which will be described in Part III���. However, Part III of Miya���s monograph of the Alpheidae of Japan was never published and Miya���s name is therefore a Nomen nudum (De Grave & Fransen 2011). It remains unknown from which locality (or localities) Miya���s material came from and if it was collected in shallow or in deep water. The only character mentioned by Miya (1974) for A. miyakei was the presence of orbital teeth, which was used by Bruce (1994) in the first couplet of his key to the A. brevirostris group. Miya���s invalid name is here tentatively listed in the synonymy of A. alaincrosnieri n. sp., although its true identity may never be uncovered., Published as part of Anker, Arthur, 2020, On two new deep-water snapping shrimps from the Indo-West Pacific (Decapoda Alpheidae: Alpheus), pp. 393-409 in Zootaxa 4845 (3) on pages 394-396, DOI: 10.11646/zootaxa.4845.3.5, http://zenodo.org/record/4406729, {"references":["Chace, F. A. Jr. (1988) The caridean shrimps (Crustacea: Decapoda) of the Albatross Philippine Expedition, 1907 - 1910, Part 5: Family Alpheidae. Smithsonian Contributions to Zoology, 466, 1 - 99. https: // doi. org / 10.5479 / si. 19436696.391.1","De Man, J. G. (1909) Diagnoses of new species of macrurous decapod Crustacea from the \" Siboga-Expedition \". Tijdschrift der Nederlandse Dierkundige Vereeniging, 2 (9), 99 - 125.","Miya, Y. (1974) The Alpheidae (Crustacea, Decapoda) of Japan and its adjacent waters. Part II. Publications from the Amakusa Marine Biology Laboratory, 3, 103 - 195, pls. 15 - 31.","Bruce, A. J. (1994) Alpheus fenneri sp. nov. and A. williamsi sp. nov., two new Indo-West Pacific alpheid shrimps of the brevirostris species group. The Beagle, Records of the Museums and Art Galleries of the Northern Territory, 11, 15 - 28.","De Grave, S. & Fransen, C. H. J. M. (2011) Carideorum catalogus: the recent species of the dendrobranchiate, stenopodidean, procarididean and caridean shrimps (Crustacea: Decapoda). Zoologische Mededelingen, Leiden, 85, 195 - 589.","Crosnier, A. & Forest, J. (1966) Crustaces decapodes: Alpheidae. In: Campagnes de la Calypso dans le Golfe de Guinee et aux Iles Principe, Sao Tome et Annobon (1956), et Campagne aux Iles du Cap Vert (1959). Part 19. Resultats Scientifiques des Campagnes de la Calypso 7 (27). Annales de l'Institut Oceanographique de Monaco, 44, 199 - 314.","Hayashi, K. - I. & Nagata, M. (2000) A new species of Alpheus (Decapoda, Caridea, Alpheidae) from Kagoshima Bay, Japan. Crustaceana, 73, 1109 - 1120. https: // doi. org / 10.1163 / 156854000505119","Alcock, A. & Anderson, A. R. S. (1894) Natural history notes from H. M. Royal Indian marine survey steamer \" Investigator \", Commander C. F. Oldham, R. N., commanding. Series II, no. 14. An account of a recent collection of deep sea Crustacea from the Bay of Bengal and Laccadive Sea. Journal of the Asiatic Society of Bengal, 63, 141 - 185, pl. 9.","Coutiere, H. (1898) Note sur Alpheus Talismani n. sp. et A. macroskeles (Alcock et Anderson) [Crust.]. Bulletin de la Societe Entomologique de France, 1898, 31 - 33. https: // doi. org / 10.5962 / bhl. part. 16849","Stebbing, T. R. R. (1915) South African Crustacea. Part VIII of S. A. Crustacea, for the marine investigations in South Africa. Annals of the South African Museum, 15, 57 - 103, pls. 13 - 25. https: // doi. org / 10.5962 / bhl. part. 22195","Boone, L. (1935) The Crustacea: Anomura, Macrura, Euphausiacea, Isopoda, Amphipoda and Echinodermata: Asteroidea and Echinoidea of the \" Alva \" world cruise, 1931, William K. Vanderbilt, commanding. Bulletin of the Vanderbilt Marine Museum, 6, 1 - 264, pls. 1 - 96.","Lewinsohn, C. & Holthuis, L. B. (1978) On a new species of Alpheus (Crustacea Decapoda, Natantia) from the eastern Mediterranean. Zoologische Mededelingen, Leiden, 53, 75 - 82.","Kensley, B. (1969) Decapod Crustacea from the south-west Indian Ocean. Annals of the South African Museum, 52, 149 - 181.","Komai, T. & Ohtomi, J. (2018) A new deep-sea species of the snapping shrimp genus Alpheus Fabricius, 1798 (Decapoda: Caridea: Alpheidae) from Kagoshima Bay, Japan. Zootaxa, 4434 (1), 099 - 110. https: // doi. org / 10.11646 / zootaxa. 4434.1.6","Alcock, A. (1901) A descriptive catalogue of the Indian deep-sea Crustacea Decapoda Macrura and Anomala, in the Indian Museum. Being a revised account of the deep-sea species collected by the Royal Indian marine survey ship Investigator. Indian Museum, Calcutta, iv + 286 pp., 3 pls. https: // doi. org / 10.5962 / bhl. title. 30840","De Man, J. G. (1911) The Decapoda of the Siboga Expedition. Part II. Family Alpheidae. In: Siboga Expeditie. 39 a 1. Brill, Leiden, pp. 133 - 465; (1915) Supplement-Explanation of plates of Alpheidae, pls. 1 - 23.","Komai, T. (2011) Deep-sea shrimps and lobsters (Crustacea: Decapoda: Dendrobranchiata and Pleocyemata) from the Sagami Sea and Izu Islands, Central Japan. Memoirs of the National Museum of Nature and Science, 47, 279 - 337.","Dworschak, P. C. & Pervesler, P. (2002) Alpheus migrans Lewinsohn & Holthuis, 1978 (Decapoda, Caridea, Alpheidae): burrow morphology and first record from the Red Sea. Crustaceana, 75, 351 - 357. https: // doi. org / 10.1163 / 156854002760095426","Banner, A. H. & Banner, D. M. (1983) An annotated checklist of the alpheid shrimp from the western Indian Ocean. Travaux et Documents de l'ORSTOM, 158, 1 - 164."]}

Alpheus vanuatu n. sp. Figs. 5���8 Type material. Holotype: male, cl 6.0 mm, MNHN-IU-2017-1953, Vanuatu, Espiritu Santo Island, inner Big Bay, Campaign BOA 1, R / V ���Alis���, Sta. CP 2445, 15��08���0���S 166��53���3���E, depth 231���285 m, leg. S. Samadi / IRD et al., 10.09.2005. Paratypes: 1 female, cl 4.8 mm, MNHN-IU-2019-3169, same collection data as for holotype; 1 male, cl 4.2 mm, MNHN-IU-2017-1952, Vanuatu, Espiritu Santo Island, inner Big Bay, Campaign BOA 1, R / V ���Alis���, Sta. CP 2444, 15��07���80���S 166��53���70���E, depth 250���331 m, leg. S. Samadi / IRD et al., 10.09.2005; 4 males, cl 3.5, 3.5, 4.0, 4.5 mm, 2 females, cl 3.6, 3.8 mm, 1 ovigerous female, cl 4.0 mm, MNHN-IU-2017-1955, same collection data as for previous paratype. Description. Small to medium-sized (present material: cl 3.5���6.0 mm) species of Alpheus. Carapace glabrous, neither setose nor pubescent. Rostrum well developed, moderately slender, about twice as long as wide at base, subacute distally, reaching mid-length of first article of antennular peduncle, slightly descendant in lateral view; rostral carina strong, rounded, continuing and noticeably widening and flattening between orbital hoods, abruptly delimited from adrostral furrows, not extending past orbital hoods, disappearing posterior to level of eyes (Fig. 5a, b). Orbital hoods swollen, slightly projecting anteriorly in lateral view, with rounded anterior margin; frontal margin between rostrum and orbital hood very shallowly concave; adrostral furrows shallow, distinct mainly in anterior portion of orbital hoods, between eyes (Fig. 5a, b). Pterygostomial angle broadly rounded (Fig. 5b); cardiac notch well developed, deep. Eyes well developed, with large, normally pigmented corneas (Fig. 5a, b). Telson broad, subrectangular, tapering distally, about 2.0���2.2 times as long as maximal width, with lateral margins slightly concave posterior to telson mid-length; dorsal surface typically with two pairs of stout cuspidate setae both inserted far from lateral margin, first pair at about telson mid-length, second pair at about 0.7���0.8 of telson length, rarely with several cuspidate missing (as in one paratype); posterior margin broadly rounded; posterolateral angles each with one pair of slender spiniform setae, mesial ones much stouter and almost three times as long as lateral ones (Fig. 5 c���e). Antennular peduncle moderately stout; stylocerite with basal part moderately broad, slightly swollen laterally, distal part slender, ending in sharp point, latter slightly exceeding distal margin of first article; ventromesial carina with large, anteriorly directed tooth; second article about twice times as long as wide; lateral antennular flagellum with secondary ramus fused to main ramus over most of its length, with numerous groups of aesthetascs distally starting from about eighth subdivision (Fig. 5a, b, f). Antenna with basicerite stout, its distoventral margin armed with anteriorly projecting, slender, sharp tooth; scaphocerite well developed, with straight to faintly concave lateral margin, broad blade and very strong distolateral tooth, latter reaching far beyond distal margin of blade, also typically reaching or slightly overreaching end of antennular peduncle (distolateral tooth of left scaphocerite curved and shorter, slightly abnormal, in illustrated specimen); carpocerite reaching well beyond scaphocerite and end of antennular peduncle (Fig. 1a, b). Mouthparts typical for genus in external observation. Mandible with incisor process bearing about 10 teeth. Third maxilliped relatively stout, moderately setose; coxa with subacutely projecting lateral plate; antepenultimate article flattened ventrolaterally, with distinct ridge running parallel to dorsal margin on lateral surface, mesial margin slightly rugose; penultimate article relatively short, distally widening, at most twice as long as maximal width; ultimate article unarmed distally, with numerous rows of shorter serrulate setae and much longer stiff setae, especially on and near apex (Fig. 5g). Major cheliped feebly sexually dimorphic, weaker in females. Male major cheliped much more robust and longer than minor cheliped; ischium short, stout, smooth, unarmed; merus moderately stout, subtrigonal in crosssection, less than three times as long as wide at base, smooth, distodorsal margin bluntly protruding, without strong tooth, ventromesial margin smooth proximally, faintly rugose distally, with very stout sharp subdistal tooth; carpus very short, cup-shaped, smooth; chela slightly elongate; palm not noticeably compressed, subcylindrical in crosssection, length / height ratio more or less equal to 2.0, without grooves, notches, smooth (no granules), ventral margin faintly concave distally, forming shallow sinus, dorsal margin somewhat convex near mid-length; fingers subequal in length, about half-length of palm, slightly twisted laterally, gaping proximally when closed in lateral view; dactylus distally rounded, plunger somewhat reduced, continuous with dactylar cutting edge anteriorly, abruptly delimited posteriorly; adhesive disks large (Fig. 6 a���f). Minor cheliped sexually dimorphic, balaeniceps in males, simple in females (Fig. 7). Male minor cheliped with ischium short, smooth; merus moderately slender, subtrigonal in cross-section, about four times as long as wide at base, smooth, distodorsal margin blunt, ventromesial margin smooth over most of its length, with stout sharp subdistal tooth; carpus short, cup-shaped, smooth; palm slender, subcylindrical in cross-section, smooth, without grooves or notches, length / height ratio about 2.5; fingers about 0.9 length of palm, slightly twisted, gaping when closed, subequal in length, strongly crossing distally; dactylus strongly flattened ventrodorsally, with lateral and mesial margins expanded, strongly curved distally, with two balaeniceps ridges, one on mesial side and one on lateral side, each furnished with densely inserted, thickened, plumose setae; both balaeniceps ridges extending from about 0.4 to 0.7 of dactylar length; mesial and lateral sides of pollex each with row of balaeniceps setae on proximal half; adhesive disks modestly developed (Fig. 7a, b). Female minor cheliped smaller and more slender than male minor cheliped; chela with fingers slightly longer than palm, not gaping when closed; dactylus not expanded, simple, without balaeniceps setae (Fig. 7c, d). Second pereiopod slender; ischium and merus subequal in length; carpus with five subdivisions, first longest, ratio of carpal subdivisions approximately equal to 3.5/2/1/1/1.6; chela longer than distal-most carpal subdivision (Fig. 8a). Third pereiopod moderately slender; ischium with stout cuspidate seta on ventrolateral surface; merus about 6.5 times as long as maximal width, unarmed distoventrally; carpus about half-length of merus, slightly slenderer than merus, unarmed; propodus much longer than carpus, with numerous (some elongate) stiff setae, ventral margin with five or six smaller or larger spiniform setae, in addition to one pair of longer slender spiniform setae near propodo-dactylar articulation; dactylus slightly more than half-length of propodus, faintly curving distally, subconical, noticeably flattened ventrally (Fig. 8b, c). Fourth pereiopod generally similar to third, somewhat slenderer. Fifth pereiopod much slenderer than third pereiopod; ischium longer than in third pereiopod, unarmed; merus slender, almost seven times as long as wide; carpus slenderer than merus, about 0.8 length of merus; propodus much longer than carpus, distal ventrolateral surface with rows of serrulate setae forming cleaning brush, ventromesial margin with about six slender spiniform setae, including longest one near propodo-dactylar articulation; dactylus similar to that of third and fourth pereiopods (Fig. 8d, e). Male second pleopod with appendix masculina shorter than appendix interna, with long stiff setae on apex and along margins (Fig. 5h). Uropod with both mesial and lateral lobes of protopod ending in sharp tooth; exopod broad, broadly rounded distally, with stout triangular distolateral tooth; diaeresis sinuous for most part, with two rounded lobes in its lateral section, one of them adjacent to slender distolateral spiniform seta, latter not reaching level of distal margin of exopod; endopod much narrower than exopod, ovate; neither exopod nor endopod with spiniform setae on their distal margins (Fig. 5i). Colour pattern unknown. Etymology. This interesting new species is named after the country of the type locality, the Republic of Vanuatu; used as a noun in apposition. Distribution. West Pacific: presently known only from Vanuatu (Espiritu Santo Island), at depths of 231��� 331 m. Remarks. Alpheus vanuatu n. sp. appears to be morphologically closest to the pantropical A. paracrinitus Miers, 1881 species complex, which contains species found mostly in waters shallower than 50 m (A. Anker, unpublished data). The assignment of A. paracrinitus and related taxa to the A. diadema Dana, 1852 group dates back to Couti��re (1905) and was followed by most subsequent workers (e.g. Miya 1974; Banner & Banner 1982; Chace 1988; Kim & Abele 1988). However, the A. diadema group sensu Banner and Banner (1982) is rather vaguely defined and contains morphologically diverse taxa, including several species that are obviously not closely related to the remaining species traditionally assigned to this group. Therefore, the present author does not recognise the A. paracrinitus complex as part of the A. diadema group. In fact, it seems most reasonable to treat this morphologically well-defined species complex as a separate species group of Alpheus, hereafter the A. paracrinitus species group. The A. paracrinitus group can be defined by the following combination of morphological characters: (1) rostrum present, often small; (2) rostral carina, if present, not flattened and abruptly delimited from adrostral furrows, rostro-orbital region, including orbital hoods, without teeth; (3) third maxilliped with antepenultimate article often projecting distodorsally; (4) cheliped merus more or less slender, ventromesial margin with strong distal or subdistal tooth; (5) major chela subcylindrical, smooth, without grooves or notches; (6) minor cheliped simple or balaeniceps in males, simple in females; (7) third pereiopod merus lacking distoventral tooth and spiniform setae on ventral margin; and (8) third pereiopod dactylus typically simple, not strongly spatulate or biunguiculate. The following species can be presently assigned to the A. paracrinitus group: A. paracrinitus Miers, 1881 (currently regarded as pantropical species, but probably restricted to the East and West Atlantic, A. Anker, unpublished data); A. rostratus Kim & Abele, 1988 (East Pacific); A. alpheopsides Couti��re, 1905; A. paralpheopsides Couti��re, 1905; A. tenuipes De Man, 1910; A. labis Banner & Banner, 1982; A. mitis Dana, 1852, and A. vanuatu n. sp. (all from the Indo-West Pacific). Two junior synonyms of A. paracrinitus will need to be resurrected, in addition to descriptions of possibly two cryptic or pseudocryptic species in the A. paracrinitus complex sensu stricto. The taxonomic identity of A. mitis remains problematic (Banner & Banner 1983), although the species seems to be distinguishable from both A. paracrinitus sensu Miers (1881) and A. paracrinitus sensu Banner & Banner (1982). The recent record of A. mitis from Iran (Ashrafi et al. 2020) has to be treated with caution. Thus, the A. paracrinitus group, when fully revised, may include at least 12 species (A. Anker, under study). Apart from A. vanuatu n. sp., all the species of the A. paracrinitus group are distributed in shallow waters, although the habitat and depth of the single known specimen of A. labis are unknown (Banner & Banner 1982). Alpheus vanuatu n. sp. can be easily separated from all of the species listed above by the strongly balaeniceps male major chela, with the dactylus strongly flattened and expanded laterally and mesially (Fig. 7a, b) (cf. Couti��re 1905; De Man 1911; Banner & Banner 1982; Kim & Abele 1988). The only species of the A. paracrinitus group, in which the minor cheliped is feebly balaeniceps, is A. paracrinitus sensu Banner & Banner (1982: fig. 36d). However, this taxon (currently under study) differs from A. vanuatu n. sp. in several other morphological features (see below). The new species also differs from A. paracrinitus sensu Miers (1881) and sensu Banner & Banner (1982), as well as A. alpheopsides, A. paralpheopsides, A. rostratus, A. labis, A. mitis by the presence of a strong, posteriorly extending rostral carina, which is not distinct in the other species (cf. Couti��re 1905; Banner & Banner 1982; Kim & Abele 1988). In A. tenuipes, a rostral carina is present, but is less developed and not extending beyond the level of eyes (cf. De Man 1911). In addition to the strong balaeniceps condition of the male minor chela and well-developed rostral carina, A. vanuatu n. sp. differs specifically from each of the above-mentioned species, e.g. from A. paracrinitus sensu Banner & Banner (1982) by the major cheliped merus with the tooth on the ventromesial margin in subdistal position (vs. more proximal, closer to merus mid-length in A. paracrinitus of Banner & Banner 1982); from A. paracrinitus sensu Miers (1881), Crosnier & Forest (1966) and Kim & Abele (1988) by the longer stylocerite, exceeding the distal margin of the first article of the antennular peduncle (vs. not reaching it in A. paracrinitus) and the third maxilliped not protruding distodorsally (cf. Crosnier & Forest 1966; Kim & Abele 1988); from the form described as A. paracrinitus bengalensis Couti��re, 1905 by the longer stylocerite, as well as the proportions of the major chela and telson; from A. alpheopsides by the relatively longer second article of the antennular peduncle, the stronger distolateral tooth of the scaphocerite and the much narrower telson; from A. paralpheopsides by the shape of the rostro-orbital margin, especially by the margin between the rostrum and orbital hood not as deeply incised as in A. paralpheopsides, and by the second pereiopod carpus with the first subdivision much longer than the second (vs. the two being subequal in A. paralpheopsides) (cf. Couti��re 1905); from A. rostratus by the longer stylocerite, exceeding the distal margin of the first article of the antennular peduncle (vs. not reaching it in A. rostratus), the distomesial margin of the major chela palm protruding forward, forming almost a right angle, the major cheliped merus with the tooth on the ventromesial margin in subdistal position (vs. distal in A. rostratus), and the third pereiopod dactylus noticeably longer and slenderer (cf. Kim & Abele 1988); from A. labis by the distolateral tooth of the scaphocerite by far overreaching the distal margin of the blade (vs. not reaching it in A. labis), the non-expanded antepenultimate article of the third maxilliped (noticeably expanded in A. labis), the major chela with a much higher palm length / dactylus length ratio (i.e. with the relatively longer dactylus in the new species), and the major cheliped merus without spiniform setae on the ventromesial margin; from A. mitis sensu Banner & Banner (1982, but see Banner & Banner 1983: p. 55) by the longer stylocerite, exceeding the distal margin of the first article of the antennular peduncle (vs. not reaching it in A. mitis) and the major cheliped merus less stout and with the tooth on the ventromesial margin clearly in subdistal position (vs. more distal in A. mitis) (cf. Banner & Banner 1982); and from A. tenuipes by the major cheliped merus much stouter and without spiniform setae along the ventromesial margin (cf. De Man 1911). Based on the comparisons above, A. vanuatu n. sp. does not seem to be very closely related to any of the species of the A. paracrinitus group, the morphologically closest taxa being A. tenuipes and A. labis, and possibly A. paracrinitus sensu Banner & Banner (1982)., Published as part of Anker, Arthur, 2020, On two new deep-water snapping shrimps from the Indo-West Pacific (Decapoda Alpheidae: Alpheus), pp. 393-409 in Zootaxa 4845 (3) on pages 401-407, DOI: 10.11646/zootaxa.4845.3.5, http://zenodo.org/record/4406729, {"references":["Miers, E. J. (1881) On a collection of Crustacea made by Baron Hermann-Maltzan at Goree island, Senegambia. The Annals and Magazine of Natural History, Series 5, 8, 204 - 220 + 259 - 281 + 364 - 377, pls. 13 - 16. https: // doi. org / 10.1080 / 00222938109487471","Dana, J. D. (1852) Conspectus Crustaceorum & c. Conspectus of the Crustacea of the Exploring Expedition under Capt. C. Wilkes, U. S. N. Macroura. Proceedings of the Academy of Natural Sciences of Philadelphia, 1852, 10 - 29.","Coutiere, H. (1905) Les Alpheidae. In: Gardiner, J. S. (Ed.), The fauna and geography of the Maldive and Laccadive Archipelagoes. Being the account of the work carried on and of the collections made by an expedition during the years 1899 and 1900. University Press, Cambridge, pp. 852 - 921, pls. 70 - 87.","Miya, Y. (1974) The Alpheidae (Crustacea, Decapoda) of Japan and its adjacent waters. Part II. Publications from the Amakusa Marine Biology Laboratory, 3, 103 - 195, pls. 15 - 31.","Banner, D. M. & Banner, A. H. (1982) The alpheid shrimp of Australia, part III: the remaining alpheids, principally the genus Alpheus, and the family Ogyrididae. Records of the Australian Museum, 34, 1 - 357. https: // doi. org / 10.3853 / j. 0067 - 1975.34.1982.434","Chace, F. A. Jr. (1988) The caridean shrimps (Crustacea: Decapoda) of the Albatross Philippine Expedition, 1907 - 1910, Part 5: Family Alpheidae. Smithsonian Contributions to Zoology, 466, 1 - 99. https: // doi. org / 10.5479 / si. 19436696.391.1","Kim, W. & Abele, L. G. (1988) The snapping shrimp genus Alpheus from the eastern Pacific (Decapoda: Caridea: Alpheidae). Smithsonian Contributions to Zoology, 454, 1 - 119. https: // doi. org / 10.5479 / si. 00810282.454","De Man, J. G. (1910) Diagnoses of new species of macrurous decapod Crustacea from the \" Siboga-Expedition \". Tijdschrift der Nederlandsche Dierkundige Vereeniging, 2 (11), 287 - 319.","Banner, A. H. & Banner, D. M. (1983) An annotated checklist of the alpheid shrimp from the western Indian Ocean. Travaux et Documents de l'ORSTOM, 158, 1 - 164.","Ashrafi, H., Dehghani, A., Sari, A. & Naderloo, R. (2020) An updated checklist of caridean shrimps of the Persian Gulf and Gulf of Oman. Zootaxa, 4747 (3), 521 - 534.","De Man, J. G. (1911) The Decapoda of the Siboga Expedition. Part II. Family Alpheidae. In: Siboga Expeditie. 39 a 1. Brill, Leiden, pp. 133 - 465; (1915) Supplement-Explanation of plates of Alpheidae, pls. 1 - 23.","Crosnier, A. & Forest, J. (1966) Crustaces decapodes: Alpheidae. In: Campagnes de la Calypso dans le Golfe de Guinee et aux Iles Principe, Sao Tome et Annobon (1956), et Campagne aux Iles du Cap Vert (1959). Part 19. Resultats Scientifiques des Campagnes de la Calypso 7 (27). Annales de l'Institut Oceanographique de Monaco, 44, 199 - 314."]}

Alpheus malleator Dana, 1852 Zoea I Carapace length: 0.52 �� 0.03 mm (n = 10 larvae) Cephalothorax (Fig. 2A, 2B): rostrum absent; pterygostomian spine present; eyes sessile. Antennule (Fig. 2C): peduncle 2-segmented; endopod as a plumose seta; exopod with 4 terminal aesthetascs (1 more robust) and 1 terminal plumose seta. Antenna (Fig. 2D): peduncle 2-segmented with 1 medial distal spiniform projection near the endopod; endopod with 1 terminal plumose seta and 1 medial distal spiniform projection; exopod (scaphocerite) 6-segmented with 11 plumose setae (2, 1, 2, 1, 1, 4), plus a medial tubercle in the proximal segment and 1 simple seta in the distal segment. Mandible: palp absent. Maxillule (Fig. 2E): coxa with 6 (1 medial simple, 3 terminal simple, 1 terminal serrulate plumose, larger in proximal region, 1 lateral simple) setae; base with 2 spiniform projections and 1 lateral simple seta; endopod with 1 terminal plumoserrulate and 1 lateral simple setae; exopod absent. Maxilla (Fig. 2F): coxa with 2 medial simple setae; base bilobed, each lobe with 5 medial simple setae (median seta shorter); endopod with 3 medial (1 proximal and 2 distal) simple setae and 2 terminal long (1 simple and 1 plumose) setae, with microtrichia; exopod (scaphognathite) with 5 plumose setae, with microtrichia. First maxilliped (Fig. 2G): coxa with 1 medial distal simple seta; base with 7 medial simple setae arranged 2+2+2+1; endopod 2-segmented with 1 medial distal simple seta, 5 [4 simple (1 medial + 1 lateral + 2 terminal) and 1 terminal serrulate] setae; exopod 4-segmented with 0, 0, 2, 2 terminal natatory plumose setae plus 1 lateral proximal simple seta on third segment. Second maxilliped (Fig. 2H): coxa without setae; base with 4 medial simple setae arranged 1+2+1; endopod 4-segmented with 1 medial distal simple, 0, 1 medial distal serrulate seta, 5 (1 medial simple, 3 lateral simple and 1 terminal serrulate) setae; exopod 4-segmented with 0, 1, 2, 2 terminal natatory plumose setae plus 1 terminal simple seta on second segment. Third maxilliped (Fig. 2I): coxa with 1 medial distal simple seta; base with 1 medial simple seta; endopod 4segmented with 0, 0, 3 (2 medial proximal and 1 lateral distal), 3 (2 lateral and 1 terminal) simple setae; exopod 4-segmented with 0, 2, 2, 2 terminal natatory plumose setae plus 2 (1 medial and 1 lateral) simple setae on second segment. Pereopod (Fig. 2J): first as a biramous bud. Pleon (Fig. 2A, 2K): with 5 somites, presence of simple dorsal setae on the somites (Figs. 2L, 3A, 3D, 3E, 3F); lateral spines absent; pleopods absent; anal spine present (Fig. 2M). Telson (Fig. 2M): triangular, with 7+7 (5 inners plumose, 2 outers plumose laterally) setae, 4 inner setae with shorter setules in the proximal region, inner pair shorter; 6 inner pairs with a row of denticles in the distal margin, 4 innermost pairs with marginal spines., Published as part of Pasinatto, Karmine, Mantelatto, Fernando L. & Terossi, Mariana, 2020, First zoeal stage of the snapping shrimps Alpheus formosus Gibbes, 1850 and Alpheus malleator Dana, 1852 (Caridea: Alpheidae), with new characters to the genus, pp. 540-550 in Zootaxa 4820 (3) on pages 542-546, DOI: 10.11646/zootaxa.4820.3.7, http://zenodo.org/record/4441079, {"references":["Dana, J. D. (1852) Conspectus Crustaceorum & c. Conspectus of the Crustacea of the Exploring Expedition under Capt. C. Wil- kes, U. S. N. Macroura. Proceedings of the Academy of Natural Sciences of Philadelphia, 1852, 10 - 29.","Gibbes, L. R. (1850) On the carcinological collections of the United States, and an enumeration of species contained in them, with notes on the most remarkable, and descriptions of new species. Proceedings of the American Association for the Advancement of Science, 3, 165 - 201."]}

Alpheus formosus Gibbes, 1850 Zoea I Carapace length: 0.42 �� 0.04 mm (n = 10 larvae) Cephalothorax (Fig. 1A, 1B): rostrum absent; pterygostomian spine present; eyes sessile. Antennule (Fig. 1C): peduncle 3-segmented; endopod as a plumose seta; exopod with 4 terminal aesthetascs (2 spatulate, one of them being more robust) and 1 terminal plumose seta. Antenna (Fig. 1D): peduncle 2-segmented with 1 medial distal spiniform projection near the endopod; endopod with 1 terminal plumose seta and 1 medial distal spiniform projection; exopod (scaphocerite) 8-segmented with 11 plumose setae (2, 1, 2, 1, 1, 1, 1, 2), distal segment with 1 simple seta. Mandible: palp absent. Maxillule (Fig. 1E): coxa with 6 (1 medial simple, 3 terminal simple, one larger in proximal region, 1 terminal serrulate plumose, 1 lateral simple) setae; base with 2 spiniform projections and 1 lateral simple seta; endopod with 1 terminal plumoserrulate and 1 lateral simple setae; exopod absent. Maxilla (Fig. 1F): coxa with 2 medial simple setae; base bilobed, each lobe with 5 medial simple setae (distal seta longer); endopod with 1 medial proximal simple and 2 terminal long (1 simple and 1 plumose) setae, with microtrichia; exopod (scaphognathite) with 5 plumose setae, without microtrichia. First maxilliped (Fig. 1G): coxa with 1 medial distal simple seta; base with 7 medial simple setae arranged 2+2+2+1; endopod 2-segmented with 1 medial distal simple seta, 5 [4 simple (1 medial + 3 terminal) and 1 terminal serrulate] setae; exopod 4-segmented with 0, 0, 2, 2 terminal natatory plumose setae plus 1 lateral distal simple seta on second segment. Second maxilliped (Fig. 1H): coxa without setae; base with 4 medial simple setae arranged 1+1+1+1; endopod 4-segmented with 1 medial distal simple, 0, 1 medial distal serrulate, 5 (1 medial simple, 3 lateral simple and 1 terminal serrulate) setae; exopod 4-segmented with 0, 1, 2, 2 terminal natatory plumose setae plus 1 terminal simple seta on second segment. Third maxilliped (Fig. 1I): coxa with 1 medial distal simple seta; base with 2 medial simple setae arranged 1+1; endopod 5-segmented with 0, 1 medial, 0, 3 (2 lateral proximal and 1 medial distal), 3 (2 medial proximal and 1 terminal) simple setae; exopod 5-segmented with 0, 0, 2, 2, 2 terminal natatory plumose setae plus 1 medial simple seta on second segment. Pereopod (Fig. 1J): first as a biramous bud. Pleon (Fig. 1A, 1K,): with 5 somites, presence of simple dorsal setae on the somites (Figs. 1L, 3B, 3C); lateral spines absent; pleopods absent; anal spine present (Fig. 1M). Telson (Fig. 1M): triangular, with 7+7 (5 inners plumose, 2 outers plumose laterally) setae, 4 inner setae with shorter setules in the proximal region, inner pair shorter; 6 inner pairs with a row of denticles in the distal margin, 4 innermost pairs with marginal spines (Fig. 3G, 3H)., Published as part of Pasinatto, Karmine, Mantelatto, Fernando L. & Terossi, Mariana, 2020, First zoeal stage of the snapping shrimps Alpheus formosus Gibbes, 1850 and Alpheus malleator Dana, 1852 (Caridea: Alpheidae), with new characters to the genus, pp. 540-550 in Zootaxa 4820 (3) on pages 541-542, DOI: 10.11646/zootaxa.4820.3.7, http://zenodo.org/record/4441079, {"references":["Gibbes, L. R. (1850) On the carcinological collections of the United States, and an enumeration of species contained in them, with notes on the most remarkable, and descriptions of new species. Proceedings of the American Association for the Advancement of Science, 3, 165 - 201."]}

Alpheus edwardsii (Audouin, 1826) sensu lato (Fig. 2) Athanas Edwarsii Audouin 1826: 91 (lap. cal.). Athanas Edwardsii. ��� Audouin 1827: 274. Alpheus Edwardsii. ��� Couti��re 1898: 249, figs. 4, 5; Nobili 1901: 2; Banner & Banner 1973a: 1141, fig. 1; Kazmi & Kazmi 1979: 154; Banner & Banner 1981: 22, 92; Banner & Banner 1982: 270, fig. 83; Lewinsohn & Galil 1982: 208; Afzal et al. 1986: 338; De Grave & Ashelby 2011: 3; Naderloo & T��rkay 2012: 9; Naderloo et al. 2013: 448; Dehghani et al. 2019a: 484, fig. 3A, B. Alpheus edwardsii sensu lato.��� Anker & De Grave 2016: 352, figs. 10, 11. (?) Alpheus Audouini Couti��re 1905: 911, pl. 87, fig. 52. Not Alpheus edwardsii. ��� Al-Maliky et al. 2017: 49, figs. 2���4 [= Alpheus lobidens De Haan, 1849]. Material examined. Kuwait: 1 male, OUMNH. ZC. 2020.01.018, Kubbar Island, 29��04���18.93���N 48��29���18.74���E, depth 2���3 m, sand flat with sparse rubble, under coral rocks, leg. S. De Grave, 26.ix.2019 [fcn KUW-017]; 2 males, 1 female, KISR, Miscan Island, sandy-rocky intertidal, leg. M. Al-Kandari et al., 15.xii.2019. General distribution. Species complex (see Remarks below), reported from throughout the Indo-West Pacific, from the Red Sea, Arabian Gulf (Iran, Kuwait, UAE) and South Africa across the Indian Ocean to Japan, Australia and French Polynesia (e.g., Banner & Banner 1973a, 1981, 1982, 1983; Chace 1988; Naderloo & T��rkay 2012; Anker & De Grave 2016; Dehghani et al. 2019a; present study); Lessepsian migrant into the eastern Mediterranean Sea (Forest & Guinot 1958; Holthuis & Gottlieb 1958). Ecology. Variable (species complex, see below); found both in the intertidal and shallow subtidal (to at least 36 m), rocky and mixed rocky-sandy shores, also on coral reefs and associated coral rubble habitats (reef flats); typically, under large rocks and coral rubble (e.g. Banner & Banner 1982; Corfield & Alexander 1995; present study). Remarks. Without a comprehensive revision of the A. edwardsii species complex (cf. Anker & De Grave 2016), it is impossible to apply a correct species name to the likely more than one species present in the Arabian Gulf, at least in its more biodiverse eastern part (see also Dehghani et al. 2019a). In addition, many literature records of A. edwardsii are unclear and may refer to closely or even not closely related taxa, exemplified by the regional record of Al-Maliky et al. (2017), who erroneously reported A. edwardsii as A. lobidens from Shatt Al Basra and Faw in Iraq. The photographed male specimen from Kuwait (Fig. 2) is characterised by the overall brown to brown-green colour, the presence of narrow longitudinal white streaks on the pleon flanks and two small dark spots on the dorsal surface of the third pleonite, as well as by the brightly coloured major chela, with conspicuous blue, yellow, olivegreen and red-orange areas on the mesial face of the palm. This colour pattern seems to correspond to that of the female from Iran illustrated by Dehghani et al. (2019a: fig. 3B), although the Iranian specimen is much paler and greener., Published as part of Al-Kandari, Manal, Anker, Arthur, Hussain, Sumaiah, Al-Yassen, Sharifa, Sattari, Zainab & Grave, Sammy De, 2020, New records of decapod crustaceans from Kuwait (Malacostraca: Decapoda), pp. 251-280 in Zootaxa 4803 (2) on page 254, DOI: 10.11646/zootaxa.4803.2.2, http://zenodo.org/record/3909077, {"references":["Audouin, V. (1826) Explication sommaire des planches de crustaces de l'Egypte et de la Syrie, publiees par Jules-Cesar Savigny, Membre de l'Institut; offrant un expos des characteres naturels des genres avec la distinction des especes. Animaux invertebres. In: Description de l'Egypte ou receuil des observations et des recherches qui ont et faites en Egypte pendant l'expedition de l'arme francaise, publi par les ordres de sa Majest l'Empereur Napoleon le Grand. Imprimerie Imperiale, Paris, pp. 77 - 98.","Audouin, V. (1827) Description de l'Egypte, ou recueil des observations et des recherches qui ont ete faites en Egypte pendant l'expedition de l'armee francaise, publie par les ordres de sa Majeste l'Empereur Napoleon le Grand. Vol. 22. Edition 2. Imprimerie Imperiale, Paris, 42 pp. [pp. 249 - 290]","Coutiere, H. (1898) Note sur quelques formes nouvelles d'Alpheides voisines de A. Bouvieri A. M. - Edwards [Crust.]. Bulletin de la Societe Entomologique de France, 1898, 131 - 134. https: // doi. org / 10.5962 / bhl. part. 16847","Nobili, G. (1901) Decapodi e Stomatopodi Eritrei del Museo Zoologico dell'Universita di Napoli. Annuario del Museo Zoologico della R. Universita di Napoli, 1, 1 - 21.","Banner, A. H. & Banner, D. M. (1973 a) The establishment of a neotype for Alpheus edwardsi (Audouin). Bulletin du Museum national d'Histoire naturelle, 3 eme serie, Zoologie, 88, 1141 - 1146.","Kazmi, M. A. & Kazmi, Q. B. (1979) A check-list of marine caridean prawns of Pakistan. Biologia, 25, 151 - 157.","Banner, D. M. & Banner, A. H. (1981) Annotated checklist of the alpheid shrimp of the Red Sea and Gulf of Aden. Zoologische Verhandelingen, Leiden, 190, 1 - 99.","Banner, D. M. & Banner, A. H. (1982) The Alpheid Shrimp of Australia, Part III: The remaining alpheids, principally the genus Alpheus, and the family Ogyrididae. Records of the Australian Museum, 34, 1 - 357. https: // doi. org / 10.3853 / j. 0067 - 1975.34.1982.434","Lewinsohn, C. & Galil, B. (1982) Notes on species of Alpheus (Crustacea Decapoda) from the Mediterranean coast of Israel. Quaderni del Laboratorio di Tecnologia della Pesca, Ancona, 3, 207 - 210.","Afzal, M., Javed, A. & Barkati, S. (1986) Checklist, synonymy and key to the genera and species of alpheid shrimps of Karachi coast. Pakistan Journal of Agriculture Research, 7, 337 - 340.","De Grave, S. & Ashelby, C. W. (2011) Notes on some shrimp species (Decapoda: Caridea) from Qatar, with an annotated checklist of marine caridean species known from the Persian Gulf. Marine Biodiversity Records, 4 (e 20), 1 - 7. https: // doi. org / 10.1017 / S 1755267211000224","Naderloo, R. & Turkay, M. (2012) Decapod crustaceans of the littoral and shallow sublittoral Iranian coast of the Persian Gulf: faunistics, biodiversity and zoogeography. Zootaxa, 3374 (1), 1 - 67. https: // doi. org / 10.11646 / zootaxa. 3374.1.1","Naderloo, R., Turkay, M. & Sari, A. (2013) Intertidal habitats and decapod (Crustacea) diversity of Qeshm Island, a biodiversity hotspot within the Persian Gulf. Marine Biodiversity, 43, 445 - 462. https: // doi. org / 10.1007 / s 12526 - 013 - 0174 - 3","Dehghani, A., Sari A. & Naderloo, R. (2019 a) Annotated checklist of the snapping shrimps of the genus Alpheus Fabricius, 1798 (Crustacea: Decapoda: Alpheidae) from the Persian Gulf and Gulf of Oman, Iran. Zootaxa, 4544 (4), 479 - 504. https: // doi. org / 10.11646 / zootaxa. 4544.4.2","Anker, A. & De Grave, S. (2016) An updated and annotated checklist of marine and brackish caridean shrimps of Singapore (Crustacea, Decapoda). Raffles Bulletin of Zoology, Supplement 34, 343 - 454.","Coutiere, H. (1905) Les Alpheidae. In: Gardiner, J. S. (Ed.), The Fauna and Geography of the Maldive and Laccadive Archipelagoes. Being the account of the work carried on and of the Collections made by an Expedition during the years 1899 and 1900. University Press, Cambridge, pp. 852 - 921.","Al-Maliky, T. H. Y., Al-Khafaji, K. K. & Khalaf, T. A. (2017) New records of the snapping shrimp Alpheus edwardsii (Audouin, 1826) (Crustacea: Alpheoidea) in Basarh, Iraq. Arthropods, 6, 47 - 53.","Banner, A. H. & Banner, D. M. (1983) An annotated checklist of the alpheid shrimp from the Western Indian Ocean. Travaux et Documents de l'ORSTOM, 158, 1 - 164.","Chace, F. A. Jr. (1988) The caridean shrimps (Crustacea: Decapoda) of the Albatross Philippine Expedition, 1907 - 1910, Part 5: family Alpheidae. Smithsonian Contributions to Zoology, 466, 1 - 99. https: // doi. org / 10.5479 / si. 19436696.391.1","Forest, J. & Guinot, D. (1958) Sur une collection de crustaces decapodes des cotes d'Israel. Bulletin of the Sea Fisheries Research Station of Israel, 15, 4 - 16.","Holthuis, L. B. & Gottlieb, E. (1958) An annotated list of the Decapod Crustacea of the Mediterranean Coast of Israel, with an appendix listing the Decapoda of the eastern Mediterranean. Bulletin of the Research Council of Israel, 7 B, 1 - 126, pls. 1 - 3.","Corfield, J. L. & Alexander, C. G. (1995) The distribution of two species of alpheid shrimp, Alpheus edwardsii and A. lobidens, on a tropical beach. Journal of the Marine Biological Association of the United Kingdom, 75, 675 - 687. https: // doi. org / 10.1017 / S 0025315400039096"]}

Alpheus samudra nov. sp. (Figs. 1���5) Type material. Holotype: female (cl 13.7 mm), bycatch from vessels operating on Quilon Bank (between 8��� 11��N and 74��� 76��E), Sakthikulangara Fishing Harbour, Kollam, Kerala, leg. A. Kumar, 07.ii.2016 (SIFAN). Paratypes: male (cl 11.7 mm), same collection data (SIFAN); female (cl 10.2 mm), same location, leg. A. Krishnan, 08.iii 2016 (DABFUK). Non-type material. Male (cl 13.0 mm), from bycatch, Neendakara fishing port, Kollam, Kerala, leg. A.B. Kumar, 25.ii.2018, from bycatch (DABFUK); 3 males (cl 10.2, 11.5, 12.2 mm), 1 female (cl 11.6 mm), from bycatch, Sakthikulangara fishing port, Kollam, Kerala, leg. A.B. Kumar, 2015���2016 (DABFUK). Description. Medium- to large-sized species of Alpheus (cl reaching 13.8 mm). Carapace tuberculate (Fig. 1 A���B). Rostrum well developed (Fig. 1 A���B), relatively long, about 1.5 times as long as broad at base, sharp distally, reaching to distal margin of first article of antenullar peduncle; rostral carina weak, not extending posterior to orbital hoods; lateral margin non-setose. Orbital hoods swollen, anterior margin lacking teeth. Pterygostomial angle rounded (Fig. 1A), not protruding; cardiac notch well developed. Telson (Fig. 1D) elongate, gradually tapering lateral margins, about twice as long as proximal width (at widest point); dorsal surface with two pairs of spiniform setae, both inserted at some distance from lateral margin, proximal pair at about 0.4 of telson length, distal-most pair at about 0.7 of telson length; posterior margin broadly convex (Fig. 1E), furnished with long plumose setae and short, spiniform setae; each posterolateral angle with single stout spiniform seta in holotype (pair of spiniform setae present in male paratype, inner pair longest). Antennular peduncle (Fig. 1 A���B) with second article elongate, about 3 times as long as wide; stylocerite with sharp point almost reaching to distal margin of first article; ventromesial carina (Fig. 1C) with sharply delineated triangular tooth. Antenna (Fig. 1 A���B) with stout basicerite, bearing blunt distoventral tooth; scaphocerite relatively stout, with marginally concave lateral margin and well-developed blade, with strong distolateral tooth, overreaching blade; carpocerite reaching to about distal margin of second article of antennular peduncle. Mouthparts typical for genus in external observation. Third maxilliped (Fig. 3A) slender, pediform; antepenultimate article granulated, penultimate article relatively short, about 1.5 times as long as wide, ultimate article richly setose, unarmed distally. Major cheliped (Fig. 2 A���B) similar in males and females, not significantly larger in males; ischium short, rugose on mesial margin, finely granulated on lateral margin; merus elongate, about 4.5 times as long as wide, distomedial margin unarmed, mesial and lateral margin granulated, distomedial margin rounded; carpus cup-shaped, with broadly rounded distal lobes, faintly granulated; chela elongate, extremely laterally compressed, about 4.0 times as long as wide, about 3.0 times as high as thick; palm about as long as fingers, densely granulated, but patchy in places, without any grooves, crest or notches, highest immediately proximal to dactylar articulation, tapering proximally, area of maximum thickness above middle with lower half of palm blade-like, inferior edge carinate; dactylus about same length as pollex, sharply carinate on inferior margin, curved slightly outwards, plunger reduced (Fig. 4), defined only by proximal lobe; pollex distally curved, continuing with inferior carinate edge of palm, fossa shallow, poorly-developed. Minor cheliped (Fig. 2C) not sexually dimorphic, nearly of equal length to major cheliped; ischium short, granulated; merus elongate, about 5 times as long as wide, granulated, distomedial margin unarmed; carpus cupshaped; chela elongate, slender, with palm sub-cylindrical in cross-section, almost half as long as fingers, surface smooth, without any grooves, crests or notches; fingers subequal in length, tapering to tips, distally slightly arched, with simple, blade-like cutting edges; long setae covering chela and palm. Second pereiopod slender (Fig. 3 B���C), ischium somewhat shorter than merus, both with rugose surfaces; carpus with five divisions, with approximate ratios (proximal to distal) of 1:1.4:0.4:0.5:0.5. Third pereiopod (Fig. 3 D���F) relatively robust; ischium somewhat granulated, merus about 6.5 times as long as wide, unarmed; carpus about half length of merus, unarmed; propodus about 1.2 length of carpus, unarmed; dactylus spatulate, about 0.6 length of propodus, lateral surface with cluster of setae; mesial surface with 5���6 short rows of setae. Fourth pereiopod (not illustrated) similar to third. Fifth pereiopod (Fig. 3 G���H) much more slender than third and fourth; ischium unarmed; merus about 6.5 times as long as wide; carpus 0.9 length of merus; propodus subequal in length to carpus, ventromesial margin unarmed, cleaning brush well-developed; dactylus subspatulate, simple, about 0.5 length of propodus. Male second pleopod (Fig. 2 D���E) with appendix masculina slightly longer than appendix interna, adorned with setae on mesial and lateral margins, as well as on apex. Uropod (Fig. 1F) with lateral lobe of protopod unarmed (mesial lobe distally damaged in holotype), mesial lobe in male paratype terminating in subacute tooth; exopod broad, somewhat truncate distally, diaresis weakly sinuous, with small triangular tooth adjacent and mesial to short spiniform seta; endopod without special features. Colour pattern (Fig. 5). On a recently deceased specimen, traces of transverse red banding remain on the pleon, whilst the chelipeds have an olive-green colour, with the fingertips of the major cheliped being peach coloured. Fresh specimens have been noted to also display red banding on the distal part of each pereiopod article, with the uropods and telson being translucent-red. Etymology. The new species name is from the Sanskrit ���samudra���, meaning the gathering together of waters, i.e. the ocean; used as a noun in apposition. Type locality. Quilon Bank, Kerala, India Distribution. Currently only known from the type locality. Ecology. The fishing vessels from which the specimens were obtained operated on a fine sandy to muddy bottom in depths of 275��� 375 m. Taxonomic remarks. The present new species can comfortably be referred to the A. brevirostris species group (sensu Bruce, 1994), on account of the major chela being strongly compressed and sub-rectangular in cross-section. This species group contains about 50 species (see Komai & Ohtomi, 2018), with several deep-water representatives. However, the extreme lateral compression of the chela of the major cheliped, at once separates the new species from all species in the group, except A. leptocheles Banner & Banner, 1975, a shallow water (Alpheus leptocheles was described on the basis of a male holotype and two female paratypes, with the description noting that the observed sexual dimorphism in the major cheliped would be unusual within the genus (Banner & Banner, 1975). Although the authors believed that they were dealing with a single species, they also noted that the male was in a different vial than the two females ������this may indicate a separate habitat or even a separate trawl haul���. As the major chela of the female paratypes of A. leptocheles are reminiscent of the usual type observed within the A. brevirostris group, it seems highly likely that two species were mixed up in the type description of A. leptocheles and further comparison is restricted to the holotype only. The major chela of the holotype of A. leptocheles is extremely laterally compressed, as is the case in the new species. Banner & Banner (1975) state that the palm is ������7 times as high as thick; ������ although their illustration (Fig. 1D) shows this to be incorrect. At our request, S. Ahyong examined the holotype in the collection of the Australian Museum and confirmed that the drawing is correct with a measured ratio of 3.37 of height vs thickness. This is somewhat similar to the ratio observed in the new species, and can probably not be used as a distinguishing feature between both species. Although Banner & Banner (1975) do not comment on the nature of the plunger-fossa, it was also confirmed that in the holotype this is similar to the new species, in having a poorly developed plunger and a shallow fossa. Both species can however be distinguished by the relative proportion and shape of the fingers of the major cheliped. In A. leptocheles, the finger to palm ratio is 0.36 and the dactylus is relatively robust, whilst in A. samudra nov. sp., the fingers are longer (about 0.50���0.55) and the dactylus is more elongate. Asides from the differences in the major cheliped, A. samudra nov. sp. also differs from A. leptocheles by the rostrum reaching the distal margin of the first article of the antennular peduncle (vs. falling short in A. leptocheles), the length of the fingers of the minor cheliped being shorter in relation to the palm (2.2 times in A. leptocheles vs. 1.7���1.8 in A. samudra nov. sp.), the general shape of the telson (more elongate and more distally concave in A. samudra nov. sp.), the strongly convex distal margin of the telson (vs. weakly convex in A. leptocheles) and the distolateral tooth of the scaphocerite over-reaching the blade (vs. tip being level with blade in A. leptocheles). A further distinct difference between these two species, is that the distal margin of the telson is furnished with a large series of short, spiniform setae in A. samudra nov. sp., whilst these are absent in the holotype of A. leptocheles (confirmed by S. Ahyong). A potential ecological difference also exists between both species. Alpheus leptocheles was obtained from depths of 1���20 m from the mouth of the Sepik River (Papua New Guinea), whilst the new species was obtained from depths of 275���375 m, albeit on a similar substrate of fine sand and mud., Published as part of Grave, Sammy De, Krishnan, Apsara S., Kumar, Anil & Christodoulou, Magdalini, 2020, A new species of Alpheus (Crustacea, Decapoda, Alpheidae) from the Arabian Sea, Kerala, India, pp. 277-285 in Zootaxa 4750 (2) on pages 278-284, DOI: 10.11646/zootaxa.4750.2.11, http://zenodo.org/record/3707303, {"references":["Bruce, A. J. (1994) Alpheus fenneri sp. nov. and A. williamsi sp. nov., two new Indo-West Pacific alpheid shrimps of the brevirostris species group. The Beagle, Records of the Museums and Art Galleries of the Northern Territory, 11, 15 - 28.","Komai, T. & Ohtomi, J. (2018) A new deep-sea species of the snapping shrimp genus Alpheus Fabricius, 1798 (Decapoda: Caridea: Alpheidae) from Kagoshima Bay, Japan. Zootaxa, 4434 (1), 99 - 110. https: // doi. org / 10.11646 / zootaxa. 4434.1.6","Banner, A. H. & Banner, D. M. (1975) Contributions to the knowledge of the alpheid shrimp of the Pacific Ocean. Part XVIII: A new species of the genus Alpheus from the mouth of the Sepik River, New Guinea. Records of the Australian Museum, 29, 261 - 266."]}

Alpheus samudra nov. sp. (Figs. 1–5) Type material. Holotype: female (cl 13.7 mm), bycatch from vessels operating on Quilon Bank (between 8– 11°N and 74– 76°E), Sakthikulangara Fishing Harbour, Kollam, Kerala, leg. A. Kumar, 07.ii.2016 (SIFAN). Paratypes: male (cl 11.7 mm), same collection data (SIFAN); female (cl 10.2 mm), same location, leg. A. Krishnan, 08.iii 2016 (DABFUK). Non-type material. Male (cl 13.0 mm), from bycatch, Neendakara fishing port, Kollam, Kerala, leg. A.B. Kumar, 25.ii.2018, from bycatch (DABFUK); 3 males (cl 10.2, 11.5, 12.2 mm), 1 female (cl 11.6 mm), from bycatch, Sakthikulangara fishing port, Kollam, Kerala, leg. A.B. Kumar, 2015–2016 (DABFUK). Description. Medium- to large-sized species of Alpheus (cl reaching 13.8 mm). Carapace tuberculate (Fig. 1 A–B). Rostrum well developed (Fig. 1 A–B), relatively long, about 1.5 times as long as broad at base, sharp distally, reaching to distal margin of first article of antenullar peduncle; rostral carina weak, not extending posterior to orbital hoods; lateral margin non-setose. Orbital hoods swollen, anterior margin lacking teeth. Pterygostomial angle rounded (Fig. 1A), not protruding; cardiac notch well developed. Telson (Fig. 1D) elongate, gradually tapering lateral margins, about twice as long as proximal width (at widest point); dorsal surface with two pairs of spiniform setae, both inserted at some distance from lateral margin, proximal pair at about 0.4 of telson length, distal-most pair at about 0.7 of telson length; posterior margin broadly convex (Fig. 1E), furnished with long plumose setae and short, spiniform setae; each posterolateral angle with single stout spiniform seta in holotype (pair of spiniform setae present in male paratype, inner pair longest). Antennular peduncle (Fig. 1 A–B) with second article elongate, about 3 times as long as wide; stylocerite with sharp point almost reaching to distal margin of first article; ventromesial carina (Fig. 1C) with sharply delineated triangular tooth. Antenna (Fig. 1 A–B) with stout basicerite, bearing blunt distoventral tooth; scaphocerite relatively stout, with marginally concave lateral margin and well-developed blade, with strong distolateral tooth, overreaching blade; carpocerite reaching to about distal margin of second article of antennular peduncle. Mouthparts typical for genus in external observation. Third maxilliped (Fig. 3A) slender, pediform; antepenultimate article granulated, penultimate article relatively short, about 1.5 times as long as wide, ultimate article richly setose, unarmed distally. Major cheliped (Fig. 2 A–B) similar in males and females, not significantly larger in males; ischium short, rugose on mesial margin, finely granulated on lateral margin; merus elongate, about 4.5 times as long as wide, distomedial margin unarmed, mesial and lateral margin granulated, distomedial margin rounded; carpus cup-shaped, with broadly rounded distal lobes, faintly granulated; chela elongate, extremely laterally compressed, about 4.0 times as long as wide, about 3.0 times as high as thick; palm about as long as fingers, densely granulated, but patchy in places, without any grooves, crest or notches, highest immediately proximal to dactylar articulation, tapering proximally, area of maximum thickness above middle with lower half of palm blade-like, inferior edge carinate; dactylus about same length as pollex, sharply carinate on inferior margin, curved slightly outwards, plunger reduced (Fig. 4), defined only by proximal lobe; pollex distally curved, continuing with inferior carinate edge of palm, fossa shallow, poorly-developed. Minor cheliped (Fig. 2C) not sexually dimorphic, nearly of equal length to major cheliped; ischium short, granulated; merus elongate, about 5 times as long as wide, granulated, distomedial margin unarmed; carpus cupshaped; chela elongate, slender, with palm sub-cylindrical in cross-section, almost half as long as fingers, surface smooth, without any grooves, crests or notches; fingers subequal in length, tapering to tips, distally slightly arched, with simple, blade-like cutting edges; long setae covering chela and palm. Second pereiopod slender (Fig. 3 B–C), ischium somewhat shorter than merus, both with rugose surfaces; carpus with five divisions, with approximate ratios (proximal to distal) of 1:1.4:0.4:0.5:0.5. Third pereiopod (Fig. 3 D–F) relatively robust; ischium somewhat granulated, merus about 6.5 times as long as wide, unarmed; carpus about half length of merus, unarmed; propodus about 1.2 length of carpus, unarmed; dactylus spatulate, about 0.6 length of propodus, lateral surface with cluster of setae; mesial surface with 5–6 short rows of setae. Fourth pereiopod (not illustrated) similar to third. Fifth pereiopod (Fig. 3 G–H) much more slender than third and fourth; ischium unarmed; merus about 6.5 times as long as wide; carpus 0.9 length of merus; propodus subequal in length to carpus, ventromesial margin unarmed, cleaning brush well-developed; dactylus subspatulate, simple, about 0.5 length of propodus. Male second pleopod (Fig. 2 D–E) with appendix masculina slightly longer than appendix interna, adorned with setae on mesial and lateral margins, as well as on apex. Uropod (Fig. 1F) with lateral lobe of protopod unarmed (mesial lobe distally damaged in holotype), mesial lobe in male paratype terminating in subacute tooth; exopod broad, somewhat truncate distally, diaresis weakly sinuous, with small triangular tooth adjacent and mesial to short spiniform seta; endopod without special features. Colour pattern (Fig. 5). On a recently deceased specimen, traces of transverse red banding remain on the pleon, whilst the chelipeds have an olive-green colour, with the fingertips of the major cheliped being peach coloured. Fresh specimens have been noted to also display red banding on the distal part of each pereiopod article, with the uropods and telson being translucent-red. Etymology. The new species name is from the Sanskrit “samudra”, meaning the gathering together of waters, i.e. the ocean; used as a noun in apposition. Type locality. Quilon Bank, Kerala, India Distribution. Currently only known from the type locality. Ecology. The fishing vessels from which the specimens were obtained operated on a fine sandy to muddy bottom in depths of 275– 375 m. Taxonomic remarks. The present new species can comfortably be referred to the A. brevirostris species group (sensu Bruce, 1994), on account of the major chela being strongly compressed and sub-rectangular in cross-section. This species group contains about 50 species (see Komai & Ohtomi, 2018), with several deep-water representatives. However, the extreme lateral compression of the chela of the major cheliped, at once separates the new species from all species in the group, except A. leptocheles Banner & Banner, 1975, a shallow water ( Alpheus leptocheles was described on the basis of a male holotype and two female paratypes, with the description noting that the observed sexual dimorphism in the major cheliped would be unusual within the genus (Banner & Banner, 1975). Although the authors believed that they were dealing with a single species, they also noted that the male was in a different vial than the two females “…this may indicate a separate habitat or even a separate trawl haul”. As the major chela of the female paratypes of A. leptocheles are reminiscent of the usual type observed within the A. brevirostris group, it seems highly likely that two species were mixed up in the type description of A. leptocheles and further comparison is restricted to the holotype only. The major chela of the holotype of A. leptocheles is extremely laterally compressed, as is the case in the new species. Banner & Banner (1975) state that the palm is “…7 times as high as thick; …” although their illustration (Fig. 1D) shows this to be incorrect. At our request, S. Ahyong examined the holotype in the collection of the Australian Museum and confirmed that the drawing is correct with a measured ratio of 3.37 of height vs thickness. This is somewhat similar to the ratio observed in the new species, and can probably not be used as a distinguishing feature between both species. Although Banner & Banner (1975) do not comment on the nature of the plunger-fossa, it was also confirmed that in the holotype this is similar to the new species, in having a poorly developed plunger and a shallow fossa. Both species can however be distinguished by the relative proportion and shape of the fingers of the major cheliped. In A. leptocheles, the finger to palm ratio is 0.36 and the dactylus is relatively robust, whilst in A. samudra nov. sp., the fingers are longer (about 0.50–0.55) and the dactylus is more elongate. Asides from the differences in the major cheliped, A. samudra nov. sp. also differs from A. leptocheles by the rostrum reaching the distal margin of the first article of the antennular peduncle (vs. falling short in A. leptocheles), the length of the fingers of the minor cheliped being shorter in relation to the palm (2.2 times in A. leptocheles vs. 1.7–1.8 in A. samudra nov. sp.), the general shape of the telson (more elongate and more distally concave in A. samudra nov. sp.), the strongly convex distal margin of the telson (vs. weakly convex in A. leptocheles) and the distolateral tooth of the scaphocerite over-reaching the blade (vs. tip being level with blade in A. leptocheles). A further distinct difference between these two species, is that the distal margin of the telson is furnished with a large series of short, spiniform setae in A. samudra nov. sp., whilst these are absent in the holotype of A. leptocheles (confirmed by S. Ahyong). A potential ecological difference also exists between both species. Alpheus leptocheles was obtained from depths of 1–20 m from the mouth of the Sepik River (Papua New Guinea), whilst the new species was obtained from depths of 275–375 m, albeit on a similar substrate of fine sand and mud.

Alpheus chiragricus H. Milne Edwards, 1837 Material. 2 females & 2 males (ZUTC 6859), Qeshm Island, Persian Gulf, 26°29’07”N, 55°08’10”E, shrimp fishing trawling vessel, 02.05.2015, coll. A. Dehghani, H. Ashrafi & A. Sari. Habitat and habit. The specimens were found in sandy-muddy bottom with abundant small rocks. General distribution. Indo-West Pacific., Published as part of Ashrafi, Hossein, Dehghani, Amir, Sari, Alireza & Naderloo, Reza, 2020, An updated checklist of caridean shrimps of the Persian Gulf and Gulf of Oman, pp. 521-534 in Zootaxa 4747 (3) on page 522, DOI: 10.11646/zootaxa.4747.3.6, http://zenodo.org/record/3696260

Alpheus mitis Dana, 1852 (Fig. 1A) Material. 2 males (ZUTC 6758), Hormuz Island, Persian Gulf, 27°02’10”N, 56°29’23”E, intertidal, sandy with patches of rock, 02.04.2016, coll. A. Dehghani & H. Ashrafi; 1 female (ZUTC 6860), Qadir Park, Abu-Musa Island, Persian Gulf, 25°89’58”N, 55°04’23”E, sandy/rocky with coral, 17.07.2019, coll. H. Ashrafi. Habitat and habit. Sandy/rocky shores, usually under rocks or in dead coral branches in intertidal zone. General distribution. Indian Ocean and Pacific Ocean., Published as part of Ashrafi, Hossein, Dehghani, Amir, Sari, Alireza & Naderloo, Reza, 2020, An updated checklist of caridean shrimps of the Persian Gulf and Gulf of Oman, pp. 521-534 in Zootaxa 4747 (3) on page 522, DOI: 10.11646/zootaxa.4747.3.6, http://zenodo.org/record/3696260

Alpheus balaenodigitus Banner & Banner, 1982 Material. 1 female & 2 males (ZUTC 6853), Abu-Musa Island, Persian Gulf, 25°53’12”N, 55°02’10”E, 04.05.2016, coll. A. Dehghani & H. Ashrafi. Habitat and habit. The specimens were found in a rocky tidal pool and in dead coral branches. General distribution. Indian Ocean and Australia. Remark. These three specimens agree with Banner & Banner’s (1982) descriptions except that they noted margin of rostrum bearing six short setae, while no setae is seen in the Persian Gulf’s materials., Published as part of Ashrafi, Hossein, Dehghani, Amir, Sari, Alireza & Naderloo, Reza, 2020, An updated checklist of caridean shrimps of the Persian Gulf and Gulf of Oman, pp. 521-534 in Zootaxa 4747 (3) on page 522, DOI: 10.11646/zootaxa.4747.3.6, http://zenodo.org/record/3696260, {"references":["Banner, D. M. & Banner, A. H. (1982) The alpheid shrimps of Australia, part III: The remaining alpheids, principally the genus Alpheus, and family Ogyrididae. Records of the Australian Museum, 34, 1 - 357. https: // doi. org / 10.3853 / j. 0067 - 1975.34.1982.434"]}

Alpheus gallicus n. sp. (Figs. 1���4) Alpheus cf. macrocheles ��� Cartes et al. 2014: 168 (Table 1) [not Alpheus macrocheles (Hailstone, 1835)]. (?) Alpheus macrocheles��� Garc��a Raso 1996: 735 [not Alpheus macrocheles (Hailstone, 1835), see discussion below]. Type material. Holotype: male (cl 11.5 mm), OUMNH. ZC.2017-01-061, Spain, Galicia Bank, expedition INDE- MARES 0711, sta. V11, 42��44.209���N 11��46.352���W, depth: 768 m, beam trawl, leg. J.E. Cartes et al., 9 August 2011. Paratypes: 1 female (CL 13 mm), OUMNH. ZC.2017-01-062, same collection data as for holotype; 1 female (cl 11.25 mm), OUMNH. ZC.2017-01-063, same collection data as for holotype; 1 female (cl 11.9 mm), ICM 002618, same collection data as for holotype; 1 male (cl 9.6 mm), ICM 002619, Spain, Galicia Bank, expedition INDE- MARES 0711, sta. GOC3, 42��38.972���N 11��49.092���W, depth: 785 m, otter trawl / 2-doors trawl, leg. J.E. Cartes et al., 30 July 2011. Diagnosis. Species of Alpheus macrocheles species group. Frontal margin of carapace with well-developed rostrum and orbital teeth; rostrum distally acute, about 1.3 times as long as wide at base, tip not reaching halflength of first article of antennular peduncle; orbital teeth narrow and acute; stylocerite of antennule not reaching distal margin of first article or antennular peduncle; second antennular article slender, four times as long as wide; scaphocerite with shallowly concave lateral margin, stout distolateral tooth, reaching far beyond distal margin of narrow blade; third maxilliped not especially modified; major cheliped much larger and stouter than minor cheliped; major cheliped ischium with small spiniform seta on ventromesial margin; major cheliped merus with several small spiniform setae on ventromesial margin and acute distomesial tooth; major chela with distinct inferior plaque and superior crests separated by superior and inferior grooves, as well as with superior and inferior transverse grooves; superior and inferior crests ending distally in acute, anteriorly directed teeth; distal portion of major cheliped pollex twisted laterally, cutting edge bearing small tooth and notch; major chela dactylus twisted laterally, dorsally convex; plunger reduced to small proximal protuberance; minor cheliped ischium bearing small stout spiniform seta on ventral margin; ventromesial margin of minor cheliped merus without spiniform setae, with acute distomesial tooth; minor cheliped dactylus compressed, twisted laterally, with high dorsal crest; length ratio of subdivisions of second pereiopod carpus approximately equal to 9: 5: 3: 3: 4; meri of third and fourth pereiopods without distoventral tooth, ventral margin without spiniform setae; dactyli of third to fifth pereiopods inconspicuously biunguiculate; male second pleopod with appendix masculina slightly overreaching appendix interna; uropod with distolateral spiniform seta flanked laterally by acute tooth, mesially by obtuse projection. Description. Carapace glabrous, not setose; frontal margin with well-developed rostrum and orbital teeth; rostrum subtriangular, distally acute, about 1.3 times as long as wide at base, without setae, tip not reaching half-length of first article of antennular peduncle; rostral carina not distinct; orbital teeth narrow, distally acute, reaching halflength of rostrum, directed anteriorly and slightly mesially; frontal margin between rostrum and orbital teeth nearly straight at oblique angle; orbital hoods moderately swollen, completely enclosing eyes (Fig. 1a, b). Pterygostomial angle rounded, not protruding anteriorly (Fig. 1b); cardiac notch deep. Telson about twice as long as basal width; lateral margin slightly constricted posterior to half-length, tapering distally; dorsal surface smooth, bearing two pairs of stout cuspidate setae; anterior pair inserted slightly anterior to telson mid-length; posterior pair inserted at about 0.7 telson length; posterior margin slightly convex, distolateral angles each bearing one pair of spiniform setae, mesial about four times longer than lateral; margin between spiniform setae without spinules (Fig. 1c); anal tubercles well developed. Antennular peduncle slender; stylocerite well developed, with acute tip, not reaching distal margin of first article of antennular peduncle (Fig. 1a, b); ventromesial carina of first article subtriangular, with minute, anteriorly directed tooth (Fig. 1d); second article about twice as long as visible portion of first, four times as long as wide; third article shortest, one-fourth length of second (Fig. 1a, b); lateral flagellum with aesthetascs starting from about 15 th subdivision. Antenna with basicerite bearing short, acute distoventral tooth; carpocerite slender, about eight times as long as broad, overreaching both antennular peduncle and scaphocerite; scaphocerite with shallowly concave lateral margin; distolateral tooth stout, reaching far beyond distal margin of blade; blade fairly narrow, gradually tapering distally (Fig. 1a, b). Mouthparts typical for genus in external view. Third maxilliped slender, not especially modified; coxa with lateral plate not strongly produced, distal margin sparsely setose; exopod not overreaching penultimate article; antepenultimate article gradually flattening ventrally, ventral margin setose; penultimate article subcylindrical, slightly broadening distally, roughly 4.5 as long as maximal width; ultimate article long, slender, about 11 times as long as maximal width, unarmed distally, with transverse rows of serrulate setae on mesial surface (Fig. 1e). Major cheliped noticeably larger and stouter and somewhat longer than minor cheliped; ischium short, bearing small spiniform seta on ventromesial margin; merus broad, relatively short, subtriangular in cross-section, about 2.5 times as long as maximal width; ventrolateral margin unarmed; ventromesial margin bearing seven or so small spiniform setae and ending in sharp distal tooth; distodorsal margin bluntly protruding; carpus short, cup-shaped; chela somewhat compressed, twisted laterally; plaque crest deviating from dorsal margin in proximal third of chela, continuing as elevation on lateral surface; superior crest lamellar, dorsally rounded, ending distally in acute tooth; inferior crest distinct, ending distally as sharp acute tooth; superior groove deep, proximally extending beyond superior transverse groove; inferior groove originating shallowly near linea impressa, gradually deepening distally; superior transverse groove deep, with overhanging proximal shoulder; inferior depression deep, sharply delimited proximally; pollex short, distal portion twisted laterally, cutting edge bearing small tooth followed by triangular notch; dactylus twisted laterally, dorsally convex, flattened; tip with deep notch bearing tufts of setae; plunger greatly reduced to small proximal protuberance (Fig. 2 a���d). Minor cheliped with ischium short, bearing small, stout spiniform seta on ventral margin; merus short, broad, subtriangular in cross-section, about 2.5 times longer than maximal width; ventrolateral margin smooth; ventromesial margin without spiniform setae, with acute distomesial tooth; distodorsal margin not protruding; carpus short, cup-shaped; chela strongly compressed, twisted laterally; crests and grooves on lateral face of chela similar to those of major chela but less distinct; fingers subequal to palm in length; pollex with ventral margin distally curving, cutting edge excavated; dactylus compressed, lamellar, twisted laterally, with high dorsal crest (Fig. 2e, f). Second pereiopod long, slender; ischium slightly longer than merus; carpus with five subdivisions with length ratio approximately equal to 9: 5: 3: 3: 4; chela simple, fingers slightly longer than palm, with distal tufts of setae (Fig. 3a). Third pereiopod long, slender; ischium with one cuspidate seta on ventrolateral surface; merus compressed, about 6.5 times as long as maximal width; ventral margin without spiniform setae and without distal tooth; carpus about half-length of merus, slenderer than latter, distal portion of dorsal margin obtusely projecting; propodus slen- der, subequal in length to carpus, armed with twelve or so slender spiniform setae on ventral margin, in addition to two distoventral spiniform setae flanking base of dactylus; dactylus inconspicuously biunguiculate, with proximal unguis much smaller and shorter than distal one, extensor margin bearing small distal notch (Fig. 3 b���d). Fourth pereiopod generally similar to third in armature and proportions of articles, somewhat slenderer. Fifth pereiopod slenderest; ischium bearing small cuspidate seta on ventrolateral surface; merus and carpus subequal in length, unarmed; propodus bearing 10 or so spiniform setae on ventral margin and setal brush on distolateral surface; dactylus generally similar to that of third pereiopod (Fig. 3 e���g). Second pleopod with lateral margin of protopod unarmed, bearing long, fine setae; in males appendix masculina slightly overreaching appendix interna, but not overreaching distal margin of endopod, with numerous long distal setae (Fig. 1f). Uropod with protopod bearing large acute distolateral tooth; exopod with sinuous transverse suture (diaeresis); distolateral spiniform seta flanked by acute distolateral tooth laterally and broad obtuse projection mesially; endopod bearing row of minute spinules along distal margin (Fig. 1g). Colouration. Body uniformly reddish, with reddish chromatophores distributed throughout the body (carapace, pleon, telson); major and minor chelae orange-reddish; antennules, antennae and walking legs pale orange-reddish (Fig. 4). Etymology. The new species��� name refers to the type locality, Galicia Bank, situated in the offshore waters of Galicia, an autonomous community of Spain (gallicus ���Latin adjective for Galician, with a masculine ending); used as an adjective. Ecology. All specimens were trawled at considerable depths ranging from 768 to 785 m (type series), and down to 909 m (other material, see Cartes et al. 2014). Remarks. The distinct pattern of crests and grooves on the chelipeds clearly place Alpheus gallicus n. sp. within the Alpheus macrocheles species group sensu Couti��re (1905). Within this species group, A. gallicus n. sp. is morphologically closest to three deep-water species, viz. A. lentiginosus, A. romensky, and A. platydactylus, but also to the shallow-water A. macrocheles. These five taxa are indeed similar in the shape of the frontal margin of the carapace and also many features of the major and minor chelipeds, as well as the walking legs (third to fifth pereiopods). The most important morphological characters uniting these taxa are: (1) the well-developed rostrum and orbital teeth; (2) the major chela palm with well-developed crests and deep grooves, including a superior transverse groove; (3) the major chela dactylus with a greatly reduced plunger; (4) the major chela pollex not or only feebly reduced; (5) the meri of the third and fourth pereiopods without a sharp distoventral tooth; (6) the second article of the antennular peduncle slender (length to width ratio>3); and (7) the dorsal margin of the cheliped meri without spiniform setae. Alpheus lentiginosus, presently known only from its type locality in the Gulf of Mexico, can be distinguished from A. gallicus n. sp. by the absence of a notch the extensor margin of the dactyli of the third to fifth pereiopods (present in A. gallicus n. sp.), and by the number of spiniform setae on the ventral margin of the cheliped ischia (three in A. lentiginosus vs. one in A. gallicus n. sp.) (cf. Anker & Nizinski 2011). Alpheus romensky, was described from a single incomplete specimen collected from the Salas y Gomez Ridge in the Central Pacific. The frontal margin and major cheliped of this species are similar to those of A. gallicus n. sp., but it can be easily distinguished from the new species by the shape of the minor chela dactylus (slender, mesially somewhat concave in A. romensky vs. greatly expanded dorsally, distinctly convex in A. gallicus n. sp.), as well as the development of the superior transverse groove of the major chela (shallow in A. romensky vs. fairly deep in A. gallicus n. sp.) (cf. Burukovsky 1990). Alpheus platydactylus is a deep-water eastern Atlantic species found at depths between 55 m and 600 m (Couti��re 1897; Crosnier & Forest 1966). Based on both previously published descriptions and the presently examined material (see comparative material), A. platydactylus is distinguishable from A. gallicus n. sp. mainly by the shape and setation of the minor chela. In A. platydactylus, the minor chela dactylus (cf. Crosnier & Forest 1966: fig. 2g) is slender and not expanded dorsally (vs. strongly expanded in A. gallicus n. sp.), and bears dense rows of elongate setae throughout its length (absent in A. gallicus n. sp.). Alpheus macrocheles is a shallow to moderately deep-water species (typically 0���200 m, but see below) that has been reported from throughout the eastern Atlantic from Great Britain to Gabon, including the Mediterranean Sea, as well as from Ascension Island and St. Helena in the central Atlantic (Holthuis 1951; Crosnier & Forest 1966; Chace 1966; Manning & Chance 1990, Soledade et al. 2019). Alpheus macrocheles may be distinguished from A. gallicus n. sp. by the shape of the dactyli of the third to fifth pereiopods (simple in A. macrocheles vs. minutely biunguiculate in A. gallicus n. sp.); the shorter and stouter antennular peduncles (length-with ratio of the second article 2.5���3 in A. macrochele s vs. closer to 4 in A. gallicus n. sp.); the length of the stylocerite (reaching or nearly reaching the distal margin of the first antennular article in A. macrocheles vs. not reaching its distal margin in A. gallicus n. sp.); and the number of spiniform setae on the ventral surface of the major cheliped ischium (three in A. macrocheles vs. one in A. gallicus n. sp.). The two species also differ by the colouration of the major chela, with A. macrocheles displaying a transverse, more or less W-shaped blotch on the distomesial face of the palm (cf. Anker & De Grave 2012: fig. 4A, B). Several other members of the A. macrocheles species group seem to be more distantly related to A. gallicus n. sp. based on their morphological characteristics. Alpheus puapeba Christoffersen, 1979 is a moderately deep-water species (45���175 m) known from Brazil, Uruguay and Argentina (Christoffersen 1979). It can be distinguished from A. gallicus n. sp. by the shape of the dactyli of the third to fifth pereiopods (simple in A. puapeba vs. minutely biunguiculate in A. gallicus n. sp.); the shape of the minor cheliped dactylus (not expanded in A. puapeba vs. dorsally expanded in A. gallicus n. sp.); and the development of the inferior transverse groove of the major chela (with strongly overhanging shoulder in A. puapeba vs. with very shallowly overhanging shoulder in A. gallicus n. sp.). The members of the Indo-West Pacific A. hailstonei species complex, which includes the non-monophyletic A. hailstonei (Scioli et al., in prep.), A. brachymerus Banner, 1953 and A. lanceostylus Banner, 1959, can be distinguished from A. gallicus n. sp. by the presence of spiniform setae on the dorsal margin of the cheliped meri (present in most members of this complex, absent in A. gallicus n. sp.), as well as by several features on the minor cheliped, such as the proportions of articles and the general shape of the minor chela (cf. Couti��re 1905; Banner 1953, 1959; Banner & Banner 1982). The poorly known A. candei Gu��rin-M��neville, 1855, originally described from Cuba, is a taxonomically highly problematic species that has not been collected since Couti��re���s (1910) tentative identification of a single specimen from Dry Tortugas, Florida. Alpheus candei, if confirmed to be valid, can be easily distinguished from A. gallicus n. sp. by multiple characters of the frontal region (shape of the rostrum and orbital teeth, proportions of the antennular articles, much shorter scaphocerite blade, etc.) and minor cheliped (e.g. with the dactylus not expanded in A. candei vs. dorsally expanded in A. gallicus n. sp.). In addition, it appears to be a shallow-water species inhabiting coral reefs. Genbank Accession No. OUMNH.ZC.2017-01-063: MN754223, ICM 002618: MN754224., Published as part of Scioli, Justin A. & Anker, Arthur, 2020, Description of Alpheus gallicus, a new deep-water snapping shrimp from Galicia Bank, northeastern Atlantic (Malacostraca, Decapoda, Alpheidae), pp. 347-358 in Zootaxa 4731 (3) on pages 349-355, DOI: 10.11646/zootaxa.4731.3.4, http://zenodo.org/record/3653608, {"references":["Cartes, J. E., Papiol, V., Frutos, I., Macpherson, E., Gonzalez-Pola, C., Punzon, A., Valeiras, X. & Serrano, A. (2014) Distribution and biogeographic trends of decapod assemblages from Galicia Bank (NE Atlantic) at depths between 700 and 1800 m, with connexions to regional water masses. Deep Sea Research II, 106, 165 - 178. https: // doi. org / 10.1016 / j. dsr 2.2013.09.034","Hailstone, S. (1835) The species of crustaceous animals discovered and described by Mr. Hailstone, and illustrated and annotated upon by Mr. Westwood. The Magazine of Natural History and Journal of Zoology, Botany, Mineralogy, Geology, and Meteorology, 8, 394 - 395.","Garcia Raso, J. E. (1996) Crustacea Decapoda (excl. Sergestidae) from Ibero-Moroccan waters. Results of Balgim- 84 expedition. Bulletin of Marine Science, 58, 730 - 752.","Coutiere, H. (1905) Les Alpheidae. In: Gardiner, J. S. (Ed.), The fauna and geography of the Maldive and Laccadive Archipelagoes. Being the account of the work carried on and of the collections made by an expedition during the years 1899 and 1900. University Press, Cambridge, pp. 852 - 921, pls. 70 - 87.","Anker, A. & Nizinski, M. (2011) Description of a new deep-water species of Alpheus Fabricius, 1798 from the Gulf of Mexico (Crustacea, Decapoda, Alpheidae). Zootaxa, 2925 (1), 49 - 56. https: // doi. org / 10.11646 / zootaxa. 2925.1.5","Burukovsky, R. N. (1990) Shrimps from the Sala-y-Gomez and Nazca Ridges [in Russian]. Trudy Instituta Okeanologii Akademii Nauk SSSR, 124, 187 - 217.","Coutiere, H. (1897) Note sur quelques Alphees nouveaux. Bulletin du Museum d'Histoire Naturelle, 3 (7), 303 - 306. https: // doi. org / 10.5962 / bhl. part. 19261","Crosnier, A. & Forest, J. (1966) Crustaces decapodes: Alpheidae. In: Campagnes de la Calypso dans le Golfe de Guinee et aux Iles Principe, Sao Tome et Annobon (1956), et Campagne aux Iles du Cap Vert (1959). Part 19. Resultats Scientifiques des Campagnes de la Calypso 7 (27). Annales de l'Institut Oceanographique de Monaco, 44, 199 - 314.","Holthuis, L. B. (1951) The caridean Crustacea of tropical west Africa. Atlantide Report Scientific Results of the Danish Expedition to the Coasts of Tropical West Africa 1945 - 1946, 2, 7 - 187.","Chace, F. A. Jr. (1966) Decapod crustaceans from St. Helena Island, South Atlantic. Proceedings of the United States National Museum, 118, 622 - 662. https: // doi. org / 10.5479 / si. 00963801.118 - 3536.623","Soledade, G. O., Scioli, J. A., Terossi, M., Mantelatto, F. L. & Almeida, A. O. (2019) Congruence between morphology and molecular data supports the distinction between the snapping shrimps Alpheus macrocheles and A. amblyonyx (Caridea, Alpheidae). Zootaxa, 4624 (1), 1 - 22. https: // doi. org / 10.11646 / zootaxa. 4624.1.1","Anker, A. & De Grave, S. (2012) Description of Alpheus cedrici sp. n., a strikingly coloured snapping shrimp (Crustacea, Decapoda, Alpheidae) from Ascension Island, central Atlantic Ocean. ZooKeys, 183, 1 - 15. https: // doi. org / 10.3897 / zookeys. 183.3073","Christoffersen, M. L. (1979) Campagne de la Calypso au large des cotes Atlantiques de l'Amerique du Sud (1961 - 1962). I. Decapod Crustacea: Alpheoidea. Annales de l'Institut Oceanographique, 55, 297 - 377.","Banner, A. H. (1953) The Crangonidae, or snapping shrimp, of Hawaii. Pacific Science, 7, 3 - 147.","Banner, A. H. (1959) Contributions to the knowledge of the alpheid shrimp of the Pacific Ocean. Part IV. Various small collections from the Central Pacific area, including supplementary notes on alpheids from Hawaii. Pacific Science, 13, 130 - 155.","Banner, D. M. & Banner, A. H. (1982) The alpheid shrimp of Australia. Part III: The remaining alpheids, principally the genus Alpheus and the family Ogyrididae. Records of the Australian Museum, 34, 1 - 357. https: // doi. org / 10.3853 / j. 0067 - 1975.34.1982.434","Coutiere, H. (1910) The snapping shrimps (Alpheidae) of the Dry Tortugas, Florida. Proceedings of the United States National Museum, 37, 485 - 487. https: // doi. org / 10.5479 / si. 00963801.37 - 1716.485"]}

Alpheus cf. floridanus Kingsley, 1878 Material examined. Petro-UFS���14 (2 M, 2 F, 1 OF); size range: 6.31 ��� CL ��� 8.23 mm; average: CL = 7.32 �� 8.23 mm; CZUFS CRU- 00030. Penaeid���1 M; CL: 6.33 mm; CZUFS CRU- 00344. Stations. Petro-UFS���8, 14, and 15; Penaeid���6. Distribution. Western Atlantic���USA (Bahamas, south of Florida), Mexico, Veracruz and Quintana Roo, Antilles, from Cuba to Cura��ao, and Brazil (Atol das Rocas, and from Piau�� to Rio Grande do Sul) (Christoffersen 1979; Mart��nez-Iglesias et al. 1996; Almeida et al. 2007 b, 2018; Soledade & Almeida 2013). Remarks. Alpheus floridanus is a species complex (Anker 2001; Soledade & Almeida 2013) and requires a revision to be better elucidated. Therefore, the species is being mentioned as A. cf. floridanus. Records from the Eastern Pacific refer to other species (Soledade & Almeida 2013). Previous records in Sergipe. Coelho et al. (1983)., Published as part of Mendon��a, Luana M. C., Guimar��es, Carmen R. P., Santos, Rafael C., Alves, Douglas F. R., Barros-Alves, Samara P., Silva, Sonja L. R. & Hirose, Gustavo L., 2019, Decapod crustaceans from the continental shelf of Sergipe, northeastern Brazil, pp. 301-344 in Zootaxa 4712 (3) on page 327, DOI: 10.11646/zootaxa.4712.3.1, http://zenodo.org/record/3586316, {"references":["Kingsley, J. S. (1878) Notes on the North American Caridea in the Museum of the Peabody Academy of Science at Salem, Mass. Proceedings of the Academy of Natural Sciences of Philadelphia, 30, 89 - 98. Avaiable from: https: // www. jstor. org / stable / 4060370 (accessed 23 September 2019)","Christoffersen, M. L. (1979) Decapod Crustacea: Alpheoidea. Cmapagne de la Calypso au large des cotes atlantiques de L`Amerique Du Sud (1961 - 1962). I. 36, Annales de l`Institite Oceonographic, Monaco, 55 (Supplement), 297 - 377.","Martinez-Iglesias, J. C., Rios, R. & Carvacho, A. (1996) Las especies del genero Alpheus (Decapoda: Alpheidae) de Cuba. Revista de Biologia Tropical, 44 (3), 401 - 429.","Almeida, A. O., Guerrazzi, M. C. & Coelho, P. A. (2007 b) Stomatopod and decapod crustaceans from Camamu Bay, state of Bahia, Brazil. Zootaxa, 1553 (1), 1 - 45. https: // doi. org / 10.11646 / zootaxa. 1553.1.1","Almeida, A. O., Terossi, M., Buranelli, R. C., Castilho, A. L., Costa, R. C., Zara, F. J. & Mantelatto, F. L. (2018) Checklist of decapods (Crustacea) from the coast of Sao Paulo State (Brazil) supported by integrative molecular and morphological data: II. Infraorder Caridea: family Alpheidae. Zootaxa, 4450 (3), 331 - 358. https: // doi. org / 10.11646 / zootaxa. 4450.3.2","Soledade, G. O. & Almeida, A. O. (2013) Snapping shrimps of the genus Alpheus Fabricius from Brazil (Caridea: Alpheidae): updated checklist and key for identification. Nauplius, 21 (1), 89 - 122. https: // doi. org / 10.1590 / S 0104 - 64972013000100010","Anker, A. (2001) Two new species of snapping shrimps from the indo-pacific, with remarks on colour patterns and sibling spesies in Alpheidae (Crustacea: Caridae). The Raffles Bulletin of Zoology, 49 (1), 57 - 72.","Coelho, P. A., Ramos-Porto, M. & Calado, T. C. S. (1983) Litoral de Alagoas e Sergipe: Decapoda. Anais da Sociedade Nordestina de Zoologia, 1 (1), 133 - 155."]}

Alpheus cf. packardii Kingsley, 1880 (Figure 7B) Material examined. Petro-UFS—2 OF; size range: 5.05 ≤ CL ≤ 6.22 mm; average: CL = 5.64 ± 0.83 mm; CZUFS CRU- 00026. Stations. Petro-UFS—5. Distribution. Western Atlantic—Bermuda, Virgin to South Carolina, Florida, Gulf of Mexico, Bahamas, Mexico (Quintana Roo and Yucatan), West Indies, Venezuela, and Brazil (Atol das Rocas, Fernando de Noronha, Trindade and Martin Vaz, from Amapá to São Paulo) (Christoffersen 1979, 1998; Rodríguez, 1980; Martínez-Iglesias et al. 1996) as Alpheus normanni (Kingsley) (Santos et al. 2012; Almeida et al. 2012, 2018; Anker et al. 2016). Ecological notes. From intertidal to a depth of 70 m. On coral reefs, sand, mud, broken shells, algae, and sponges. On crevices of coral rubble and Halimeda sp. clumps. Reported in crevices of coral rubble from southern Bahia (Christoffersen 1979; Róman-Contreras & Martínez-Mayén 2010; Almeida et al. 2012; Santos et al. 2012). Remarks. Previously recorded as Alpheus normanni (Kingsley). Later A. packardii was considered a junior synonym of A. normanni (Chace 1937, 1972; Christoffersen 1979). Kim & Abele (1988) removed A. packardii from the synonymy of A. normanni. However, the taxonomy of A. normanni and A. packardii remains unsettled due to the presence of cryptic taxa on both sides of the Americas, and, the true identity of the material from Brazil remains unknown (Williams et al. 2001; Almeida et al. 2012; Soledade & Almeida 2013), requiring a taxonomic revision. Because of that the specie is reported as A. cf. packardii for Brazilian waters. Found in the present study associated with algae banks. Previous records in Sergipe. None., Published as part of Mendonça, Luana M. C., Guimarães, Carmen R. P., Santos, Rafael C., Alves, Douglas F. R., Barros-Alves, Samara P., Silva, Sonja L. R. & Hirose, Gustavo L., 2019, Decapod crustaceans from the continental shelf of Sergipe, northeastern Brazil, pp. 301-344 in Zootaxa 4712 (3) on pages 327-328, DOI: 10.11646/zootaxa.4712.3.1, http://zenodo.org/record/3586316, {"references":["Kingsley, J. S. (1880) On a collection of Crustacea from Virginia, North Carolina, and Florida, with a revision of the genera of Crangonidae and Palaemonidae. Proceedings of the Academy of Natural Sciences of Philadelphia, 31 (for 1879), 383 - 427. https: // doi. org / 10.5962 / bhl. title. 62608","Christoffersen, M. L. (1979) Decapod Crustacea: Alpheoidea. Cmapagne de la Calypso au large des cotes atlantiques de L`Amerique Du Sud (1961 - 1962). I. 36, Annales de l`Institite Oceonographic, Monaco, 55 (Supplement), 297 - 377.","Christoffersen, M. L. (1998) Malacostraca. Eucarida. Caridea. Crangonidae and Alpheidae (Except Glyphocrangonidae and Crangonidae). In: Young, P. S. (Ed.), Catalogue of Crustacea of Brazil. Museu Nacional, Rio de Janeiro, pp. 351 - 372.","Rodriguez, G. (1980) Los Crustaceos Decapodos de Venezuela. Instituto Venezuelano de Investigaciones Cientificas, Caracas, 494 pp.","Martinez-Iglesias, J. C., Rios, R. & Carvacho, A. (1996) Las especies del genero Alpheus (Decapoda: Alpheidae) de Cuba. Revista de Biologia Tropical, 44 (3), 401 - 429.","Santos, P. S., Soledade, G. O. & Almeida, A. O. (2012) Decapod crustaceans on dead coral from reef areas on the coast of Bahia, Brazil. Nauplius, 20 (2), 145 - 169. https: // doi. org / 10.1590 / S 0104 - 64972012000200007","Almeida, A. O., Boehs, G., Araujo-Silva, C. L. & Bezerra, L. E. A. (2012) Shallow-water caridean shrimps from southern Bahia, Brazil, including the first record of Synalpheus ul (Rios & Duffy, 2007) (Alpheidae) in the southwestern Atlantic Ocean. Zootaxa, 3347 (1), 1 - 35. https: // doi. org / 10.11646 / zootaxa. 3347.1.1","Almeida, A. O., Terossi, M., Buranelli, R. C., Castilho, A. L., Costa, R. C., Zara, F. J. & Mantelatto, F. L. (2018) Checklist of decapods (Crustacea) from the coast of Sao Paulo State (Brazil) supported by integrative molecular and morphological data: II. Infraorder Caridea: family Alpheidae. Zootaxa, 4450 (3), 331 - 358. https: // doi. org / 10.11646 / zootaxa. 4450.3.2","Anker, A., Tavares, M. & Mendonca, J. B. (2016) Alpheid shrimps (Decapoda: Caridea) of the Trindade & Martin Vaz Archipelago, off Brazil, with new records, description of a new species of Synalpheus and remarks on zoogeographical patterns in the oceanic islands of the tropical southern Atlantic. Zootaxa, 4138 (1), 1 - 58. https: // doi. org / 10.11646 / zootaxa. 4138.1.1","Roman-Contreras, R. & Martinez-Mayen, M. (2010) Notes on some alpheid shrimps (Decapoda: Caridea) of Thalassia testudinum meadows, from the central-Southern Mexican Caribbean. Hidrobiologica, 20 (3), 222 - 229.","Chace Jr., F. A. (1937) The Templeton Crocker Expedition, Part 7: Caridean Decapod Crustacea from the Gulf of California and the West Coast of Lower California. Zoologica, 22 (2), 109 - 138.","Chace Jr., F. A. (1972) The shrimps of the Smithsonian-Bredin Caribbean Expeditions with a summary of the West Indian shallow-water species (Crustacea: Decapoda: Natantia). Smithsonian Contributions to Zoology, 98, 1 - 179. https: // doi. org / 10.5479 / si. 00810282.98","Kim, W. & Abele, L. G. (1988) The Snapping Shrimp Genus Alpheus from the Eastern Pacific (Decapoda: Caridea: Alpheidae). Smithsonian Contributions to Zoology, 454, 1 - 119. https: // doi. org / 10.5479 / si. 00810282.454","Williams, S. T., Knowlton, N., Weigt, L. A. & Jara, J. A. (2001) Evidence for three major clades within the snapping shrimp genus Alpheus inferred from nuclear and mitochondrial gene sequence data. Molecular Phylogenetics and Evolution, 20, 375 - 389. https: // doi. org / 10.1006 / mpev. 2001.0976","Soledade, G. O. & Almeida, A. O. (2013) Snapping shrimps of the genus Alpheus Fabricius from Brazil (Caridea: Alpheidae): updated checklist and key for identification. Nauplius, 21 (1), 89 - 122. https: // doi. org / 10.1590 / S 0104 - 64972013000100010"]}

Alpheus ramosportoae sp. nov. urn:lsid:zoobank.org:act: B9 DFFA 34-24 DA- 4672-8A56-A474B3406B71 Figs 1–4 Alpheus macrocheles – Ramos-Porto 1979: 118, figs. 1–6 [not Alpheus macrocheles (Hailstone, 1835)]. (?) Alpheus macrocheles – Coelho & Ramos-Porto 1980: 135; 1995: 116 — Coelho et al. 1980: 55; 1986: 84 — Fausto Filho 1980: 113 — Ramos-Porto et al. 1996: 219 — Christoffersen 1998: 359 — Barros & Pimentel 2001: 21 — Guterres et al. 2005: 231, fig. 1 — Coelho et al. 2006: 51 — Coelho Filho 2006: 8 — Alves et al. 2008: 49 — Souza et al. 2011: 47 — Almeida et al. 2012: 27 — Soledade & Almeida 2013: 101 [not Alpheus macrocheles (Hailstone, 1835)]. Diagnosis Rostrum without setae, reaching well beyond half length of first segment of antennular peduncle; scaphocerite with lateral margin slightly concave, lateral tooth strong, surpassing distal margin of blade; blade reaching middle of third article of antennular peduncle; distal tooth distinctly exceeding distal margin of peduncle. Major cheliped with ischium armed with four spiniform setae; ventromesial margin of merus with five spiniform setae similar in size and shape, spaced almost equidistantly along length of mesial surface of merus; dactylus flattened, laterally twisted, dorsally convex and slightly bulbous distally; plunger developed, with anterior surface distinctly angular. Minor cheliped with ischium armed with four spiniform setae; ventromesial margin of merus with seven spiniform setae similar in size and shape, spaced almost equidistantly along length of mesial surface of merus; dactylus with strong crest on extensor margin. Second pereiopod elongated, slender; ischium slightly longer than merus; carpus with five articles with ratio approximately equal to 4: 2: 1: 1.5: 2; distolateral margin of uropodal exopod with one mesial tooth and one lateral tooth adjacent to spiniform seta. Etymology The name of this new species is our tribute to the Brazilian carcinologist Dr Marilena Ramos-Porto, retired professor of the Universidade Federal Rural de Pernambuco, Recife. Dr Marilena contributed substantially to the knowledge of the caridean shrimps of the northern and northeastern Brazil. Material examined Holotype BRAZIL • ♂; Pernambuco, off Recife, approximately 20 nautical miles from the coast, REC I, dredge 4; 8°08′51.5″ S, 34°34′08.0″ W; 65 m depth; 7 Feb. 2018; agglomerations of sponges, rhodoliths and calcareous algae; GenBank 16S gene MK918616; MOUFPE 19470. Paratypes BRAZIL – Ceará • 1 ♂; Ceará, seamounts of the North Chain, REVIZEE NE III, stn 75A; 54 m depth; 7 Apr. 1998; calcareous algae; GenBank 16S gene MK918615; MOUFPE 13703. – Pernambuco • 1 ♂; exact locality unknown; 31–33 m depth; 7 Mar. 1969; calcareous algae; MOUFPE 8826 • 2 ♂♂; exped. Pesquisador IV, stn 5; 54 m depth; 12 Apr. 1971; MOUFPE 8856 • 1 ♀; off Recife, REC I, dredge 6; 8°09′06.8″ S, 34°34′28.4″ W; 53 m depth; 7 Feb. 2018; rhodolite crevice; GenBank 16S gene MK918618; CCDB 6120 • 1 ♀ ov; off Recife, REC I, dredge 3; 8°08′44.2″ S, 34°34′23.2″ W; 55 m depth; 7 Feb. 2018; rhodolith crevice; GenBank 16S gene MK918617; MZUSP 39146. Additional material BRAZIL – Amapá • 3 ♂♂; exped. GEOMAR II, stn 116; 81.5 m depth; 13 Sep. 1970; calcareous algae; MOUFPE 8848 – Pará • 1 ♂, 1 ♀ ov; exped. GEOMAR II, stn 2441; 1970; MOUFPE 8855 • 1 ♂, 1 ♀ ov; exped. GEOMAR II, stn 2528; 60 m depth; 1971; MOUFPE 8836 – Maranhão • 1 ♀; Tutóia, RV Almirante Saldanha, stn 1732A; 69 m depth; 30 Oct. 1967; calcareous algae; MOUFPE 8829 • 9 NI; RV Almirante Saldanha, stn 1750; 52 m depth; 6 Nov. 1967; calcareous algae; MOUFPE 8830 • 1 ♂, 1 ♀; São Luis, RV Almirante Saldanha, stn 1875; 49 m depth; 23 Apr. 1968; calcareous algae; MOUFPE 8831 • 1 NI; Tutóia, RV Almirante Saldanha, stn 1813; 83 m depth; 28 Nov. 1967; calcareous algae; MOUFPE 8837 • 1 ♀; RV Almirante Saldanha, stn 1751; 44 m depth; 6 Nov. 1967; calcareous algae; MOUFPE 8845 • 1 ♂; RV Almirante Saldanha, stn 1750; 52 m depth; 6. Nov. 1967; calcareous algae; MOUFPE 8847 – Ceará • 1 ♂; REVIZEE NE III, stn 72A; 2°4′48’’ S, 38°12′0’’ W; 8 Jun. 1998; MOUFPE 13647. Comparative material analyzed Alpheus amblyonyx Chace, 1972 FRANCE – Guadeloupe • 1 ♂; KARUBENTHOS, stn GB20; 16 m depth; 18 May 2012; MNHN- IU-12112 • 1 ♀ ov; KARUBENTHOS, stn GB24; 16 m depth; 18 May 2012; GenBank 16S gene MK714203; MNHN-IU-12026. MEXICO • 1 NI; Quintana Roo, southeast of Arrecife Chinchorro; 28 Aug. 1990; GenBank 16S gene MK714201; UNAM-CNCR 21271. Alpheus macrocheles (Hailstone, 1835) ANGOLA • 3 ♀♀ (2 ov), 1 ♂; Cuanza River; 31 Jan. 1949; RMNH. Crus.D. 7821. CAPE VERDE • 1 ♀ ov, 3 NI; “Tydeman” Cancap–VI, Cape Verde Islands exped., stn 6.109, off Santa Luzia; 16°44′ N, 24°46′ W; 55–80 m depth; 16 Jun. 1982; calcareous algae; RMNH. Crus.D. 51345 • 1 ♂; Tarrafal Bay; 1959; MNHN 3163. FRANCE • 1 ♀; Roscoff; RMNH. Crus.D. 34508 • 1 ♂; Roscoff; RMNH. Crus.D. 29789 • 1 ♂; Guéthary; 27 Oct. 1947; RMNH. Crus.D. 1869. MOROCCO • 1 ♀; 33°50′ N, 07°07′ W; 25 Jan. 1956; RMNH. Crus.D. 34495 • 1 ♂; Témara; 1952; RMNH. Crus.D. 34496 • 1 ♂; Témara; 1954; RMNH. Crus.D. 34497. PORTUGAL – Azores • 1 ♂, 1 NI; stn 5.142; 108–118 m depth; 7 Jun. 1981; GenBank 16S gene MK714206; RMNH. Crus.D. 51342. – Madeira • 1 ♂; Cais de Porto Novo; 18 Aug. 1993; under rocks; RMNH. Crus.D. 42706. SPAIN – Canary Islands • 1 ♂; southeast of Lanzarote, stn 4.070; 41–50 m depth; 20 May 1980; sand and calcareous algae; GenBank 16S gene MK714207; RMNH. Crus.D. 51363. Description CARAPACE. Smooth, frontal margin with well-developed sharp rostrum; orbital hoods with well-developed acute teeth originating from anterior margin of orbital hoods (Fig. 1A); rostrum slightly flattened dorsally, distinctly longer than wide, distally tapering and ending in acute point, without setae, reaching well beyond half-length of first article of antennular peduncle (Fig. 1 A–B); margin between orbital teeth and rostrum V-shaped with slightly rounded angle (Fig. 1A); orbital process weak. Pterygostomial angle rounded, not anteriorly protruding (Fig. 1B); cardiac notch deep. HEAD. Eyes with well-developed corneas; ocellar beak projecting, acute, visible in lateral view. Antennular peduncle moderately slender; stylocerite with acute tip reaching distal margin of first article of antennular peduncle; ventromesial carina with very large, triangular tooth bearing small acute point (Fig. 1C); second article of antennular peduncle much longer than visible part of first article, approx. three times longer than wide, twice as long as third article (Fig. 1 A–B); lateral flagellum with several groups of aesthetascs extending to article 10. Antenna with basicerite ending in sharp ventrolateral tooth, extending just beyond orbital tooth; carpocerite slightly overreaching scaphocerite and antennular peduncle; scaphocerite with lateral margin slightly concave, ending in strong distolateral tooth distinctly overreaching antennal peduncle and surpassing distal margin of blade (Fig. 1A), latter reaching to middle of third article. Mouthparts typical for Alpheus, as illustrated (Fig. 1 D–I). Third maxilliped slender; coxa with lateral plate slightly truncate distally; exopod overreaching antepenultimate article when extended; antepenultimate article somewhat flattened, approximately four times longer than wide, ventral surface sparsely setose; penultimate article about three times longer than wide, slightly broadened distally and densely setose; ultimate article unarmed, distally tapering, with dense transverse rows of long setae (Fig. 1I). CHELIPEDS. Major cheliped with ischium short, robust, ventromesial surface with four small spiniform setae (Fig. 2F); merus robust, about twice as long as wide, subtriangular in cross-section; ventrolateral margin unarmed; ventromesial margin straight, bearing five small spiniform setae, ending in robust sharp tooth; carpus short, cup-shaped, slightly compressed (Fig. 2F); lateral surface of palm with low crest (inferior crest) starting at approximately 0.6 of palm length, ending in sharp distolateral tooth, latter slightly directed laterally (Fig. 2A); mesial surface convex, smooth, without grooves (Fig. 2B); ventral surface with rounded smooth shoulder slightly projecting into adjacent deep notch, latter extending transversely to groove on mesial surface (inferior groove); dorsal margin with subcylindrical elevation (plaque crest) ending distally in large adhesive disk; distomesial surface with deep transversally notched crest (superior crest) ending in sharp tooth; distal third of ventral margin with long, robust setae extending to distomesial margin; pollex shorter than dactylus, strongly curved laterally, cutting edge slightly excavate, bearing one small rounded tooth proximally (Fig. 2A); dactylus flattened, laterally twisted, dorsally convex, slightly bulbous distally (Fig. 2A); plunger moderately developed, its proximal margin with sharp angle (Fig. 2E); adhesive disks of dactylus well-developed (Fig. 2A). Minor cheliped not sexually dimorphic; ischium, short, stout, its distomesial margin armed with four spiniform setae; merus broad, subtriangular in cross-section; ventrolateral surface unarmed; ventromesial margin slightly convex, ending in small acute tooth, with seven spiniform setae similar in size and shape, spaced almost equidistantly along entire length of mesial margin; carpus cup-shaped (Fig. 2G); chela strongly compressed; palm with grooves and notches on distal half of lateral surface (Fig. 2 C–D); lateral surface with low crest (inferior crest) starting at about middle of palm and ending in acute distolateral tooth (Fig. 2C); mesial surface convex and devoid of depressions or grooves (Fig. 2D); ventral surface smooth, ending in well-defined notch; dorsal margin also smooth, with sub-cylindrical elevation (plaque crest) ending distally in small adhesive disk; distomesial surface with high ridge ending in strong sharp tooth (superior crest); fingers as long as palm (Fig. 2 C–D); pollex lightly excavate on cutting edge; dactylus slightly flattened and twisted laterally, with broad crest on extensor margin, bearing small adhesive disk proximally (Fig. 2 C–D). PEREIOPODS. Second pereiopod elongate, slender; ischium slightly longer than merus; carpus with five subdivisions with ratio approximately equal to 4: 2: 1: 1.5: 2; chela fingers as long as palm, with small tufts of setae on distal region (Fig. 3A). Third pereiopod with ischium armed with one strong spiniform seta on ventrolateral surface (Fig. 3B); merus five times longer than broad, unarmed distoventrally; carpus slender, about half-length of merus (Fig. 3B); propodus approximately 1.2 times as long as carpus, with about nine robust spiniform setae along ventral margin and one pair of spiniform setae at propodo-dactylar articulation (Fig. 3E); dactylus slightly shorter than half-length of propodus, conical, slightly curved, acute, extensor margin with one subdistal denticle (Fig. 3E, H). Fourth pereiopod similar to third in shape and proportion of articles, slightly less robust (Fig. 3C); ischium armed with one strong spiniform seta on ventrolateral surface (Fig. 3C); propodus with about eight robust spiniform setae along ventral margin and one pair of spiniform setae at propodo-dactylar articulation, extensor margin with one subdistal denticle (Fig. 3F, I). Fifth pereiopod more slender than third and fourth (Fig. 3D); merus slightly longer than carpus; ischium with one spiniform seta; propodus with about eight spiniform setae along ventral margin and nine well-developed rows of setae distolaterally (omitted) (Fig. 3G). PLEON. Pleonites 1–4 with posteroventral margins broadly rounded, fifth pleonite slightly angular. Telson subrectangular, tapering to posterior margin; proximal margin twice as wide as distal margin; lateral margin slightly convex; dorsal surface with two pairs of spiniform setae, first pair located at telson halflength, second pair at approximately 0.7 of telson length; posterior margin broadly convex, with two pairs of spiniform setae, mesial pair about three times as long as lateral pair (Fig. 1J); anal tubercles well-developed. PLEOPODS. With sparse setae on lateral margin of protopod. First pleopod reduced; distal margin of endopod with setae. Second pleopod of male with appendix masculina subequal in length to appendix interna, not reaching distal margin of endopod, with numerous rigid setae distally. Uropod with lateral lobe of protopod ending in strong sharp tooth (Fig. 1J); diaeresis sinuous; distolateral margin of exopod with one mesial tooth and one lateral tooth flanking spiniform seta (Fig. 1K); endopod with row of spiniform setae along distal margin, mesial ones stronger than lateral ones (Fig. 1L). Color in life Body predominantly reddish; carapace uniformly reddish; pleon reddish with whitish transverse bands and whitish spots on lateral surface; major cheliped palm reddish, with three transverse white patches on both lateral and mesial surfaces, more defined on mesial surface; fingers of both chelae reddish; telson and uropods intensely reddish; protopods of uropods whitish; pereiopods (2–5) pale reddish (Fig. 4 A– C). This color pattern was homogeneous in all specimens collected. Habitat On sand mixed with calcareous algae, rhodoliths and sponges (Fig. 4 D–F), at depths ranging from 33 to 90 m (Ramos-Porto 1979, as A. macrocheles; present study). Distribution Brazil: Amapá, Pará, Maranhão, Ceará and Pernambuco (Ramos-Porto 1979, as A. macrocheles; present study). Molecular analysis The best-fit substitution model selected with a corrected Bayesian information criterion was TPM3uf+I, assuming the nucleotide frequencies A = 0.2847, C = 0.1194, G = 0.2561, T = 0.3398, replacement rates AC = 0.0010, AG = 4.5910, AT = 1, CG = 0.0010, CT = 4.5910, GT = 1, proportion of invariable sites = 0.5410. The Bayesian Inference analysis (Fig. 5) retrieved two clades, one being formed by A. amblyonyx and another including A. ramosportoae sp. nov. as sister to a clade formed by A. crockeri (Armstrong, 1941), A. puapeba and A. macrocheles. Genetic distance estimates (Table 2) support these results. The range of the genetic distance values between A. ramosportoae sp. nov. and other species was 15.9–22.9% according to the TPM3 model. Alpheus ramosportoae sp. nov., as well as A. amblyonyx and A. macrocheles, show no significant intraspecific divergence in 16S sequences.

Alpheus ramosportoae sp. nov. urn:lsid:zoobank.org:act: B9 DFFA 34-24 DA- 4672-8A56-A474B3406B71 Figs 1���4 Alpheus macrocheles ��� Ramos-Porto 1979: 118, figs. 1���6 [not Alpheus macrocheles (Hailstone, 1835)]. (?) Alpheus macrocheles ��� Coelho & Ramos-Porto 1980: 135; 1995: 116 ��� Coelho et al. 1980: 55; 1986: 84 ��� Fausto Filho 1980: 113 ��� Ramos-Porto et al. 1996: 219 ��� Christoffersen 1998: 359 ��� Barros & Pimentel 2001: 21 ��� Guterres et al. 2005: 231, fig. 1 ��� Coelho et al. 2006: 51 ��� Coelho Filho 2006: 8 ��� Alves et al. 2008: 49 ��� Souza et al. 2011: 47 ��� Almeida et al. 2012: 27 ��� Soledade & Almeida 2013: 101 [not Alpheus macrocheles (Hailstone, 1835)]. Diagnosis Rostrum without setae, reaching well beyond half length of first segment of antennular peduncle; scaphocerite with lateral margin slightly concave, lateral tooth strong, surpassing distal margin of blade; blade reaching middle of third article of antennular peduncle; distal tooth distinctly exceeding distal margin of peduncle. Major cheliped with ischium armed with four spiniform setae; ventromesial margin of merus with five spiniform setae similar in size and shape, spaced almost equidistantly along length of mesial surface of merus; dactylus flattened, laterally twisted, dorsally convex and slightly bulbous distally; plunger developed, with anterior surface distinctly angular. Minor cheliped with ischium armed with four spiniform setae; ventromesial margin of merus with seven spiniform setae similar in size and shape, spaced almost equidistantly along length of mesial surface of merus; dactylus with strong crest on extensor margin. Second pereiopod elongated, slender; ischium slightly longer than merus; carpus with five articles with ratio approximately equal to 4: 2: 1: 1.5: 2; distolateral margin of uropodal exopod with one mesial tooth and one lateral tooth adjacent to spiniform seta. Etymology The name of this new species is our tribute to the Brazilian carcinologist Dr Marilena Ramos-Porto, retired professor of the Universidade Federal Rural de Pernambuco, Recife. Dr Marilena contributed substantially to the knowledge of the caridean shrimps of the northern and northeastern Brazil. Material examined Holotype BRAZIL ��� ♂; Pernambuco, off Recife, approximately 20 nautical miles from the coast, REC I, dredge 4; 8��08���51.5��� S, 34��34���08.0��� W; 65 m depth; 7 Feb. 2018; agglomerations of sponges, rhodoliths and calcareous algae; GenBank 16S gene MK918616; MOUFPE 19470. Paratypes BRAZIL ��� Cear�� ��� 1 ♂; Cear��, seamounts of the North Chain, REVIZEE NE III, stn 75A; 54 m depth; 7 Apr. 1998; calcareous algae; GenBank 16S gene MK918615; MOUFPE 13703. ��� Pernambuco ��� 1 ♂; exact locality unknown; 31���33 m depth; 7 Mar. 1969; calcareous algae; MOUFPE 8826 ��� 2 ♂♂; exped. Pesquisador IV, stn 5; 54 m depth; 12 Apr. 1971; MOUFPE 8856 ��� 1 ♀; off Recife, REC I, dredge 6; 8��09���06.8��� S, 34��34���28.4��� W; 53 m depth; 7 Feb. 2018; rhodolite crevice; GenBank 16S gene MK918618; CCDB 6120 ��� 1 ♀ ov; off Recife, REC I, dredge 3; 8��08���44.2��� S, 34��34���23.2��� W; 55 m depth; 7 Feb. 2018; rhodolith crevice; GenBank 16S gene MK918617; MZUSP 39146. Additional material BRAZIL ��� Amap�� ��� 3 ♂♂; exped. GEOMAR II, stn 116; 81.5 m depth; 13 Sep. 1970; calcareous algae; MOUFPE 8848 ��� Par�� ��� 1 ♂, 1 ♀ ov; exped. GEOMAR II, stn 2441; 1970; MOUFPE 8855 ��� 1 ♂, 1 ♀ ov; exped. GEOMAR II, stn 2528; 60 m depth; 1971; MOUFPE 8836 ��� Maranh��o ��� 1 ♀; Tut��ia, RV Almirante Saldanha, stn 1732A; 69 m depth; 30 Oct. 1967; calcareous algae; MOUFPE 8829 ��� 9 NI; RV Almirante Saldanha, stn 1750; 52 m depth; 6 Nov. 1967; calcareous algae; MOUFPE 8830 ��� 1 ♂, 1 ♀; S��o Luis, RV Almirante Saldanha, stn 1875; 49 m depth; 23 Apr. 1968; calcareous algae; MOUFPE 8831 ��� 1 NI; Tut��ia, RV Almirante Saldanha, stn 1813; 83 m depth; 28 Nov. 1967; calcareous algae; MOUFPE 8837 ��� 1 ♀; RV Almirante Saldanha, stn 1751; 44 m depth; 6 Nov. 1967; calcareous algae; MOUFPE 8845 ��� 1 ♂; RV Almirante Saldanha, stn 1750; 52 m depth; 6. Nov. 1967; calcareous algae; MOUFPE 8847 ��� Cear�� ��� 1 ♂; REVIZEE NE III, stn 72A; 2��4���48������ S, 38��12���0������ W; 8 Jun. 1998; MOUFPE 13647. Comparative material analyzed Alpheus amblyonyx Chace, 1972 FRANCE ��� Guadeloupe ��� 1 ♂; KARUBENTHOS, stn GB20; 16 m depth; 18 May 2012; MNHN- IU-12112 ��� 1 ♀ ov; KARUBENTHOS, stn GB24; 16 m depth; 18 May 2012; GenBank 16S gene MK714203; MNHN-IU-12026. MEXICO ��� 1 NI; Quintana Roo, southeast of Arrecife Chinchorro; 28 Aug. 1990; GenBank 16S gene MK714201; UNAM-CNCR 21271. Alpheus macrocheles (Hailstone, 1835) ANGOLA ��� 3 ♀♀ (2 ov), 1 ♂; Cuanza River; 31 Jan. 1949; RMNH. Crus.D. 7821. CAPE VERDE ��� 1 ♀ ov, 3 NI; ���Tydeman��� Cancap���VI, Cape Verde Islands exped., stn 6.109, off Santa Luzia; 16��44��� N, 24��46��� W; 55���80 m depth; 16 Jun. 1982; calcareous algae; RMNH. Crus.D. 51345 ��� 1 ♂; Tarrafal Bay; 1959; MNHN 3163. FRANCE ��� 1 ♀; Roscoff; RMNH. Crus.D. 34508 ��� 1 ♂; Roscoff; RMNH. Crus.D. 29789 ��� 1 ♂; Gu��thary; 27 Oct. 1947; RMNH. Crus.D. 1869. MOROCCO ��� 1 ♀; 33��50��� N, 07��07��� W; 25 Jan. 1956; RMNH. Crus.D. 34495 ��� 1 ♂; T��mara; 1952; RMNH. Crus.D. 34496 ��� 1 ♂; T��mara; 1954; RMNH. Crus.D. 34497. PORTUGAL ��� Azores ��� 1 ♂, 1 NI; stn 5.142; 108���118 m depth; 7 Jun. 1981; GenBank 16S gene MK714206; RMNH. Crus.D. 51342. ��� Madeira ��� 1 ♂; Cais de Porto Novo; 18 Aug. 1993; under rocks; RMNH. Crus.D. 42706. SPAIN ��� Canary Islands ��� 1 ♂; southeast of Lanzarote, stn 4.070; 41���50 m depth; 20 May 1980; sand and calcareous algae; GenBank 16S gene MK714207; RMNH. Crus.D. 51363. Description CARAPACE. Smooth, frontal margin with well-developed sharp rostrum; orbital hoods with well-developed acute teeth originating from anterior margin of orbital hoods (Fig. 1A); rostrum slightly flattened dorsally, distinctly longer than wide, distally tapering and ending in acute point, without setae, reaching well beyond half-length of first article of antennular peduncle (Fig. 1 A���B); margin between orbital teeth and rostrum V-shaped with slightly rounded angle (Fig. 1A); orbital process weak. Pterygostomial angle rounded, not anteriorly protruding (Fig. 1B); cardiac notch deep. HEAD. Eyes with well-developed corneas; ocellar beak projecting, acute, visible in lateral view. Antennular peduncle moderately slender; stylocerite with acute tip reaching distal margin of first article of antennular peduncle; ventromesial carina with very large, triangular tooth bearing small acute point (Fig. 1C); second article of antennular peduncle much longer than visible part of first article, approx. three times longer than wide, twice as long as third article (Fig. 1 A���B); lateral flagellum with several groups of aesthetascs extending to article 10. Antenna with basicerite ending in sharp ventrolateral tooth, extending just beyond orbital tooth; carpocerite slightly overreaching scaphocerite and antennular peduncle; scaphocerite with lateral margin slightly concave, ending in strong distolateral tooth distinctly overreaching antennal peduncle and surpassing distal margin of blade (Fig. 1A), latter reaching to middle of third article. Mouthparts typical for Alpheus, as illustrated (Fig. 1 D���I). Third maxilliped slender; coxa with lateral plate slightly truncate distally; exopod overreaching antepenultimate article when extended; antepenultimate article somewhat flattened, approximately four times longer than wide, ventral surface sparsely setose; penultimate article about three times longer than wide, slightly broadened distally and densely setose; ultimate article unarmed, distally tapering, with dense transverse rows of long setae (Fig. 1I). CHELIPEDS. Major cheliped with ischium short, robust, ventromesial surface with four small spiniform setae (Fig. 2F); merus robust, about twice as long as wide, subtriangular in cross-section; ventrolateral margin unarmed; ventromesial margin straight, bearing five small spiniform setae, ending in robust sharp tooth; carpus short, cup-shaped, slightly compressed (Fig. 2F); lateral surface of palm with low crest (inferior crest) starting at approximately 0.6 of palm length, ending in sharp distolateral tooth, latter slightly directed laterally (Fig. 2A); mesial surface convex, smooth, without grooves (Fig. 2B); ventral surface with rounded smooth shoulder slightly projecting into adjacent deep notch, latter extending transversely to groove on mesial surface (inferior groove); dorsal margin with subcylindrical elevation (plaque crest) ending distally in large adhesive disk; distomesial surface with deep transversally notched crest (superior crest) ending in sharp tooth; distal third of ventral margin with long, robust setae extending to distomesial margin; pollex shorter than dactylus, strongly curved laterally, cutting edge slightly excavate, bearing one small rounded tooth proximally (Fig. 2A); dactylus flattened, laterally twisted, dorsally convex, slightly bulbous distally (Fig. 2A); plunger moderately developed, its proximal margin with sharp angle (Fig. 2E); adhesive disks of dactylus well-developed (Fig. 2A). Minor cheliped not sexually dimorphic; ischium, short, stout, its distomesial margin armed with four spiniform setae; merus broad, subtriangular in cross-section; ventrolateral surface unarmed; ventromesial margin slightly convex, ending in small acute tooth, with seven spiniform setae similar in size and shape, spaced almost equidistantly along entire length of mesial margin; carpus cup-shaped (Fig. 2G); chela strongly compressed; palm with grooves and notches on distal half of lateral surface (Fig. 2 C���D); lateral surface with low crest (inferior crest) starting at about middle of palm and ending in acute distolateral tooth (Fig. 2C); mesial surface convex and devoid of depressions or grooves (Fig. 2D); ventral surface smooth, ending in well-defined notch; dorsal margin also smooth, with sub-cylindrical elevation (plaque crest) ending distally in small adhesive disk; distomesial surface with high ridge ending in strong sharp tooth (superior crest); fingers as long as palm (Fig. 2 C���D); pollex lightly excavate on cutting edge; dactylus slightly flattened and twisted laterally, with broad crest on extensor margin, bearing small adhesive disk proximally (Fig. 2 C���D). PEREIOPODS. Second pereiopod elongate, slender; ischium slightly longer than merus; carpus with five subdivisions with ratio approximately equal to 4: 2: 1: 1.5: 2; chela fingers as long as palm, with small tufts of setae on distal region (Fig. 3A). Third pereiopod with ischium armed with one strong spiniform seta on ventrolateral surface (Fig. 3B); merus five times longer than broad, unarmed distoventrally; carpus slender, about half-length of merus (Fig. 3B); propodus approximately 1.2 times as long as carpus, with about nine robust spiniform setae along ventral margin and one pair of spiniform setae at propodo-dactylar articulation (Fig. 3E); dactylus slightly shorter than half-length of propodus, conical, slightly curved, acute, extensor margin with one subdistal denticle (Fig. 3E, H). Fourth pereiopod similar to third in shape and proportion of articles, slightly less robust (Fig. 3C); ischium armed with one strong spiniform seta on ventrolateral surface (Fig. 3C); propodus with about eight robust spiniform setae along ventral margin and one pair of spiniform setae at propodo-dactylar articulation, extensor margin with one subdistal denticle (Fig. 3F, I). Fifth pereiopod more slender than third and fourth (Fig. 3D); merus slightly longer than carpus; ischium with one spiniform seta; propodus with about eight spiniform setae along ventral margin and nine well-developed rows of setae distolaterally (omitted) (Fig. 3G). PLEON. Pleonites 1���4 with posteroventral margins broadly rounded, fifth pleonite slightly angular. Telson subrectangular, tapering to posterior margin; proximal margin twice as wide as distal margin; lateral margin slightly convex; dorsal surface with two pairs of spiniform setae, first pair located at telson halflength, second pair at approximately 0.7 of telson length; posterior margin broadly convex, with two pairs of spiniform setae, mesial pair about three times as long as lateral pair (Fig. 1J); anal tubercles well-developed. PLEOPODS. With sparse setae on lateral margin of protopod. First pleopod reduced; distal margin of endopod with setae. Second pleopod of male with appendix masculina subequal in length to appendix interna, not reaching distal margin of endopod, with numerous rigid setae distally. Uropod with lateral lobe of protopod ending in strong sharp tooth (Fig. 1J); diaeresis sinuous; distolateral margin of exopod with one mesial tooth and one lateral tooth flanking spiniform seta (Fig. 1K); endopod with row of spiniform setae along distal margin, mesial ones stronger than lateral ones (Fig. 1L). Color in life Body predominantly reddish; carapace uniformly reddish; pleon reddish with whitish transverse bands and whitish spots on lateral surface; major cheliped palm reddish, with three transverse white patches on both lateral and mesial surfaces, more defined on mesial surface; fingers of both chelae reddish; telson and uropods intensely reddish; protopods of uropods whitish; pereiopods (2���5) pale reddish (Fig. 4 A��� C). This color pattern was homogeneous in all specimens collected. Habitat On sand mixed with calcareous algae, rhodoliths and sponges (Fig. 4 D���F), at depths ranging from 33 to 90 m (Ramos-Porto 1979, as A. macrocheles; present study). Distribution Brazil: Amap��, Par��, Maranh��o, Cear�� and Pernambuco (Ramos-Porto 1979, as A. macrocheles; present study). Molecular analysis The best-fit substitution model selected with a corrected Bayesian information criterion was TPM3uf+I, assuming the nucleotide frequencies A = 0.2847, C = 0.1194, G = 0.2561, T = 0.3398, replacement rates AC = 0.0010, AG = 4.5910, AT = 1, CG = 0.0010, CT = 4.5910, GT = 1, proportion of invariable sites = 0.5410. The Bayesian Inference analysis (Fig. 5) retrieved two clades, one being formed by A. amblyonyx and another including A. ramosportoae sp. nov. as sister to a clade formed by A. crockeri (Armstrong, 1941), A. puapeba and A. macrocheles. Genetic distance estimates (Table 2) support these results. The range of the genetic distance values between A. ramosportoae sp. nov. and other species was 15.9���22.9% according to the TPM3 model. Alpheus ramosportoae sp. nov., as well as A. amblyonyx and A. macrocheles, show no significant intraspecific divergence in 16S sequences., Published as part of Soledade, Guidomar O., Terossi, Mariana, Scioli, Justin A., Mantelatto, Fernando Luis & Almeida, Alexandre O., 2019, A new western Atlantic snapping shrimp of the Alpheus macrocheles group (Caridea, Alpheidae) revealed by morphological, molecular and color data, pp. 1-21 in European Journal of Taxonomy 581 on pages 5-11, DOI: 10.5852/ejt.2019.581, http://zenodo.org/record/3660185, {"references":["Ramos-Porto M. 1979. Ocorrencia de Alpheus macrocheles (Hailstone) (Crustacea: Decapoda: Alpheidae) no norte e nordeste do Brasil. Trabalhos Oceanograficos da Universidade Federal de Pernambuco 14: 117 - 130. https: // doi. org / 10.5914 / tropocean. v 14 i 1.2570","Hailstone S. 1835. The species of crustaceous animals discovered and described by Mr. Hailstone, and illustrated and annotated upon by Mr. Westwood. The Magazine of Natural History and Journal of Zoology, Botany, Mineralogy, Geology and Meteorology 8: 394 - 395.","Coelho P. A. & Ramos-Porto M. A. 1980. Crustaceos decapodos da costa do Maranhao, Brasil. Boletim do Instituto Oceanografico 29: 135 - 138. https: // doi. org / 10.1590 / S 0373 - 55241980000200028","Coelho P. A. & Ramos-Porto M. A. 1995. Distribuicao ecologica dos crustaceos decapodos marinhos do Nordeste do Brasil. Trabalhos Oceanograficos da Universidade Federal de Pernambuco 23: 113 - 127. https: // doi. org / 10.5914 / tropocean. v 23 i 1.2682","Coelho P. A., Ramos-Porto M. & Calado T. C. S. 1986. Litoral do Rio Grande do Norte: Decapoda. Caderno Omega da Universidade Federal Rural de Pernambuco, Serie Ciencias Aquaticas, Recife 2: 79 - 105.","Fausto Filho J. 1980. Crustaceos estomatopodos e decapodos dos substratos de cascalho do nordeste brasileiro. Ciencias Agronomicas 10: 109 - 124.","Ramos-Porto M., Torres M. F. A. & Viana G. F. S. 1996. Crustaceos decapodos coletados durante as expedicoes Nordeste III e Pavasas I (Penaeidea e Caridea). Trabalhos Oceanograficos da Universidade Federal de Pernambuco 24: 211 - 227. https: // doi. org / 10.5914 / tropocean. v 24 i 1.2709","Christoffersen M. L. 1998. Malacostraca. Eucarida. Caridea. Crangonoidea and Alpheoidea (Except Glyphocrangonidae and Crangonidae). In: Young P. S. (ed) Catalogue of Crustacea of Brazil, Serie livros n. 6: 351 - 372. Museu Nacional, Rio de Janeiro.","Barros M. P. & Pimentel F. R. 2001. A fauna de Decapoda (Crustacea) do Estado do Para, Brasil: Lista preliminar das especies. Boletim do Museu Paraense Emilio Goeldi, serie Zoologia 17: 15 - 41.","Guterres L. F. R., Melo G. A. S. & Guterres P. M. C. 2005. Novos registros da ocorrencia de Alpheus macrocheles (Crustacea, Caridea, Alpheidae) na costa do Brasil. Biociencias 13: 231 - 233.","Coelho P. A., Almeida A. O., Souza-Filho J. F., Bezerra L. E. A. & Giraldes B. W. 2006. Diversity and distribution of the marine and estuarine shrimps (Dendrobranchiata, Stenopodidea and Caridea) from North and Northeast Brazil. Zootaxa 1221: 41 - 62. https: // doi. org / 10.11646 / zootaxa. 1221.1.5","Coelho Filho P. A. 2006. Checklist of the Decapods (Crustacea) from the outer continental shelf and seamounts from Northeast of Brazil - REVIZEE Program (NE III). Zootaxa 1184: 1 - 27. https: // doi. org / 10.11646 / zootaxa. 1184.1.1","Alves M. L., Ramos-Porto M. & Viana G. F. S. 2008. Checklist of the decapods (Crustacea) from the Fernando de Noronha Archipelago, Brazil. Zootaxa 1881: 43 - 68. https: // doi. org / 10.11646 / zootaxa. 1881.1.2","Souza J. A. F., Schwamborn R., Barreto A. V., Farias I. D., Fernandes L. M. G. & Coelho P. A. 2011. Marine and estuarine shrimps (Dendrobranchiata, Stenopodidea, and Caridea), of Pernambuco state (Brazil) and northeastern Brazilian oceanic islands. Atlantica 33: 33 - 63. https: // doi. org / 10.5088 / atl. 2011.33.1.33","Almeida A. O., Boehs G., Araujo-Silva C. L. & Bezerra L. E. A. 2012. Shallow-water caridean shrimps from southern Bahia, Brazil, including the first record of Synalpheus ul (Rios & Duffy, 2007) (Alpheidae) in the southwestern Atlantic Ocean. Zootaxa 3347: 1 - 35. https: // doi. org / 10.11646 / zootaxa. 3347.1.1","Soledade G. O. & Almeida A. O. 2013. Snapping shrimps of the genus Alpheus Fabricius, 1798 from Brazil (Caridea: Alpheidae): updated checklist and key for identification. Nauplius 21: 89 - 122. https: // doi. org / 10.1590 / S 0104 - 64972013000100010","Chace Jr. F. A. 1972. The shrimps of the Smithsonian-Bredin Caribbean Expeditions with a summary of the West Indian shallow-water species (Crustacea: Decapoda: Natantia). Smithsonian Contributions to Zoology 98: 1 - 179. https: // doi. org / 10.5479 / si. 00810282.98"]}

Alpheus peasei (Armstrong, 1940) (Figs. 1, 2) Crangon peasei Armstrong, 1940: 1. [Castle Harbor Reefs, Bermuda] Alpheus peasei — Chace 1956: 146; 1972: 69; Hazlett 1962: 82; Hazlett & Winn 1962: 26; Knowlton & Moulton 1963: 313, fig. 3; Pequegnat & Ray 1974: 247; Ray 1974: 120, figs. 100–104; Abele 1976: 273, tab. 9; Carvacho 1979: 454; Abele & Kim 1986: 197, 210–211, figs. A–C; Rodriguez 1986: 152; Sterrer, 1986: 325, text-pl. 105; Martinez-Iglesias et al. 1993: 11; 1996: 34; 1997: 424, fig. 15; Santos et al. 2012: 152, fig. 3J; Soledade & Almeida 2013: 104, fig. 6C; Cunha et al. 2015: 48, fig. 1; Anker et al. 2016: 25, figs. 7A, B, 8A, B; Pachelle et al. 2016: 9. Holotype. Bermuda, Castle Harbor Reefs, AMNH 8184. Not examined. Material examined. Bermuda. Paratypes: 2 males, 2 ovig. females, Castle Harbor, VIII.1938, lot 123, AMNH 8186 ; 9 males, 4 (3 ovig.) females, Grazbury’s Island, VIII.1938, lot 127, AMNH 8186; 5 males, 4 (2 ovig.) females, Castle Harbor, on reef, VIII.1938, lot 133, AMNH 8186; 2 males, St. Davids, on reef, VIII.1938, lot 138, AMNH 8186. Other material. Bermuda. 4 males, 4 (2 ovig.) females, Ferry Beach, on rocks, coll. B.A. Hazlett, 02.VIII.1960, USNM 107080 ; 2 ovig. females, South Shore, on rocks, coll. B.A. Hazlett, 10.VIII.1960, USNM 107079; 2 males, Castle Harbor, in calcareous rocks, coll. B.A. Hazlett, 21.VII.1960, USNM 107081; 2 males, 1 ovig. female, Castle Harbor, off Charles Island, in calcareous rocks, coll. J.M. Moulton, 25.VII.1958, USNM 102370. United States of America. 1 male, Florida, St. Lucie County, in limestone reef and Phragmatopoma sp., 6 m, USNM 170085. Mexico. 1 male, 1 ovig. female, 1 juv, Quintana Roo, Ascension Bay, Center of Nicchehabin Reef, coll. E.L. Bousfield, 1–2 m, 14.IV.1960, USNM 136040; 1 male, Quintana Roo, Puerto Morelos, María Irene Beach, CNCR 9631; 1 male, Quintana Roo, Isla Cozumel, Playa Aventura, CNCR 20292; 1 ovig. female, Quintana Roo, Arrecife Cacarracas, Frente Puerto Morelos, CNCR 5733; 1 ni, Quintana Roo, Punta El Placer, Campaña Edwin Link, CNCR 10872; 1 ni, Quintana Roo, Punta Pulticub, CNCR 20306. Belize. 1 male, Carrie Bow Cay, coll. K. Sandved, 1 m, IV.1974, USNM 184488. Costa Rica. 2 males, 1 ovig. female, Parque Nacional Cahuita, Limón, Puerto Vargas, colls. R. Heard, L. Rodriguez & R. Vargas, 19.XI.1999, MZUCR 2312-33. Panama. 1 male, Isla Grande, off W point, in coral rocks, colls. A. Anker & C. Hurt, 1–2 m, 06.X.2005, MZUSP 33123; 1 male, Bocas del Toro, Bocas del Drago, colls. F.L. Mantelatto, A. Baeza, I. Miranda & M. Terossi, 17. II.2009, CCDB 3493; Colombia. 4 males, 3 ovig. females, old Providence, in reef, coll. W.L. Schmitt, 06.VIII.1938, USNM 77890. Netherlands Antilles: 1 male, Curaçao, Piscaderabaai, coll. J.G den Hartog, 26.I.1972, RMNH. CRUS. D.30635; 1 ovig. female, Bonaire, Kralendijk, 09.VIII.1994, RMNH. CRUS. D.46177; 1 ovig. female, coll. R.V. Harrison, 2–3 m, II.1975, USNM 155674. Saint Kitts and Nevis. 1 male, 3 (2 ovig.) females, St. Kitts, Windward Beach, on coral reef, colls. W. L. Schmitt & D.V. Nicholson, 12. VI.1956, USNM 136030. Antigua and Barbuda. 2 males, 3 ovig. females, Antigua Island, English Harbor, Charlotte point, 02. VI.1956, USNM 136029. Guadeloupe. 1 male, 1 ovig. female, Guadeloupe, Karubenthos, GB 09, MNHN-IU-2013-12050. Dominica . 1 male, 2 ovig. females, 5 juv, West of Portsmouth, between rocks and dead coral Acropora sp., 1 m, 19. VI.1959, USNM 136035. Saint Lucia. 1 male, 1 ovig. female, the outside Pigeon Island Reef, in coral, 16. VI.1959, USNM 136034. Barbados. 1 male, off the Castle E, side Barbados, coll. C.C. Nutting, 1–4 m, USNM 68724. Trinidad and Tobago. 5 (3 ovig.) females, Tobago Island, Buccoo Reef, coll. W.L. Schmitt, 05. VI.1959, USNM 136033. Brazil. 1 male, 1 ovig. female, Pernambuco, Recife, 8°13’52.1”S 34°37’41.2”W, REC II, dredge 8, coral rubble, colls. A.M.S. Cunha, C.A.L. Santos Júnior, G.O. Soledade, F.D. Silva, R. Guéron & P.S. Santos, 51.8 m, 27. II.2018, MOUFPE 19427; 1 ovig. female, 8°13’33.0”S 34°37’40.3”W, dredge 5, 50.6 m, same collection data, MOUFPE 19428; 1 ovig. female, 8°13’52.1”S 34°37’42.7”W, dredge 10, 50 m, same collection data, MOUFPE 19429; 1 male, 8°13’52.1”S 34°37’42.7”W, dredge 10, in dead coral, 50 m, same collection data, MOUFPE 19430; 1 male, 8°13’52.1”S 34°37’41.2”W, dredge 8, 51.8 m, same collection data, MOUFPE 19431; 1 male, 1 ovig. female, Bahia, Maraú, Taipús de Fora Beach, colls. P.S. Santos, G.O. Soledade & A.O. Almeida, in dead coral, salinity 38, 30.VII.2011, MOUFPE 19432; 2 males, 3 (2 ovig.) females, Bahia, Maraú, Algodões Beach, colls. P.S. Santos, G.O. Soledade & A.O. Almeida, salinity 38, 01.VIII.2011, MOUFPE 19433; 2 males, 4 (3 ovig.) females, Rio de Janeiro, Campos Basin, Petrobrás, Biofouling on Platform P-55 (Old SS-06), coll. J.B. Mendonça, 16 m, 30.III.2010, MZUSP 25346; 1 male, 1 ovig. female, Rio de Janeiro, Búzios, Tartaruga Beach, MNRJ 3478; 1 male, Trindade Island, between Crista do Galo and Ponta Norte, 20°29’14.8”S 29°20’13.9” W, scuba diving, coll. J.B. Mendonça, 15 m, 21.V.2014, MZUSP 33263; 1 male, Trindade Island, Enseada dos Portugueses, Ponta da Calheta, 20°30’18.7”S 29°18’31.6”W, scuba diving, coll. J.B. Mendonça, 16.3 m, 21.V.2014, MZUSP 33295; 1 male, Trindade Island, Enseada dos Portugueses, SECON / ECIT, 20º30’20.9”S 29º18’43.7”W, scuba diving, 9 m, 21.V.2014, MZUSP 33246; 3 males, 2 ovig. females, Trindade Island, Enseada da Cachoeira, Farrilhões, 20°31’22.4”S 29°19’52.0”W, scuba diving, 14.4 m, 5.V.2014, MZUSP 33336. Diagnosis. Rostrum dorsally carinate. Orbital hoods toothed. Ventrolateral tooth on basicerite overreaching stylocerite. Third and fourth pereiopods with spiniform setae on ventrolateral surface of ischium and one tooth on ventrodistal margin of merus. Redescription. (Based on an ovigerous female, Recife, Pernambuco, MOUFPE 19428 and compared with the examined type material). Carapace smooth; rostrum triangular, carinate, tip acute, reaching half-length of first article of antennular peduncle (Fig. 1A). Ocular hoods dorsally inflated, with strong acute tooth arising from dorsomesial margin, margin between rostrum and orbital hoods sinuous. Eyes well-pigmented, completely covered by ocular hoods in dorsal and lateral views (Fig. 1A, B). Cardiac notch well-developed. Pleonal somites smooth; ventral and posterior margins of pleurae 1–4 broadly rounded and pleura of fifth segment subtriangular; sixth pleura without articulated plate. Telson (Fig. 1C) tapering distally, proximal region wider than distal; dorsal surface with two pairs of strong spiniform setae inserted at some distance from lateral margin, first pair anterior and second pair posterior to telson mid-length; distal margin of telson slightly convex, with two pairs of posterolateral spiniform setae, lateral about 2.5 times as long as mesial; anal tubercles well-developed. Uropods (Fig. 1C) with bifid protopods, each lobe ending in acute tooth; exopod longer than endopod; lateral margin of exopod almost straight; diaeresis sinuous, with one adjacent and internal tooth, separated by stout distolateral spiniform setae, overreaching distal margin of exopod; three very small spiniform setae before distal margin; endopod with row of small spiniform setae along distolateral margin. Antennular peduncle moderately slender. Stylocerite distally acute, not reaching distal margin of first article of antennular peduncle; ventromesial carina subtriangular, large, with small tooth in ventrodistal margin (Fig. 1D); second article about 1.2 times longer than dorsally visible portion of first article, about 2.2 times length of third article; third article about 1.5 times longer than wide (Fig. 1A). Antenna basicerite broad, distolateral tooth directed forward; carpocerite slightly exceeding both scaphocerite and distal portion of third antennular article; scaphocerite with shallowly concave lateral margin and large, acute distolateral tooth reaching far beyond distal margin of narrow blade (Fig. 1A). Mandible (Fig. 1E) with 2-jointed palp; incisor process robust, with 10 distinct teeth, mesial tooth conspicuously wider and longer than others; molar process stout, with semicircular rows of short setae. First maxilla (Fig. 1F) with palp bilobed, inferior lobe bearing 1 slender seta, superior lobe not setose. Second maxilla (Fig. 1G) palp well-developed, slender; with upper lacinia cleft, lower lacinia with single lobe; scaphognathite larger than both laciniae, extremities rounded and lateral margin sinuous. First maxilliped (Fig. 1H) palp approximately 2.2 times longer than wide; two laciniae separated by distinct cleft, exopod well-developed with long flagellum and distinct exopodal lobe; epipod about 2.4 times longer than wide. Second maxilliped (Fig. 1I) exopod well-developed, flagellum 2.6 times longer than epipod; epipod about 2.1 times as long as wide. Third maxilliped (Fig. 1J) slender, when extended forward, slightly overreaching distal margin of third article of antennular peduncle; antepenultimate article not flattened or expanded, 1.6 times longer than ultimate article; penultimate article shortest of all, about 2.2 times as long as wide, lateral margin smooth, with tufts of setae, distomesial margin with two thick setae; ultimate article tapering distally, smooth, with several rows of setae, 1.4 times longer than penultimate article, very setose; exopod overreaching distal margin of antepenultimate article, ending in long flagellum; artrobranch well-developed. Epipod present on third maxilliped and from pereiopods 1–4. Major cheliped (Fig. 2A, B) palm strongly sculptured, twisted. Palm about 2.3 times as long as wide, ventral notch broad and shallow, dorsal notch deep and narrow; linea impressa well-marked; lateral surface with low crest starting at about 0.6 times length of palm and ending in sharp distodorsal tooth; ventral margin with rounded, smooth, non-projecting shoulder adjacent to shallow notch, notch continuing transversely into shallow groove on mesial surface; distomesial margin with subcylindrical elevation ending in large adhesive disk distally; distomesial surface with transversally deeply notched crest ending in sharp tooth; dactylus twisted laterally, opening and closing in nearly horizontal plane; strongly arched; compressed proximally; slightly bulbous distally; plunger short (Fig. 2I). Pollex shorter than dactylus; ischium short, stout and unarmed; merus about three times longer than carpus, with distal margin unarmed, ventromesial surface with three small spiniform setae; carpus short and unarmed, cup-shaped. Minor cheliped (Fig. 2C, D) chela slightly compressed, with palm sculptured distally; lateral surface with short crest starting at distal third of palm and ending in sharp distodorsal tooth; ventral margin with blunt, non-protruding shoulder, distomesial margin with subcylindrical elevation ending distally in small adhesive disk; distomesial surface with crest ending in short sharp tooth; fingers as long as palm; pollex shallowly excavated on cutting edge; dactylus somewhat flattened and twisted laterally, slightly convex dorsally, proximally with small adhesive disk; ischium short, stout and unarmed; merus about 2.5 times longer than carpus, with distal margin unarmed, ventromesial surface unarmed; carpus short and unarmed, cup-shaped. Second pereiopod slender (Fig. 2E). Merus slightly longer than ischium; carpus five-segmented, segment ratio (proximal to distal) 3: 2.5: 1: 1: 1.5; distalmost carpal article shorter than chela; chela simple; fingers slightly longer than palm. Third (Fig. 2F) and fourth (Fig. 2G) pereiopods similar in shape and length; both with ischium armed with spiniform seta on ventrolateral surface; merus with ventrodistal tooth; carpus with pair of ventrodistal spiniform setae; propodus with six pairs of strong spiniform setae along ventral margin; dactylus biunguiculate, tooth on flexor margin shorter than extensor margin. Fifth pereiopod (Fig. 2H) shorter than P3 and P4; ischium, merus and carpus unarmed; merus slightly longer than carpus; propodus with five single spiniform setae and one distal pair on ventral margin; dactylus biunguiculate, tooth on flexor margin shorter than extensor margin. Morphological variation. There was no solid morphological variation in the examined characters, considering all specimens and localities. However, the following variations were detected: ventromesial carina of the first article of antennular peduncle was unarmed or armed with a tooth that ranged from small to well-developed. The inner margin of the teeth on ocular hoods ranged from almost straight to slightly convex. The distolateral tooth of the scaphocerite may be either shorter, subequal, or slightly overreaching the distal margin of carpocerite and third article of antennular peduncle. The number of small spiniform setae near the distal margin of the uropodal exopod varied from one to nine (most frequently five). The number of thickened setae on the penultimate article of the third maxilliped varied from one to three (frequently two). The fingers of the minor chela are slightly longer or shorter than the palm. The number of spiniform setae on the ventromesial margin of the merus of the major chela varied from none to six (most frequently four); these setae seem to be easily lost. The form of the molar tooth (Fig. 2I) may be either subtriangular or more subquadrate. The number of pairs of ventral spiniform setae on pereiopods 3 and 4 varied from five to eight (most frequently six pairs in both appendages). The number of single spiniform setae on the ventral surface of pereiopod 5 ranged from three to eight (most frequently five). No sexual dimorphism was observed. Distribution. Western Atlantic: Bermuda; United States of America: Florida, Texas; Mexico: Yucatan, Isla of Cozumel, Bahía de la Ascension, Bahía del Espíritu Santo; Belize; Cuba; Costa Rica; Panama; Colombia: Providence Island; Curaçao; Bonaire; Porto Rico; Anguilla; Saint Kitts and Nevis; Barbuda; Antigua; Guadeloupe; Dominica; Saint Lucia; Barbados; Tobago Cays; Trinidad and Tobago; Venezuela; Brazil: Ceará, Pernambuco, Bahia, Rio de Janeiro, Trindade and Martin Vaz Archipelago (Armstrong 1940; Chace 1972; Rodríguez 1980; Santos et al. 2012; Cunha et al. 2015; Anker et al. 2016; Pachelle et al. 2016; this study) (Fig. 3). Ecology. Found in intertidal areas up to 51.8 m depth (Martínez-Iglesias et al. 1997; Anker et al. 2016; this study). Inhabits polychaete tubes, coral rubble, dead corals, and calcareous algae, and it is found under rocks or in rock crevices, among mangrove roots near the coast (Carvacho 1979; Santos et al. 2012), and occasionally in sponges of the genus Callyspongia Duchassaing & Michelotti, 1864 (Chace 1956). Colour in life. Body color ranges from greenish-yellow to reddish (Rodriguez 1986; Sterrer 1986; Anker et al. 2016). According to Anker et al. (2016), the large white spots with brown chromatophores on the major chela are an accurate diagnostic character. Fresh specimens examined by us were predominantly greenish-yellow. However, we also observed specimens with body and appendages transparent with reddish chromatophores. Eggs in initial stage are olive green., Published as part of Santos, Patricia S., Terossi, Mariana, Mantelatto, Fernando L. & Almeida, Alexandre O., 2019, Using integrative taxonomy to establish the status of Alpheus peasei (Armstrong, 1940) (Decapoda: Alpheidae) as a single species throughout its distribution, pp. 51-68 in Zootaxa 4629 (1) on pages 55-59, DOI: 10.11646/zootaxa.4629.1.4, http://zenodo.org/record/3268287, {"references":["Armstrong, J. C. (1940) New species of Caridea from the Bermudas. American Museum Novitates, 1096, 1 - 10.","Chace, F. A. Jr. (1956) Crustaceos Decapodos y Stomatopodos del Archipelago de los Roques e Isla de la Orchila. In: Mendez, A. (Ed.), Archipelago de Los Roques y la Orchila. Sociedad de Ciencias Naturales La Salle, Editorial Sucre, Caracas, pp. 145 - 168, pl. 4.","Chace, F. A. Jr. (1972) The shrimps of the Smithsonian-Bredin Caribbean expeditions with a summary of the West Indian shallow water species (Crustacea: Decapoda: Natantia). Smithsonian Contributions to Zoology, 98, 1 - 179. https: // doi. org / 10.5479 / si. 00810282.98","Hazlett, B. A. & Winn, H. E. (1962) Sound production and associated behavior of Bermuda crustaceans (Panulirus, Gonodactylus, Alpheus, and Synalpheus). Crustaceana, 4, 25 - 38. https: // doi. org / 10.1163 / 156854062 X 00030","Knowlton, R. E. & Moulton, J. M. (1963) Sound production in the snapping shrimps Alpheus (Crangon) and Synalpheus. The Biological Bulletin, 125, 311 - 331. https: // doi. org / 10.2307 / 1539406","Pequegnat, L. H. & Ray, J. P. (1974) Crustacea and other arthropods. In: Bright, T. J. & Pequegnat, L. H. (Eds.), Biota of the Fest Flower Garden Bank. Gulf Publishers Co., Houston, Texas, pp. 232 - 261.","Ray, J. P. (1974) A study of the coral reef crustaceans (Decapoda and Stomatopoda) of two Gulf of Mexico reef systems: West Flower Garden, Texas and Isla de lobos, Veracruz, Mexico. PhD Dissertation, Texas A & M University, College Station, Texas, 323 pp.","Abele, L. G. (1976) Comparative species composition and relative abundance of decapod crustaceans in marine habitats of Panama. Marine Biology, 38, 263 - 275. https: // doi. org / 10.1007 / BF 00388939","Carvacho, A. (1979) Les crevettes Carides de la mangrove guadeloupeenne. Bulletin du Museum National d'Histoire Naturelle, Serie 4, 1, Section A, 2, 445 - 470.","Abele, L. G. & Kim, W. (1986) An illustrated guide to the marine decapod crustaceans of Florida. State of Florida, Department of Environmental Regulation, 8, 1 - 196.","Rodriguez, B. (1986) Los camarones (Crustacea; Decapoda; Natantia) del Parque Nacional Archipielago de los Roques. PhD dissertation, Universidad Central de Venezuela, Caracas, 350 pp.","Sterrer, W. (1986) Marine fauna and flora of Bermuda: a systematic guide to the identification of marine organisms. John Wiley & Sons, 742 pp.","Martinez-Iglesias, J. C., Gomez, O., Carvacho, A. & Rios, R. (1993) Nuevos registros de crustaceos decapodos (Crustacea: Decapoda) en la plataforma marina de Cuba. Avicennia, 9 - 13.","Martinez-Iglesias, J. C., Carvacho, A. & Rios, R. (1996) Catalogo de los carideos marinos (Crustacea, Decapoda, Caridea) de las aguas someras de Cuba. Avicennia, 4, 27 - 40.","Martinez-Iglesias, J. C., Rios, R. & Carvacho, A. (1997) Las especies del genero Alpheus (Decapoda: Alpheidae) de Cuba. Revista de Biologia Tropical, 44, 401 - 429.","Santos, P. S., Soledade, G. O. & Almeida, A. O. (2012) Decapod crustaceans on dead coral from reef areas on the coast of Bahia, Brazil. Nauplius, 20, 145 - 169. https: // doi. org / 10.1590 / S 0104 - 64972012000200007","Soledade, G. O. & Almeida, A. O. (2013) Snapping shrimps of the genus Alpheus Fabricius, 1798 from Brazil (Caridea: Alpheidae): updated checklist and key for identification. Nauplius, 21, 89 - 122. https: // doi. org / 10.1590 / S 0104 - 64972013000100010","Cunha, A. M., Soledade, G. O., Boos, H. & Almeida, A. O. (2015) Snapping shrimps of the genus Alpheus Fabricius, 1798 (Caridea: Alpheidae) from Brazil: range extensions and filling distribution gaps. Nauplius, 23, 47 - 52. https: // doi. org / 10.1590 / S 0104 - 64972015002305","Anker, A., Tavares, M. & Mendonca Jr., J. B. (2016) Alpheid shrimps (Decapoda: Caridea) of the Trindade & Martin Vaz Archipelago, off Brazil, with new records, description of a new species of Synalpheus and remarks on zoogeographical patterns in the oceanic islands of the tropical southern Atlantic. Zootaxa, 4138, 1 - 58. https: // doi. org / 10.11646 / zootaxa. 4138.1.1","Pachelle, P. P., Anker, A., Mendes, C. B. & Bezerra, L. E. (2016) Decapod crustaceans from the state of Ceara, northeastern Brazil: an updated checklist of marine and estuarine species, with 23 new records. Zootaxa, 4131 (1), 1 - 63. https: // doi. org / 10.11646 / zootaxa. 4131.1.1","Rodriguez, G. (1980) Crustaceos decapodos de Venezuela. Instituto Venezolano de Investigaciones Cientificas, Caracas, 494 pp."]}

Alpheus amblyonyx Chace, 1972 (Figs. 4–6) Alpheus amblyonyx Chace, 1972: 59; fig. 16. [Near center of Arrecife Nicchehabin, Baía de la Ascensión, Territorio de Quintana Roo, Mexico]; Christoffersen 1979: 302, fig. 1 (in part); Abele & Kim 1986: 197, 208, 209, figs. b–d; Martínez-Iglesias et al. 1997: 403, fig. 2; 356; Coelho Filho 2006: 7; Serejo et al. 2007: 198; Alves et al. 2008: 48; Souza et al. 2011: 46; Soledade & Almeida 2013: 94 (in part); Anker et al. 2016: 14, fig. 2. Type material. 1 OVF, Mexico, Quintana Roo, Bahía de la Ascención, #52–60, USNM 135356 (Not examined). Material examined. WEST ATLANTIC— Mexico. 1 NI, Quintana Roo, southwest of the Chinchorro Reef, 28.VIII.1990, CNCR 21271 (topotype, genetic voucher); 1 NI, Gulf of Mexico, 34–36 m, R/ V Pelican, Box Dredge 14.XI.2005, 022°6.78’N 91°26.43’W, ULLZ 6615 (genetic voucher); Belize. 1 OVF, Stann Creek, Carrie Bow Cay, 16.80251°N 88.081747°W, 3–4 m, 28.IV.2015, ULLZ 16584 (genetic voucher); 1 M, same locality, 04.IV.2007, ULLZ17432; 1 OVF, same locality, 21.IV.2015, ULLZ16509, Guadeloupe. 1 M, KARUBENTHOS, #GB20, 16 m, 18.V.2012, MNHN 12112; 1 OVF, KARUBENTHOS, #GB24, 16 m, 18.v.2012, MNHN 12026 (genetic voucher); Saint Martin. 1 OVF, Chicot, windward side of Tintamarre Island, 12–20 m, 23.IV.2012, UF 32400 (genetic voucher); Brazil. 1 OVF, Espírito Santo, REVIZEE Benthos #34, 20°24’10”S 39°55’35”W, R/ V Astro Garoupa, 50 m, dredge, 15.VII.2001, MNRJ 17471; 1 OVF, Trindade Island, Ilha da Racha, 20°30’26.5”S 29°20’40.0”W, 32 m, under rocks, 30.I.2012, MZUSP 30317; 1 OVF, Trindade Island, Ilha da Racha, 20°30’26.5”S 29°20’40.0”W, 25 m, under rocks, 12.XI.2014, MZUSP 30262; 1 OVF, off Rio de Janeiro, REVIZEE Benthos #Y1, R/ V Astro Garoupa, 60 m, dredge, 11. VI.2002, MNRJ 17467. Type locality. WEST ATLANTIC—Mexico, Quintana Roo, Bahía de la Ascensión (Chace 1972). Distribution. WESTERN ATLANTIC–Gulf of Mexico, Yucatan Peninsula, Florida, Caribbean Sea, Brazil (Ceará to Espírito Santo, North Chain Seamounts, Fernando de Noronha, Atol das Rocas, Vitória-Trindade Seamount Chain, Trindade & Martin Vaz Archipelago) (Chace 1972; Abele & Kim 1986; Christoffersen 1979; Martínez-Iglesias et al. 1997; Soledade & Almeida 2013 and references therein; Anker et al. 2016; present study). Diagnosis. Rostrum almost reaching distal margin of first article of antennular peduncle; scaphocerite blade reaching half length of second article of antennular peduncle; distal tooth reaching distal limit of antennular peduncle. Minor cheliped with ischium armed with 4 spiniform setae; ventromesial margin of merus with 6 spiniform setae similar in size and shape, spaced almost equidistantly along length of mesial surface of merus; dactylus without crest on extensor margin. Major cheliped with ischium armed with 3 spiniform setae; ventromesial margin of merus with 6 spiniform setae similar in size and shape, spaced almost equidistantly along length of mesial surface of merus; dactylus flattened, laterally twisted, dorsally convex and strongly bulbous distally; plunger developed, with anterior surface distinctly angular. Second pereiopod elongated, slender; ischium slightly longer than merus; carpus subdivided in 5 articles with ratio approximately equal to 3: 2: 1: 1.3: 1.5. Distolateral margin of uropodal exopod with 1 mesial and 1 lateral teeth adjacent to spiniform setae. Morphological characterization (bold characters represent the main differences regarding A. macrocheles). Carapace smooth, frontal margin with strong rostrum, ocular hoods with well-developed teeth originating from anterior margin of hoods; rostrum slightly flat in dorsal view, distinctly longer than broad, distally tapering and ending in sharp point, without setae, almost reaching distal margin of first article of antennular peduncle; margin between orbital teeth and rostrum V-shaped (Fig. 4A, B); orbital process present. Pterygostomial angle rounded, not anteriorly protruding (Fig. 4B); cardiac notch deep. Pleonal segments with posteroventral margins broadly rounded, those of fifth segment slightly angular; sixth segment without articulated plate in posterolateral margin. Eyes with well-developed corneas; ocellar beak projecting anteriorly, acute, visible in lateral view. Antennular peduncle moderately slender; stylocerite with acute tip reaching distal limit of first article of antennular peduncle; ventromesial carina well-developed, triangular in shape, with small pointed tooth (Fig. 4C); second article of antennular peduncle much longer than visible part of first article, about 2 times longer than broad (Fig. 4A); lateral flagellum with groups of aesthetascs extending to article 10. Antennal basicerite with acute lateral tooth; carpocerite slightly exceeding distal end of scaphocerite and antennular peduncle; lateral margin of scaphocerite slightly concave, ending in strong lateral tooth overpassing distal margin of blade (Fig. 4A); blade reaching half length of second article of antennular peduncle; distal tooth reaching distal limit of antennular peduncle. Mouthparts as illustrated by Chace (1972). Third maxilliped slender; coxa with lateral plate slightly truncated distally; exopod reaching beyond distal limit of antepenultimate article when extended; antepenultimate article somewhat flattened, approximately 4 times longer than broad, ventral surface sparsely setose; penultimate article approximately 3 times longer than broad, slightly broadened distally and densely setose; ultimate article unarmed, distally tapering, with dense rows of long setae (Fig. 4I). Minor cheliped with short and stout ischium; distomesial margin broad, sub-triangular in cross-section, armed with 4 spiniform setae; ventrolateral surface unarmed; ventromesial margin of merus slightly convex, ending in small acute tooth, with 6 spiniform setae similar in size and shape, spaced almost equidistantly along length of mesial surface of merus; carpus cup-shaped; chelae strongly compressed, palm with grooves and notches from distal half of lateral surface; mesial surface convex and devoid of depressions or grooves; lateral surface with low crest starting at about middle of palm and ending in acute distolateral tooth; ventral surface smooth, ending in welldefined notch; dorsal margin also smooth, with sub-cylindrical elevation ending distally in small adhesive disk; distomesial surface with high ridge ending in strong sharp tooth; fingers as long as palm; pollex lightly excavated on cutting edge; dactylus slightly flattened and twisted laterally, without crest on extensor margin, with small adhesive disk proximally (Fig. 5C, D). Major cheliped with ischium short and robust, ventromesial surface with 3 small spiniform setae; merus robust, not elongated, about 2 times longer than wide, sub-triangular in cross section; ventrolateral margin unarmed; ventromesial margin straight, with 6 small spiniform setae, ending in robust and sharp tooth; carpus very short, cupshaped, slightly compressed; lateral surface with low crest starting at approximately 0.6 of length of palm, ending in distolateral sharp tooth, slightly directed laterally; linea impressa well-marked; mesial surface convex, devoid of sculptures and grooves; ventral surface with rounded smooth shoulder slightly projecting into adjacent deep notch; notch extending transversely to groove on mesial surface; dorsal margin with sub-cylindrical elevation ending distally in large adhesive disk; distomesial surface with deep transversally notched crest ending in sharp tooth; distal third of ventral margin of palm with strong and long setae extending to distomesial margin of palm; pollex shorter than dactylus, strongly curved laterally, cutting edge slightly excavated and with small rounded tooth in proximal portion; dactylus flattened, laterally twisted, dorsally convex and strongly bulbous distally; plunger developed, with anterior surface distinctly angular (Fig. 5A, B). Second pereiopod elongated, slender; ischium slightly longer than merus; carpus with 5 articles with ratio approximately equal to 3: 2: 1: 1.3: 1.5; chelae not sculptured, fingers as long as palm, with small tufts of setae (Fig. 6A). Third pereiopod with ischium armed with strong spiniform setae on ventrolateral surface; merus 4 times longer than broad, without tooth at distoventral angle; carpus slender, about half length of merus; propodus about 1.5 times longer than carpus, with about 9 robust spiniform setae along ventral margin and 1 pair of spiniform setae near to articulation with dactylus; dactylus slightly shorter than half length of propodus, conical, slightly curved, ending in sharp tip; extensor margin with 1 sub-terminal denticle (Fig. 6B). Fourth pereiopod similar to third in shape and proportion of articles, but slightly less robust; ischium armed with strong spiniform setae on ventrolateral surface; propodus with about 7 robust spiniform setae along ventral margin and 1 pair of spiniform setae near to articulation with dactylus (Fig. 6C). Fifth pereiopod slenderer than third and fourth (Fig. 6D); merus slightly longer than carpus; ischium with 1 spiniform setae; propodus with about 8 spiniform setae along ventral margin and 10 well-developed distolateral setal brushes. Pleopods with sparse setae on lateral margin of protopod; endopods and exopods with edges densely setose; first pleopod very reduced, distal margin of endopod with setae; second pleopod in male with appendix masculina subequal in length to appendix interna, not reaching distal margin of endopod, with numerous rigid setae at distal end; uropod with lateral lobe of protopod ending in strong and sharp tooth; diaeresis sinuous, ending in 1 mesial and 1 lateral teeth adjacent to spiniform setae; endopod with row of small spiniform setae along distal margin (Fig. 4I). Telson subretangular, tapering to posterior margin; proximal margin 2.0 times broader than distal margin; lateral margins slightly convex; dorsal surface with 2 pairs of spiniform setae, first pair located at half-length of telson, second pair located approximately 0.7 of length of telson; posterior margin broadly convex, with 2 pairs of spiniform setae, mesial pair about 3 times size of lateral pair (Fig. 4J); anal tubercles well-developed. Morphological variation. Little morphological variation has been observed (see Table 2). No significant differences between sexes have been observed. Comparative material analyzed. Alpheus crockeri. EAST ATLANTIC— Cape Verde. 1 NI, São Vicente Island, 01.X.2013, OUMNH 2014-01 - 0030 (genetic voucher). Alpheus pouang. WEST ATLANTIC— Brazil. 1 M (Holotype), off São Paulo, R/ V Wladimir Besnard #1019, 23°39’S 43°37’W, 120–121 m, mud and sand, 27.V.1970, MZUSP 4543; 1 F, 1 M, off Rio Grande do Sul, GEDIP I #330, 34°35’S 52°00’W, sand, 30. VI.1968, MNRJ 4388. Alpheus puapeba: WEST ATLANTIC— Brazil. 1 OVF (Paratype), off Rio de Janeiro, R/ V Wladimir Besnard #1632, 23°33’S 44°11’W, 72 m, 17.XII.1971, MZUSP 31525; 1 OVF, off Rio de Janeiro, HAB 2 #66, 23°11’15.741”S 41°37’17.184”W, trawl, continental shelf of the Campos Basin, 26.IV.2008, MNRJ 21836; 1 M, Rio de Janeiro, Ilha Grande, #302, 49 m, 16.II.1968, MNRJ 4387; 1 F, 1 M, HAB 1, # 66, trawl, 98.7 m, 26.IV.2008, MNRJ 21836 (genetic voucher); 4 F, 10 M, off Santa Catarina, #61 — CB102, 27°07’S 42°03’W, 100 m, 01. VI.1987, MZUSP 31900. Alpheus platydactylus: WEST ATLANTIC— Azores. 1 OVF, 1 NI, # 5.010, 26.V.1981, RMNH. Crus. D.51229; 2 OVF, # 5.129, 06.VI.1981, RMNH. Crus. D.51237; Canary Islands. 1 OVF, 1 M, # 4.07, 20.I.1980, RMNH. Crus. D.51238; 2 F (1 OV), # 4.17, 09.I.1980, RMNH. Crus. D.51239; Cape Verde. 2 F (1 OV), 1 M, “Tydeman” Cancap- VII, # 7.122, 01.IX.1986, RMNH. Crus. D.51240. Molecular analysis. A total of 10 new 16S rRNA gene sequences were obtained from species of the A. macrocheles species complex (see Table 1). The substitution model that best fit the data was the TPM3uf + I (= K81) assuming the nucleotide frequencies A = 0.2812, C = 0.1185, G = 0.2601, T = 0.3402, replacement rates AC = 0.1431, AG = 4.6735, AT = 1, CG = 0.1431, CT = 4.6735, GT = 1, and invariable sites = 0.54. The topology generated from the Bayesian Inference Analysis shows the presence of two main clades, clearly separating the specimens of A. macrocheles and A. amblyonyx (Fig. 8). In the first clade A. crockeri (Cape Verde) is placed as a sister group of A. puapeba (Brazil) + A. macrocheles (Azores and Canary Islands), in the second clade are the specimens of A. amblyonyx, including the topotypic material (Quintana Roo, Mexico). In both clades the groupings are well supported by the posterior probability values (Fig. 8). The values of interspecific genetic distance (K81 model) ranged from 14 (A. macrocheles X A. puapeba) to 29% (A. amblyonyx X A. crockeri). There was no genetic intraspecific divergence among specimens of A. macrocheles or A. amblyonyx. The two species of interest (A. macrocheles and A. amblyonyx) had a genetic distance of 19%., Published as part of Soledade, Guidomar O., Scioli, Justin A., Terossi, Mariana, Mantelatto, Fernando L. & Almeida, Alexandre O., 2019, Congruence between morphology and molecular data supports the distinction between the snapping shrimps Alpheus macrocheles and A. amblyonyx (Caridea, Alpheidae), pp. 1-22 in Zootaxa 4624 (1) on pages 9-15, DOI: 10.11646/zootaxa.4624.1.1, http://zenodo.org/record/3258424, {"references":["Chace Jr., F. A. (1972) The shrimps of the Smithsonian-Bredin Caribbean Expeditions with a summary of the West Indian shallow-water species (Crustacea: Decapoda: Natantia). Smithsonian Contributions to Zoology, 98, 1 - 179. https: // doi. org / 10.5479 / si. 00810282.98","Christoffersen, M. L. (1979) Decapod Crustacea: Alpheoida. Campagne de la Calypso au large des cotes atlantiques de l'Amerique du Sud (1961 - 1962). I. 36, Annales de l'Institute Oceanographique, Monaco, 55 (Supplement), 297 - 377.","Abele, L. G. & Kim, W. (1986) An illustrated guide to the marine decapod crustaceans of Florida. Technical Series, Department of Environmental Regulations. Vol. 8. No. 1 & 2. Department of Biological Sciences, Florida State University, Tallahassee, Florida, 760 pp.","Martinez-Iglesias, J. C., Rios, R. & Carvacho, A. (1997) Las especies del genero Alpheus (Decapoda: Alpheidae) de Cuba. Revista de Biologia Tropical, 44, 401 - 429.","Coelho Filho, P. A. (2006) Checklist of the Decapods (Crustacea) from the outer continental shelf and seamounts from Northeast of Brazil - REVIZEE Program (NE III). Zootaxa, 1184 (1), 1 - 27. https: // doi. org / 10.11646 / zootaxa. 1184.1.1","Serejo, C. S., Cardoso, I. A., Tavares, C. R., Abreu Jr., C. R., Amancio, I. C. & Senna, A. R. (2007) Filo Arthropoda: Subfilo Crustacea. In: Lavrado, H. & Viana, M. S. (Eds.), Atlas de invertebrados marinhos da regiao central da Zona Economica Exclusiva Brasileira. Parte 1. Serie Livros 25. Museu Nacional, Rio de Janeiro, pp. 165 - 214.","Alves, M. L., Ramos-Porto, M. & Viana, G. F. S. (2008) Checklist of the decapods (Crustacea) from the Fernando de Noronha Archipelago, Brazil. Zootaxa, 1881, 43 - 68. https: // doi. org / 10.11646 / zootaxa. 1881.1","Souza, J. A. F., Schwamborn, R., Barreto, A. V., Farias, I. D., Fernandes, L. M. G. & Coelho, P. A. (2011) Marine and estuarine shrimps (Dendrobranchiata, Stenopodidea, and Caridea), of Pernambuco state (Brazil) and northeastern Brazilian oceanic islands. Atlantica, 33, 33 - 63. https: // doi. org / 10.5088 / atl. 2011.33.1.33","Soledade, G. O. & Almeida A. O. (2013) Snapping shrimps of the genus Alpheus Fabricius, 1798 from Brazil (Caridea: Alpheidae): updated checklist and key for identification. Nauplius, 21 (1), 89 - 122. https: // doi. org / 10.1590 / S 0104 - 64972013000100010","Anker, A., Tavares, M. & Mendonca, J. B. (2016) Alpheid shrimps (Decapoda: Caridea) of the Trindade & Martin Vaz Archipelago, off Brazil, with new records, description of a new species of Synalpheus and remarks on zoogeographical patterns in the oceanic islands of the tropical southern Atlantic. Zootaxa, 4138 (1), 1 - 58. https: // doi. org / 10.11646 / zootaxa. 4138.1.1"]}

Alpheus alpheopsides Couti��re, 1905 (Fig. 1B) Alpheus alpheopsides Couti��re, 1905: 901; Anker & De Grave, 2016: 344. Material examined. PG: 1 male (ZUTC 6682), Bushehr, Coll. R. Naderloo, 3 Apr. 2015; 1 male and 1 female (ZUTC 6683), Hormuz I., on dead corals, Coll. A. Dehghani, 15 May 2016; GO: 2 males, 2 ovig. females (ZUTC 6684) Chabahar Bay, Tis, on dead corals, Coll. A. Sari and A. Kazemi, 2 May 2014. Distribution. Indo-West Pacific: from Red Sea to Micronesia and Samoa. Regional records. PG and GO: Iran (present study). Habitat. Found under coral rubble or in crevices of dead coral heads. Remark. This species belongs to the Alpheus paracrinitus Miers, 1881 species complex, but shows a distinct distribution pattern compared to A. paracrinitus. This species is distinguishable from A. paracrinitus by absence of balaeniceps on minor chela. Geographically, A. paracrinitus is an amphi-Atlantic species while A. alpheopsides occurs in the Indo-West Pacific., Published as part of Dehghani, Amir, Sari, Alireza & Naderloo, Reza, 2019, Annotated checklist of the snapping shrimps of the genus Alpheus Fabricius, 1798 (Crustacea: Decapoda: Alpheidae) from the Persian Gulf and Gulf of Oman, Iran, pp. 479-504 in Zootaxa 4544 (4) on pages 481-482, DOI: 10.11646/zootaxa.4544.4.2, http://zenodo.org/record/2618475, {"references":["Coutiere, H. (1905) Marine Crustaceans. Les Alpheidae. XV. (Plates LXX-LXXXVII et Texte-Figures 127 - 139). In: Gardiner, J. S. (Ed.), The Fauna and Geography of the Maldive and Laccadive Archipelagoes. 2 (4). University Press, Cambridge and London, pp. 852 - 921.","Anker, A. & De Grave, S. (2016) An updated and annotated checklist of marine and brackish caridean shrimps of Singapore (Crustacea, Decapoda). Raffles Bulletin of Zoology, 34, 343 - 454.","Miers, E. J. (1881) On a Collection of Crustacea made by Baron Hermann Maltzam at Goree Island, Senegambia. The Annals and Magazine of Natural History, 5, 8 (45), 204 - 220 + 259 - 281 + 364 - 377."]}

Alpheus gracilis Heller, 1861 (Fig. 3 C���D) Alpheus gracilis Heller, 1861: 271; De Man, 1897: 733; Tiwari, 1963: 283. Alpheus gracilis alluaudi ��� Coutiere, 1905a: 882. Alpheus gracilis luciparensis ��� De Man, 1911: 337. Alpheus gracilis gracilis ���Banner & Banner, 1968: 280. Crangon gracilis simplex ��� Banner, 1953: 75. Alpheus gracilis simplex ��� Banner & Banner, 1966b: 97. Material examined. PG : 3 males, 3 ovig. females (ZUTC 6674), Abu-Musa I., sandy/boulder shores, Coll. A. Dehghani & H. Ashrafi, 10 Oct. 2017; 1 male, 1 female (SMF 51380), Abu-Musa I., sandy/boulder shores, Coll. A. Dehghani & H. Ashrafi, 10 Oct. 2017; GO: 2 males, 2 ovig. females (ZUTC 6673), Chabahar Bay, Tis, in large rocks and coral rubble, Coll. A. Dehghani, 2 Oct. 2016. Distribution. Indo-West Pacific: South Africa, Red Sea, Australia, central Pacific to Hawaii, and Society Islands. Regional records. PG and GO: Iran (present study). Habitat. All specimens were collected from crevices in dead coral heads and mostly found in water from 3���6 m deep. This species is syntopic with Alpheus parvirostris. Remark: Heller (1861) described Alpheus gracilis from the Red Sea. Couti��re (1905) described a subspecies Alpheus gracilis alluaudi based on material collected in the Maldives. According to Couti��re (1905), the subspecies shows no secondary unguis on the dactylus of the third pereiopod. De Man (1911) described a variety of the species from the Lucipara Island as A. gracilis luciparensis. Banner & Banner (1983) studied various collections from Red Sea, Thailand, southern Philippines and Christmas Island, Maui and other Hawaiian Islands and listed three morphological variations for the species. These variations are categorized as: 1) Presence of deep or shallow orbito-rostral grooves, 2) Presence or absence of a projection on dactylar articulation of small chela, 3) Simple or bi-ungulate dactylus of third pereiopod. Based on these features, Banner & Banner (1983) concluded that this species shows a tendency to develop morphologically separated geographic lineages. All of our examined specimens show shallow orbito-rostral grooves, a projection on the dactylar articulation of small chela, and a biungulate dactylus of the third pereiopod., Published as part of Dehghani, Amir, Sari, Alireza & Naderloo, Reza, 2019, Annotated checklist of the snapping shrimps of the genus Alpheus Fabricius, 1798 (Crustacea: Decapoda: Alpheidae) from the Persian Gulf and Gulf of Oman, Iran, pp. 479-504 in Zootaxa 4544 (4) on pages 485-486, DOI: 10.11646/zootaxa.4544.4.2, http://zenodo.org/record/2618475, {"references":["Heller, C. (1861) Synopsis der im rothen Meere vorkommenden Crustaceen. Ferhandlungen der kaiserlich-koniglichen zoologisch-botanischen Gesellschaft in Wien, 11, 3 - 31.","De Man, J. G. (1897) Bericht uber die von Herrn Schiffscapitan Storm zu Atjeh, an den westlichen Kusten von Malakka, Borneo und Celebes sowie in der Java-See gesammelten Decapoden und Stomatopoden. Funfter Theil. Zoologische Jahrbucher. Abtheilung fur Systematik, Geographie und Biologie der Thiere, 9, 725 - 790. https: // doi. org / 10.5962 / bhl. title. 16084","Tiwari, K. K. (1963) Alpheid shrimps (Crustacea: Decapoda: Alpheidae) of Vietnam. Annales de la Faculte des Sciences de Saigon, 1963, 269 - 362.","Coutiere, H. (1905) Marine Crustaceans. Les Alpheidae. XV. (Plates LXX-LXXXVII et Texte-Figures 127 - 139). In: Gardiner, J. S. (Ed.), The Fauna and Geography of the Maldive and Laccadive Archipelagoes. 2 (4). University Press, Cambridge and London, pp. 852 - 921.","De Man, J. G. (1911) The Decapoda of the Siboga Expedition. Part II. Family Alpheidae. Siboga Expeditie, 39 a (2), 133 - 465. https: // doi. org / 10.5962 / bhl. title. 10512","Banner, A. H. (1953) The Crangonidae, or snapping shrimp, of Hawaii. Pacific Science, 7, 3 - 147.","Banner, A. H. & Banner, D. M. (1966 b) Contributions to the knowledge of the alpheid shrimp of the Pacific Ocean. Part X. Collections from Fiji, Tonga and Samoa. Pacific Science, 20, 145 - 188.","Banner, A. H. & Banner, D. M. (1983) An annotated checklist of the alpheid shrimp from the Western Indian Ocean. Travaux et Documents de l'O. R. S. T. O. M., 158, 1 - 164."]}

Alpheus pachychirus Stimpson, 1860 (Fig. 5F) Alpheus pachychirus Stimpson, 1861: 30; De Man, 1890:116; Ortmann, 1890: 487; Cowles, 1913: 121. Material examined. PG : 1 male (ZUTC 6681), Abu-Musa I., on dead coral, Coll. A. Dehghani, 10 May 2016. Distribution. Indo-West Pacific: from Red Sea, PG, and India to the Philippines. Regional records. PG : Iran (present study). Habitat. In dead coral heads and live sponges; from lower intertidal to over 3 m. Remarks. Alpheus pachychirus seems to be either a morphologically variable species or a species complex (Anker & De Grave, 2016). Therefore, there is a need for special taxonomic treatment using diverse character analyses., Published as part of Dehghani, Amir, Sari, Alireza & Naderloo, Reza, 2019, Annotated checklist of the snapping shrimps of the genus Alpheus Fabricius, 1798 (Crustacea: Decapoda: Alpheidae) from the Persian Gulf and Gulf of Oman, Iran, pp. 479-504 in Zootaxa 4544 (4) on page 489, DOI: 10.11646/zootaxa.4544.4.2, http://zenodo.org/record/2618475, {"references":["Stimpson, W. (1860) Prodromus descriptionis animalium evertebratorum, quae in Expeditione ad Oceanum Pacificum Septentrionalem, a Republica Federata missa, Cadwaladore Ringgold et Johanne Rodgers Ducibus, observavit et descripsit. Pars VIII, Crustacea Macrura. Proceedings of the Academy of Natural Sciences of Philadelphia, 1860, 22 - 47.","Ortmann, A. (1890) Die Decapoden-Krebse des Strassburger Museums, mit besonderer Berucksichtigung der von Herrn Dr. Doderlein bei Japan und bei den Liu-Kiu-Inseln gesammelten und z. Z. im Strassburger Museum aufbewahrten Formen. I. Theil. Die Unterordnung Natantia (Boas) (Abtheilungen: Penaeidae und Eucyphidea Caridae der Autoren). Zoologische Jahrbucher. Abtheilung fur Systematik, Geographie und Biologie der Thiere, 5, 437 - 542.","Anker, A. & De Grave, S. (2016) An updated and annotated checklist of marine and brackish caridean shrimps of Singapore (Crustacea, Decapoda). Raffles Bulletin of Zoology, 34, 343 - 454."]}

Alpheus spongiarum Couti��re, 1897 Alpheus spongiarum Couti��re, 1897c: 236; De Man, 1911: 362; Johnson, 1962: 52; 1963: 283; 1979: 41; Banner & Banner, 1978: 230; 1982: 116; 1985: 31; Chace, 1988: 54; Anker & De Grave, 2016: 369. Material examined. GO: 3 males (ZUTC 6688), Abu-Musa I., in passageways of dead sponges, Coll. A. Dehghani, 15 June 2016; 4 males (ZUTC 6689), Siri I., in passageways of dead sponges, Coll. A. Sari, 14 Nov. 2016; GO: 2 males, 2 ovig. females (ZUTC 6680) Chabahar Bay, Tis, in passageways of recently dead sponges washed to the beach, Coll. A. Dehghani, 2 Oct. 2016. Distribution. Indo-West Pacific: from Red Sea, PG, GO to Singapore, Japan, Papua New Guinea and Australia. Regional records. PG and GO: Iran (present study). Habitat. This species is associated with living sponges and found in the passageways of dead sponges washed to the beach; from lower intertidal to over 3 m deep (Anker & De Grave, 2016). Remark. This species is easily distinguished by a distinct projection on the first abdominal pleura. All material collected from Siri Island shows the following features which agree well with Anker & De Grave���s (2016) material: 1) Ischium of third pereiopod unarmed; 2) First carpal article of second pereiopod about 0.4 times as length of second article; 3) First pleura projected and second to fifth pleura angular; 4) Scaphocerite blade short; 5) Major chela ovoid, not compressed; and 7) Dactylus of minor chela longer than palm (see Banner & Banner, 1982, Anker & De Grave, 2016)., Published as part of Dehghani, Amir, Sari, Alireza & Naderloo, Reza, 2019, Annotated checklist of the snapping shrimps of the genus Alpheus Fabricius, 1798 (Crustacea: Decapoda: Alpheidae) from the Persian Gulf and Gulf of Oman, Iran, pp. 479-504 in Zootaxa 4544 (4) on page 498, DOI: 10.11646/zootaxa.4544.4.2, http://zenodo.org/record/2618475, {"references":["Coutiere, H., (1897 c) Note sure Betaeus Jousseaumei, nouvelle espece d'Alphee de la Mer Rouge. Bulletin du Museum d'Histoire Naturelle, 2, 236 - 237.","De Man, J. G. (1911) The Decapoda of the Siboga Expedition. Part II. Family Alpheidae. Siboga Expeditie, 39 a (2), 133 - 465. https: // doi. org / 10.5962 / bhl. title. 10512","Johnson, D. S. (1962) A synopsis of the Decapoda Caridea and Stenopodidea of Singapore, with notes on their distribution and a key to the genera of Caridea occurring in Malaysian waters. Bulletin of the National Museum Singapore, 30, 44 - 79.","Banner, D. M. & Banner, A. H. (1978) Annotated checklist of alpheid and ogyridid shrimp from the Philippine Archipelago and the South China Sea. Micronesica, 14 (2), 215 - 257.","Chace, F. A. Jr. (1988) The caridean shrimps (Crustacea: Decapoda) of the Albatross Philippine Expedition, 1907 - 1910, Part 5: family Alpheidae. Smithsonian Contributions to Zoology, 466, 1 - 99.","Anker, A. & De Grave, S. (2016) An updated and annotated checklist of marine and brackish caridean shrimps of Singapore (Crustacea, Decapoda). Raffles Bulletin of Zoology, 34, 343 - 454.","Banner, D. M. & Banner, A. H. (1982) The alpheid shrimp of Australia. Part III: The remaining alpheids, principally the genus Alpheus and the family Ogyrididae. Records of the Australian Museum, 34, 1 - 357. https: // doi. org / 10.3853 / j. 0067 - 1975.34.1982.434"]}

Alpheus splendidus Couti��re, 1897 (Fig. 6F) Alpheus splendidus Couti��re, 1897a: 235; De Man, 1924: 41; 1929: 23; Crosnier & Forest, 1966: 361; Banner & Banner, 1982: 56; Chace, 1988: 54; Anker & De Grave, 2016: 368. Material examined. PG : 1 male, 1 female (ZUTC 6672) Hormuz I., under large boulder, Coll. A. Sharifi, 4 Dec. 2015; 3 males (ZUTC 6670), Abu-Musa I., under large boulder, Coll. A. Dehghani, 10 May 2016; GO: 1 male, 1 female (ZUTC 6671), Chabahar Bay, Tis, under large boulder, Coll. A. Dehghani, 2 Oct. 2016. Distribution. Indo-West Pacific: Red Sea, PG, GO, Singapore, Malaysia, Indonesia, Thailand, the Philippines, Hong Kong, Australia and Southern China. Regional records. PG and GO: Iran (present study). Habitat. This species was collected from sandy-boulder shores and under dead corals and boulders in lower intertidal to 3 m deep. Remarks. Anker & De Grave (2016) recorded two color patterns for this species in Singapore and mentioned it as a cryptic or pseudo-cryptic species in the Indo-West Pacific. However, the specimens of current study show uniformity in the color pattern and mostly agree with material from Singapore figured by Anker & De Grave (2016: fig. 29c)., Published as part of Dehghani, Amir, Sari, Alireza & Naderloo, Reza, 2019, Annotated checklist of the snapping shrimps of the genus Alpheus Fabricius, 1798 (Crustacea: Decapoda: Alpheidae) from the Persian Gulf and Gulf of Oman, Iran, pp. 479-504 in Zootaxa 4544 (4) on page 491, DOI: 10.11646/zootaxa.4544.4.2, http://zenodo.org/record/2618475, {"references":["Coutiere, H. (1897 a) Note sur quelques especes du genre Alpheus du Musee de Leyde. Notes from the Leyden Museum, 19, 195 - 207.","De Man, J. G. (1924) On a collection of macrurous decapod Crustacea, chiefly Penaeidae and Alpheidae from the Indian Archipelago. Archiv fur Naturgeschichte, Abteilung A, 90, 1 - 60.","Crosnier, A. & Forest, J. (1966) Resultats Scientifiques des Campagnes de la \" Calypso \". Fascicule VII. Campagne de la Calypso dans la Golfe de Guinee et aux Iles Principe, Sao Tome et Annobon (1956), et campagne aux Iles du Cap Vert (1959) (suite). 19. Crustaces decapodes: Alpheidae. Annales de l'Institut Oceanographique, Monaco, 44, 199 - 314.","Banner, D. M. & Banner, A. H. (1982) The alpheid shrimp of Australia. Part III: The remaining alpheids, principally the genus Alpheus and the family Ogyrididae. Records of the Australian Museum, 34, 1 - 357. https: // doi. org / 10.3853 / j. 0067 - 1975.34.1982.434","Chace, F. A. Jr. (1988) The caridean shrimps (Crustacea: Decapoda) of the Albatross Philippine Expedition, 1907 - 1910, Part 5: family Alpheidae. Smithsonian Contributions to Zoology, 466, 1 - 99.","Anker, A. & De Grave, S. (2016) An updated and annotated checklist of marine and brackish caridean shrimps of Singapore (Crustacea, Decapoda). Raffles Bulletin of Zoology, 34, 343 - 454."]}

Alpheus lobidens De Haan, 1849 (Fig. 4 A���D) Alpheus lobidens De Haan, 1849: 179; Banner & Banner, 1985: 19; Chace, 1988: 34; Hayashi, 1998: 394; Naderloo & T��rkay, 2012:10; Anker & De Grave, 2016: 364. Alpheus lobidens lobidens ��� Banner & Banner, 1974: 430; Banner & Banner, 1978: 223; Banner & Banner, 1982: 252. Alpheus lobidens polynesica ��� Banner & Banner, 1974: 429; Banner & Banner, 1982: 256. Alpheus crassimanus Heller, 1862a: 526; Johnson, 1962: 53; Banner & Banner, 1966a: 138; Johnson, 1979: 36. Material examined. PG : 2 males, 3 females (ZUTC 6758), Bandar-Mahshar, under rock in turbid water, Coll. A. Dehghani, 26 Nov. 2015; 1 male, 1 female (ZUTC 6759), Arvand-Kenar, under rock in turbid water, Coll. A. Dehghani, 25 Nov. 2015; 3 males, 2 ovig. females (ZUTC 6760), Bandar-Deylam, under rock, Coll. R. Naderloo, 28 Oct. 2014; 3 males, 3 females (ZUTC 6761), Bandar-Genaveh, under rocks and from burrows, Coll. A. Sari and H. Salehi, 28 Nov. 2012; 1 male (ZUTC 6762), Bandar-Dayyer, under rocks and from burrows, Coll. R. Naderloo, 1 Oct. 2012; 2 males, 2 ovig. females (ZUTC 6763), Bandar-Lengeh, under rock, Coll. A. Dehghani and H. Ashrafi, 2 May 2015; 3 males, 1 female (ZUTC 6764) Hormuz I., under rocks and from burrows, Coll. A. Sari, 2 Dec. 2014; 3 males (ZUTC 6765) Abu-Musa I., under large boulder or its inside holes and passageways, Coll. A. Dehghani, 10 May 2016; 1 male, 1 female, (SMF 51382), Abu-Musa I., under large boulder or inside holes and passageways, Coll. A. Dehghani, 10 May 2016; GO: 2 males, 2 ovig. females (ZUTC 6766), Abd Village, rocky/ cobble, Coll. A. Dehghani and H. Ashrafi, 2 May 2016; 1 male, 1 female (ZUTC 6767) Khalasi Estuary, mangroves, muddy/boulder, Coll. A. Sari, 2 Dec. 2013; 1 male, 1 female (ZUTC 6768) Chabahar Bay, Tis, rocky/ cobble, Coll. A. Dehghani, 2 May 2016; 2 males, 3 females (ZUTC 6769), Tang Village, sandy, Coll. S. Ebrahimnezhad, 25 Nov. 2015; 3 males, 2 females (ZUTC 6770) Djod Village, rocky/cobble, Coll. A. Dehghani and H. Ashrafi, 25 May 2017; 1 male, 2 females (ZUTC 6771) Gwatr Bay, sandy/muddy, Coll. A. Dehghani and H. Ashrafi, 25 May 2017. Distribution. Indo-West Pacific: from South Africa to PG, and GO to Japan, Australia and Hawaii. Regional records. PG: Iran (Banner & Banner 1981; Naderloo & T��rkay 2012); GO: Iran (Naderloo et al. 2015). Habitat. Rocky/cobble intertidal, usually under rocks and large pieces of coral rubble, muddy intertidal, estuaries, mangroves. This species is typically found in the intertidal and shallow subtidal 0���3 m deep. Remark. This species complex is common in PG and GO, especially under rocks and boulders in intertidal habitats. Alpheus lobidens shows several different color patterns and currently is a puzzling group. This species needs a thorough comparison of morphology and observed color pattern variations among individuals, which the authors have started. At least two species are currently distinguishable by the presence or absence of spine on merus of large chela and conspicuous black spots on the abdomen (see Anker & De Grave, 2016)., Published as part of Dehghani, Amir, Sari, Alireza & Naderloo, Reza, 2019, Annotated checklist of the snapping shrimps of the genus Alpheus Fabricius, 1798 (Crustacea: Decapoda: Alpheidae) from the Persian Gulf and Gulf of Oman, Iran, pp. 479-504 in Zootaxa 4544 (4) on pages 486-487, DOI: 10.11646/zootaxa.4544.4.2, http://zenodo.org/record/2618475, {"references":["De Haan, W. (1849) Crustacea. In: von Siebold, P. F. (Ed.), Fauna Japonica, sive Descriptio animalium, quae in itinere per Japoniam, jussu et auspiciis superiorum, qui summum in India Batava imperium tenent, suscepto, annis 1823 - 1830 collegit, notis, observationibus et adumbrationibus illustravit P. F. de Siebold. Conjunctis studiis C. J. Temminck et H. Schlegel pro Fertebratis atque W. de Haan pro Invertebratis elaborata Regis aupicus edita. Decas FI. Lugduni- Batavorum, Leiden, pp. 165 - 196.","Banner, D. M. & Banner, A. H. (1985) The alpheid shrimp of Indonesia, based upon J. G. De Man's \" The Decapoda of the Siboga Expedition, Part II. Family Alpheidae. \" (1911). Marine Research in Indonesia, 25, 1 - 79.","Chace, F. A. Jr. (1988) The caridean shrimps (Crustacea: Decapoda) of the Albatross Philippine Expedition, 1907 - 1910, Part 5: family Alpheidae. Smithsonian Contributions to Zoology, 466, 1 - 99.","Hayashi, K. I. (1998) Prawns, shrimps and lobsters from Japan (81). Family Alpheidae - Genus Alpheus. Aquabiology, 118, 390 - 395.","Naderloo, R. & Turkay, M. (2012) Decapod crustaceans of the littoral and shallow sublittoral Iranian coast of the Persian Gulf: faunistics, biodiversity and zoogeography. Zootaxa, 3374 (1), 1 - 67.","Anker, A. & De Grave, S. (2016) An updated and annotated checklist of marine and brackish caridean shrimps of Singapore (Crustacea, Decapoda). Raffles Bulletin of Zoology, 34, 343 - 454.","Banner, A. H. & Banner, D. M. (1974) Contributions to the knowledge of the alpheid shrimp of the Pacific Ocean Part XVII. Additional notes on the Hawaiian alpheids: new species, subspecies, and some nomenclatorial changes. Pacific Science, 28, 423 - 437.","Banner, D. M. & Banner, A. H. (1978) Annotated checklist of alpheid and ogyridid shrimp from the Philippine Archipelago and the South China Sea. Micronesica, 14 (2), 215 - 257.","Banner, D. M. & Banner, A. H. (1982) The alpheid shrimp of Australia. Part III: The remaining alpheids, principally the genus Alpheus and the family Ogyrididae. Records of the Australian Museum, 34, 1 - 357. https: // doi. org / 10.3853 / j. 0067 - 1975.34.1982.434","Heller, C. (1862) Neue Crustaceen, gesammelt wahrend der Weltumseglung der k. k. Fregatte Novara. Zweiter vorlaufiger Bericht. Ferhandlungen des Kaiserlich-koniglichen Zoologischbotanischen Gesellschaft in Wien, 12, 519 - 528.","Johnson, D. S. (1962) A synopsis of the Decapoda Caridea and Stenopodidea of Singapore, with notes on their distribution and a key to the genera of Caridea occurring in Malaysian waters. Bulletin of the National Museum Singapore, 30, 44 - 79.","Banner, A. H. & Banner, D. M. (1966 a) The alpheid shrimp of Thailand. The Siam Society Monograph Series, 3, 1 - 168.","Johnson, D. S. (1979) Prawns of the Malacca Straits and Singapore waters. Journal of the Marine Biological Association of India, 18, 1 - 54. [for 1976]","Banner, A. H. & Banner, D. M. (1981) Annotated checklist of the alpheid shrimp of the Red Sea and Gulf of Aden. Zoologische Ferhandelingen, 190, 1 - 99.","Naderloo, R., Ebrahimnezhad, S. & Sari, A. (2015) Annotated checklist of the decapod crustaceans of the Gulf of Oman, northwestern Indian Ocean. Zootaxa, 4028 (3), 397 - 412. https: // doi. org / 10.11646 / zootaxa. 4028.3.5"]}

Alpheus lutosus Anker & De Grave, 2009 (Fig. 4F) Alpheus lutosus Anker & De Grave, 2009; De Grave & Ashelby, 2011, Naderloo & T��rkay, 2012: 10. Material examined. PG : 2 males, 3 females (ZUTC 6706) Bandar-Mahshar, muddy flat, Coll. R. Naderloo and A. Dehghani, 26 Nov. 2015; 3 males, 2 females (ZUTC 6707), Bandar-Deylam, muddy flat, Coll. R. Naderloo & A. Dehghani, 28 Nov. 2015; 1 male, (SMF 51383), Bandar-Mahshar, muddy flat, Coll. R. Naderloo & A. Dehghani, 26 Nov. 2015. Distribution. It seems to be endemic to the Persian Gulf (Iran and Kuwait). Regional records. PG: Iran (Naderloo & T��rkay, 2012, present study), Kuwait (Anker & De Grave 2008); GO: No record. Habitat. In intertidal and estuarine mudflats, in burrows made under boulders or in muddy banks. Remarks. According to Anker and De Grave (2009), David Jones collected a series of Alpheus from the intertidal mudflats of Kuwait in 2004 and provisionally identified these as A. euphrosyne De Man, 1897. Later, this material was examined and was designated to a new species A. lutosus Anker & De Grave (2009). This species is easily distinguished from its PG/GO congeners by the presence of a distinct spine-like tooth on the mesial surface of the major chela palm. The species also has a distinctive blue patch on the distal region of uropods. The collected specimens agree well with the description and illustrations provided by Anker & De Grave (2009). These authors also mentioned that A. lutosus is syntopic with A. lobidens De Haan, 1849, and A. rapax Fabricius, 1798, in the muddy estuaries of Kuwait. In our samples, A. lutosus was only found syntopic with A. euphrosyne De Man, 1897., Published as part of Dehghani, Amir, Sari, Alireza & Naderloo, Reza, 2019, Annotated checklist of the snapping shrimps of the genus Alpheus Fabricius, 1798 (Crustacea: Decapoda: Alpheidae) from the Persian Gulf and Gulf of Oman, Iran, pp. 479-504 in Zootaxa 4544 (4) on pages 487-488, DOI: 10.11646/zootaxa.4544.4.2, http://zenodo.org/record/2618475, {"references":["Anker, A. & De Grave, S. (2009) A new snapping shrimp (Crustacea Decapoda, Alpheidae, Alpheus) from the estuarine mudflats of Kuwait. Zoologische Mededelingen (Leiden), 83 (25), 811 - 817.","De Grave, S. & Ashelby, C. W. (2011) Notes on some shrimp species (Decapoda: Caridea) from Qatar, with an annotated checklist of marine caridean species known from the Persian Gulf. Marine Biodiversity Records, 4, 1 - 7.","Naderloo, R. & Turkay, M. (2012) Decapod crustaceans of the littoral and shallow sublittoral Iranian coast of the Persian Gulf: faunistics, biodiversity and zoogeography. Zootaxa, 3374 (1), 1 - 67.","De Man, J. G. (1897) Bericht uber die von Herrn Schiffscapitan Storm zu Atjeh, an den westlichen Kusten von Malakka, Borneo und Celebes sowie in der Java-See gesammelten Decapoden und Stomatopoden. Funfter Theil. Zoologische Jahrbucher. Abtheilung fur Systematik, Geographie und Biologie der Thiere, 9, 725 - 790. https: // doi. org / 10.5962 / bhl. title. 16084","De Haan, W. (1849) Crustacea. In: von Siebold, P. F. (Ed.), Fauna Japonica, sive Descriptio animalium, quae in itinere per Japoniam, jussu et auspiciis superiorum, qui summum in India Batava imperium tenent, suscepto, annis 1823 - 1830 collegit, notis, observationibus et adumbrationibus illustravit P. F. de Siebold. Conjunctis studiis C. J. Temminck et H. Schlegel pro Fertebratis atque W. de Haan pro Invertebratis elaborata Regis aupicus edita. Decas FI. Lugduni- Batavorum, Leiden, pp. 165 - 196.","Fabricius, J. C. (1798) Entomologia Systematica Emendata et Aucta, Secundum Classes, Ordines, Genera, Species Adjectis Synonimis Locis Observationibus Descriptionibus. Supplementum Entomologiae Systematicae. Proft et Storch, Hafniae, 572 pp."]}