taxonID	type	description	language	source
03A00F6FFFEEFFBBFF795170FE7EF97D.taxon	description	Crangon nigrospinatus — Schmitt, 1924: 71.	en	Anker, Arthur, Hurt, Carla, Knowlton, Nancy (2008): Revision of the Alpheus websteri Kingsley, 1880 species complex (Crustacea: Decapoda: Alpheidae), with revalidation of A. arenensis (Chace, 1937). Zootaxa 1694: 51-68, DOI: 10.5281/zenodo.180675
03A00F6FFFEEFFBBFF795170FE7EF97D.taxon	materials_examined	Material examined. — Dominican Republic. 1 male (CL 7.9), MNHN-Na 16362, Bayahibe, from crevices in coral rocks, depth 1 – 2 m, coll. A. Anker, 5 Jan 2005 [fcn 05 - 020, specimen sequenced]. Guadeloupe. 1 male (CL 9.2), MNHN-Na 16241, Grand Cul de Sac, shallow subtidal, coll. F. Fasquel, 2002 [02 - 006]. Panama (Caribbean coast). 1 ovig. female (CL 7.2), USNM 1109181, Isla Grande, south of Miraculo, from coral rocks, coll. N. Knowlton et al., 15 – 16 Jun 1995 [fcn C- 1403, B- 421, specimen dissected]; 1 male (CL 8.6), 1 ovig. female (CL 6.9), USNM 1109182, same collection data as previous specimen [fcn C- 1398 + C- 1399, B- 426]; 1 male (CL 7.5), 1 ovig. female (CL 6.4), USNM 1109183, same collection data as previous specimens [fcn C- 1401 + C- 1402, B- 428]; 1 male (CL 9.5), USNM 1109184, same collection data as previous specimens [fcn C- 1410, B- 432]; 1 ovig. female (CL 6.9), USNM 1109185, Isla Grande, from coral rocks, coll. N. Knowlton et al., 15 Nov 1991 [fcn C- 171, B- 36]; 1 male (CL 7.5), USNM 1109186, same collection data as previous specimen [fcn C- 172, B- 37]; 1 male (CL 4.7), OUMNH-ZC 2007 - 13 - 033, Isla Grande, NE side, exposed rocky platform, from crevices in rocks, coll. A. Anker, J. Luque and J. Vera Caripe, 16 Sep 2007 [fcn 07 - 263]; 1 male (CL 8.7), 1 ovig. female (CL 7.3), UP, Panama, Caribbean coast, N. Knowlton et al. coll. 15 – 16 Jun 1995 [fcn B- 427, B- 430]; 1 specimen (sex and CL not determined), Panama, Bocas del Toro, Isla Colon, Bocas del Drago [fcn 06 - 500, specimen preserved in RNAlater and sequenced, not deposited]. Honduras. 1 male (CL 4.4), OUMNH-ZC 2007 - 20 - 002, Bay Islands, Utila, southern shore, Coral View reef, 1605.326 ’ N 086 º 54.652 ’ W, hand collecting, from coral rocks, depth 1 m, coll. A. Anker and S. De Grave, 5 Jul 2007 [fcn H 173]. Brazil. 1 ovig. female (CL 9.5), MNRJ 20149, Atol das Rocas, Piscina das Rocas, in calcareous algae, coll. P. S. Young, P. C. Paiva and A. A. Aguiar, 26 Oct 2000 [specimen dissected and sequenced]. Comparative material of Alpheus thomasi: Panama (Caribbean coast). 1 male (CL 5.8), USNM 1109187, Bocas del Toro, Isla Colón, between Big Creek and Playa Bluff, from crevices in coral rocks, depth 1 – 2 m, coll. A. Anker, 18 Oct 2005 [fcn 05 - 158]; 1 ovig. female (CL 7.7), USNM 1109188, Bocas del Toro, hospital Point, under large rock, depth 0.5 – 1 m, coll. A. Anker, 16 Oct 2005 [05 - 159]; 2 males (CL 8.0, 7.3), 2 ovig. females (CL 8.1, 7.8), MNHN-Na 16707, Bocas del Toro, Isla Colón, Boca del Drago, from crevices in coral rocks, depth 0.5 – 1 m, coll. A. Anker, 20 Oct 2005 [fcn 05 - 160]; 1 ovig. female (CL 6.5), USNM 1109189, Bocas del Toro, Isla Colón, Boca del Drago, from crevices in coral rocks, depth 0.5 – 1 m, coll. A. Anker, 20 Oct 2005 [fcn 05 - 157]; 1 ovig. female (CL 7.9), OUMNH-ZC 2007 - 13 - 032, Isla Grande, southern shore, from coral rubble conglomerate, depth 0.5 – 1 m, coll. A. Anker, 4 Sep 2006 [fcn 06 - 430]. FIGURE. 2. Alpheus websteri Kingsley, 1880, ovigerous female from Atol das Rocas, Brazil (MNRJ 20149); A, frontal region, dorsal view; B, same, lateral view; C, orbito-rostral process, frontal view; D, tooth on ventromesial carina of first segment of antennular peduncle, lateral view; E, lateral antennular flagellum, lateral view; F, scaphocerite of antenna, dorsal view; G, mandible, mesial view; H, third maxilliped, lateral view; I, same, detail of coxa and arthrobranch; J, major cheliped, mesial view; K, same, chela and carpus, lateral view; L, same, coxa to carpus, lateral view; M, same, detail of pollex and dactylus with setae omitted, lateral view; N, minor cheliped, lateral view; O, same, chela and carpus, mesial view; P, second pereiopod, lateral view; Q, third pereiopod, lateral view; R, same, propodus and dactylus, lateral view; S, fifth pereiopod, lateral view; T, same, propodus and dactylus, ventrolateral view; U, right uropod, dorsal view; V, telson, dorsal view. Scale bars: 1 mm.	en	Anker, Arthur, Hurt, Carla, Knowlton, Nancy (2008): Revision of the Alpheus websteri Kingsley, 1880 species complex (Crustacea: Decapoda: Alpheidae), with revalidation of A. arenensis (Chace, 1937). Zootaxa 1694: 51-68, DOI: 10.5281/zenodo.180675
03A00F6FFFEEFFBBFF795170FE7EF97D.taxon	description	Description. — Carapace smooth, not setose, laterally not compressed. Rostrum relatively short, tip slightly descendant, subacute distally, fringed with long setae (Fig. 1 A); rostral carina bluntly rounded, broad, reaching beyond eyes posteriorly (Fig. 1 A); orbito-rostral process well developed, broadly W-shaped in frontal view, with deep, rounded median notch (Fig. 2 B, C). Orbital hoods moderately inflated, distally with acute teeth (Fig. 1 A), rarely without teeth (Fig. 2 A). Frontal margin between orbital teeth and rostrum concave and sometimes slightly angular (Fig. 1 A, R). Adrostral grooves moderately deep and narrow, not abruptly delimited from rostrum and orbital hood (Fig. 1 A, R). Pterygostomial angle rounded, not protruding (Fig. 1 B); cardiac notch well developed. Eyes completely concealed in dorsal, lateral and frontal view. Ocellar beak partly concealed by orbito-rostral process, vertically protruding towards rostrum, not visible in lateral view. Antennular peduncles relatively stout, second segment distinctly longer than first, twice or more as long as wide (Fig. 1 A, R); stylocerite distally subacute, usually reaching or slightly overreaching distal margin of first segment (Fig. 1 A), sometimes reaching only to about 4 / 5 length of first segment (Fig. 2 A); mesioventral carina of first segment with minute subacute tooth as illustrated (Fig. 1 C), sometimes blunt (Fig. 2 D); lateral flagellum with numerous tufts of aesthetascs (Fig. 1 B), secondary ramus rudimentary, composed of one or two fused segments (Fig. 2 E). Antenna with basicerite bearing strong, acute ventrolateral tooth, reaching to level of stylocerite tip (Fig. 1 B); carpocerite stout, reaching slightly beyond distolateral spine of scaphocerite; distolateral tooth of scaphocerite strong, exceeding distal margin of antennular peduncle, and reaching well beyond blade (Figs. 1 A, 2 F); lateral margin of scaphocerite slightly concave (Fig. 1 A); blade narrow, subtriangular, separated from distolateral tooth by deep cleft (Figs. 1 A, 2 F). Mouthparts (mandible, maxillule, maxilla, first and second maxillipeds) typical for Alpheus; mandible with incisor process bearing 10 – 11 rounded or subtriangular distal teeth (Fig. 2 G). Third maxilliped moderately stout (Fig. 1 D); antepenultimate segment slightly flattened, ventral margin straight; penultimate segment twice as long as wide, slightly broadening distally; ultimate segment setose, with very long setae, especially on apex (Fig. 1 D); coxa with lateral plate ear-shaped distally, furnished with small setae (Fig. 1 D); exopod almost reaching penultimate segment; arthrobranch well developed. Male major cheliped (Fig. 1 F – I, S) with short, robust ischium; merus stout, about three times as long as wide proximally, dorsal margin distally blunt (Fig. 1 F, H); ventrolateral and ventromesial margins straight, latter distally with small acute tooth (Fig. 1 H), rarely without tooth (Fig. 2 J); carpus cup-shaped, without pronounced distal lobes (Fig. 1 F); chela ovate, slightly compressed laterally; mesial surface of palm smooth, very setose (Fig. 1 G); lateral surface of palm with shallow, longitudinal depression near pollex (Fig. 1 E, S), ventral and dorsal margins of palm with shallow transverse constrictions subdistally, near fingers (Fig. 1 E); linea impressa conspicuous (Fig. 1 E); fingers about half-length of palm; dactylus distally rounded, with relatively short plunger, latter distally with stamen-shaped sensillae (Fig. 1 I); adhesive discs large (Fig. 1 E, S). Female major cheliped (Fig. 2 J – M) similar to male major cheliped except for proportions of fingers to palm and general shape of palm. Male minor cheliped with merus about 2.5 times as long as wide proximally, ventrolateral and ventromesial margins straight, distally unarmed; carpus cup-shaped, distal margin with blunt process dorsomesially (Fig. 1 K); chela palm with smooth mesial and lateral surfaces, mesial surface very setose (Fig. 1 K); distomesial margin with triangular tooth (Fig. 1 J, K); linea impressa conspicuous (Fig. 1 J); fingers slightly longer than palm, not balaeniceps, with sharp, blade-like cutting margins (Fig. 1 T), tips crossing when chela closed. Female minor cheliped (Figs. 2 N, O) with merus about three times as long as wide proximally (Fig. 2 N); carpus and chela as in male but distinctly more slender (Fig. 2 O). Second pereiopod slender (Fig. 1 L); ischium slightly shorter than merus (Fig. 1 L); carpus five-segmented, ratio of carpal segments (from proximal to distal) approximately equal to 5: 3: 1.5: 1.4: 2.2 (Figs. 1 L, 2 P); chela simple, with fingers slightly longer than palm. Third and fourth pereiopods similar in shape and length; third pereiopod with ventrally unarmed ischium (Figs. 1 M, 2 Q); merus unarmed, about four times as long as wide; carpus unarmed; propodus armed with seven spines on ventral margin, one spine on distodorsal margin and two spines on distoventral margin (Figs. 1 N, 2 R); dactylus conical, gradually curved towards acute tip, biunguiculate, secondary unguis situated on flexor margin at about 3 / 4 of dactylus length (Figs. 1 N, 2 R). Fifth pereiopod (Fig. 1 S) smaller and more slender than third and fourth pereiopods; ischium unarmed ventrally; merus about six times as long as wide; propodus ventrally with six spines (including distoventral spine (Fig. 2 T), ventrolaterally with at least nine rows of grooming setae (Fig. 2 T); dactylus biunguiculate, conical, as in third and fourth pereiopods (Fig. 2 T). Abdominal segments with posteroventral margins broadly rounded; sixth segment without articulated flap, posterior margin straight, dorsolateral projections rounded; preanal plate rounded. Male second pleopod with appendix masculina reaching far beyond appendix interna, with slender, mostly apical setae (Fig. 1 O); in larger males setae present along outer margin, from about mid-length to apex. Uropod (Fig. 1 P) with protopod bearing one robust acute lateral tooth and one subacute mesial tooth distally; exopod with lateral portion of diaeresis bearing four teeth: one blunt tooth at about mid-length, one strong acute tooth mesial to distolateral spine and two acute teeth lateral to distolateral spine = doubled distolateral tooth (Fig. 1 P); distolateral spine very stout, especially in males, reaching beyond posterior margin of exopod, black or dark brown (Figs. 1 P, 2 U); distal margins of exopod without spinules. Telson (Figs. 1 Q, 2 V) moderately broad, more or less tapering towards posterior margin; dorsal surface without median groove, with two pairs of strong dorsal spines, situated at some distance from lateral margins, anterior and posterior to telson mid-length, respectively (Figs. 1 Q, 2 V); posterior margin almost straight (Fig. 1 Q) to slightly convex (Fig. 2 V), with two pairs of posterolateral spines, mesial being three times longer than lateral; anal tubercles well developed. Gill formula typical for Alpheus. Color pattern. — Background color pale grey to cream whitish; carapace with two broad transverse brown bands, one band at about mid-length, extending anteriorly on mid-dorsal line, and one more posterior band, extending anteriorly dorsolaterally (Fig. 3 A); both bands abruptly changing direction on carapace flanks (Fig. 3 D); two small short brown bands present anterolaterally, ventrally and posteroventrally to orbital hoods (Fig. 3 D); rostral area also brownish, in particular rostral carina; inner organs visible through carapace dorsally as large brownish patch; abdomen with six broad (except first) straight transverse brown bands extending to ventral margin of pleurae, abruptly changing direction laterally (towards posterior); second abdominal band bifurcating laterally, reverse V-shaped (Fig. 3 D); sixth abdominal somite almost completely brown; white bands between bands with sparse but quite large spots of egg-yellow chromatophores (Fig. 3 A, C); tail fan mostly purple-brown, with large white areas and bright egg-yellow spots, proximal portion of telson and uropods whitish, with egg-yellow spots, dorsal spines yellowish or whitish; distolateral spine black; walking legs and second pereiopods appearing pinkish semitransparent speckled with minute red chromatophores except on articulations; cheliped merus and carpus brown laterally; mesial face of palm of major chela mostly pale brown with white areas and brown patches and fairly large dark brown spots on distal half (Fig. 3 B, D), most proximal portion of palm white; pollex brown with whitish tip; dactylus brown-white with dark spots proximally, pink distally (Fig. 3 B); palm of minor chela brown-orange, with white areas and brown spots; fingers dark brown, white towards tip (Fig. 3 B, D); antennular peduncles brown distally, whitish proximally; basicerite and scaphocerite mostly whitish, with occasional brown-orange spots, lateral margin and distolateral tooth of scaphocerite brown-orange; antennular and antennal flagella pale brown; corneas grey-brown (Fig. 3 A – D); juveniles similar, but with more contrasting pattern on chelae and narrower bands on abdomen (Fig. 4 A). Size. — The two largest examined specimens are a male from Isla Grande, Panama, with CL 9.5 mm and TL 25.3 mm, and the aberrant female from Atol das Rocas, Brazil with CL 9.5 mm and TL 30 mm. Ecology. — Intertidal and shallow subtidal, probably to about 10 m, in coral and rock crevices on sand and coral-sand bottoms.	en	Anker, Arthur, Hurt, Carla, Knowlton, Nancy (2008): Revision of the Alpheus websteri Kingsley, 1880 species complex (Crustacea: Decapoda: Alpheidae), with revalidation of A. arenensis (Chace, 1937). Zootaxa 1694: 51-68, DOI: 10.5281/zenodo.180675
03A00F6FFFEEFFBBFF795170FE7EF97D.taxon	materials_examined	Type locality. — Florida Keys, Florida, USA.	en	Anker, Arthur, Hurt, Carla, Knowlton, Nancy (2008): Revision of the Alpheus websteri Kingsley, 1880 species complex (Crustacea: Decapoda: Alpheidae), with revalidation of A. arenensis (Chace, 1937). Zootaxa 1694: 51-68, DOI: 10.5281/zenodo.180675
03A00F6FFFEEFFBBFF795170FE7EF97D.taxon	distribution	Distribution. — Western Atlantic: southeastern USA: Florida Keys (Kingsley 1880); Bahamas: New Providence (Rankin 1898); eastern Mexico: Quintana Roo (Bahía de la Ascensión, Bahía del Espíritu Santo) and Cozumel (Chace 1972; Hernández Aguilera et al. 1996; McClure 2005); Honduras: Utila (present study); Cuba: Batabano Gulf (Martínez-Iglesias et al. 1997); Dominican Republic: Bayahibe; Panama: Bocas del Toro, Isla Grande, Colón (Cubit & Williams 1983; present study); Venezuela: Los Roques and Distrito Federal (Rodríguez 1980); Lesser Antilles: Anguilla, Antigua, Saint Lucia, Tobago (Chace 1972); Guadeloupe (present study); Barbados (Schmitt 1924); northeastern Brazil: Fernando do Noronha (Pocock 1890) and Atol das Rocas (present study). An unpublished record from Alagoas, Brazil (Souza 2001) needs confirmation; the almost schematic illustrations in this study may actually represent A. thomasi and not A. websteri (A. Anker, pers. obs.).	en	Anker, Arthur, Hurt, Carla, Knowlton, Nancy (2008): Revision of the Alpheus websteri Kingsley, 1880 species complex (Crustacea: Decapoda: Alpheidae), with revalidation of A. arenensis (Chace, 1937). Zootaxa 1694: 51-68, DOI: 10.5281/zenodo.180675
03A00F6FFFEEFFBBFF795170FE7EF97D.taxon	discussion	Remarks. — The ovigerous female from Atol das Rocas (MNRJ 20149) (Fig. 2) is remarkable in lacking orbital teeth (Fig. 2 A), bearing a short, ascendant rostrum (Fig. 2 B) and having no trace of a distomesial tooth on the merus of the major cheliped (Fig. 2 J). In all other features of the chelipeds (Fig. 2 J – O), second to fifth pereiopods (Fig. 2 P – S), tail fan (Fig. 2 U, V), third maxilliped (Fig. 2 H, I), antennules and antenna (Fig. 2 A, B, F), this specimen agrees well with the Caribbean A. websteri specimens. The more slender minor chela (Fig. 2 N, O) and the less stout uropodal spine (Fig. 2 U) appear to occur generally in females of A. websteri. Moreover, the color of the uropodal spine in females of A. websteri apparently varies from brown, as in the female from Atol das Rocas (Fig. 2 U), to shiny black, as in most Caribbean females (Fig. 1 P). The shape of the ventromesial carina of the first segment of the antennular peduncle – blunt in the specimen from Atol das Rocas (Fig. 2 D) vs. with small subacute tooth in the specimen from Panama (Fig. 1 C) – may be somewhat variable, as observed in other specimens of A. websteri from the Caribbean, as well as other Alpheus species (A. Anker, pers. obs.). A comparison of the COI gene sequence between the Caribbean specimens and the female from Atol das Rocas resulted in the latter falling within the variation range of the Caribbean samples. Therefore, this single specimen is considered as an aberrant individual of A. websteri. The lack of distinct orbital teeth and distomesial tooth on the chelipeds in large specimens could be due to wearing away from use or from bluntening (polishing) during molting (see also under A. rugimanus). The abnormal frontal region can also be result of a physical damage (injury). Unfortunately, no other specimens of A. websteri were collected at Atol das Rocas. In the field, A. websteri may be confused with the sympatric A. thomasi, which ranges from southern Florida to Panama and Brazil (Hendrix & Gore 1973; Christoffersen 1998; present study). However, the latter species can be easily distinguished from A. websteri by the frontal margin between the rostrum and the orbital hoods being more deeply notched; the slenderer major and minor chelae; the uropod with a distinctly narrower distolateral spine and with a simple distolateral tooth on the diaeresis (doubled in A. websteri); and several features on the third pereiopod, including the distinctly slenderer merus, a simple dactylus (biunguiculate in A. websteri), and most importantly, the presence of a ventrolateral spine on the ischium (absent in A. websteri) (see Hendrix & Gore 1973). Alpheus thomasi can also be distinguished from A. websteri by several details of the color pattern, the most important being the absence of spots on the major chela (compare Figs. 5 D, E and 4 A – D). GenBank number. — EU 339467 [05 - 020, MNHN-Na 16362, Dominican Republic], EU 339468 [06 - 500, not deposited, Panama], EU 339469 [F 119, not deposited, Panama], EU 339470 [AA-AC- 42, MNRJ 20149, Atol das Rocas, Brazil].	en	Anker, Arthur, Hurt, Carla, Knowlton, Nancy (2008): Revision of the Alpheus websteri Kingsley, 1880 species complex (Crustacea: Decapoda: Alpheidae), with revalidation of A. arenensis (Chace, 1937). Zootaxa 1694: 51-68, DOI: 10.5281/zenodo.180675
03A00F6FFFE5FFB8FF7956BDFB39FCCA.taxon	materials_examined	Material examined. — Panama (Pacific coast). 1 male (CL 4.1), USNM 1109190, Las Perlas Islands, Saboga, shallow subtidal, from rock crevices, coll. A. Anker, J. Jara and C. Hurt, 15 Nov 2005 [fcn 05 - 156]; 1 male (CL 9.7), 1 female (CL 9.3), USNM 1109191, Amador Causeway near Panama City, Isla Perico, intertidal, coll. N. Knowlton et al., 22 Jan 1992 [fcn C- 292 and C- 293, B- 68]; 1 male (CL 9.3), USNM 1109192, Río Mar, intertidal, coll. N. Knowlton et al., 16 – 20 Feb 1992 [fcn C- 367, B- 103]; 1 male (CL 8.7), USNM 1109193, same collection data as previous specimen [fcn C- 363, B- 99]; 1 male (CL 6.7), USNM 1109194, same collection data as previous specimen [fcn C- 365, B- 101]; 1 ovig. female (CL 6.5), MNHN-Na 16387, Coiba Marine National Park, Coibita, mud-rock intertidal, under rock, coll. A. Anker, I. Marin, J. Jara, E. Gómez and E. Tóth, 20 Mar 2007 [fcn 07 - 111].	en	Anker, Arthur, Hurt, Carla, Knowlton, Nancy (2008): Revision of the Alpheus websteri Kingsley, 1880 species complex (Crustacea: Decapoda: Alpheidae), with revalidation of A. arenensis (Chace, 1937). Zootaxa 1694: 51-68, DOI: 10.5281/zenodo.180675
03A00F6FFFE5FFB8FF7956BDFB39FCCA.taxon	description	Description. — See Chace (1937) as Crangon arenensis; Kim & Abele (1988) as Alpheus websteri. Color pattern. — Background color cream white or pale grey; carapace with two broad transverse chestnut to brown bands, one band at about mid-length, extending anteriorly on mid-dorsal line, and one more posterior band, constricted on anterior and posterior margins, and extending anteriorly dorsolaterally, almost reaching first band (Fig. 3 E), defining “ bikini ” pattern in dorsal view; both bands abruptly changing direction on carapace flanks (Fig. 3 E); two small short brown bands present anterolaterally, ventrally and posteroventrally to orbital hoods (Fig. 3 E); rostral area also brownish, in particular rostral carina; inner organs visible through carapace dorsally as large brownish patch; articulation zone between carapace and abdomen and posterior carapace sharing white rhomboid patch on brown background (Fig. 3 E); abdomen with six broad (except first) chestnut to brown transverse bands with concave margins, extending to ventral margin of pleurae, abruptly changing direction laterally (towards posterior) and nearly touching on flanks (Fig. 3 E); second abdominal band bifurcating laterally, reverse V-shaped (Fig. 3 D); sixth abdominal somite almost completely brown except for transversely elongate white patch (Fig. 3 E); white bands between brown bands with sparse spots of pale-yellow chromatophores; tail fan mostly purple-brown with large white areas, proximal portion of telson and uropods whitish, dorsal spines whitish; distolateral spine black; walking legs and second pereiopods appearing pinkish semitransparent speckled with minute red chromatophores except on articulations; cheliped merus and carpus brown laterally; mesial face of palm of major chela mostly pale brown with white areas and brown patches and fairly large dark brown spots on distal half (Fig. 3 E), most proximal portion of palm white; pollex brown with whitish tip; dactylus brown-white with dark spots proximally, pink distally (Fig. 3 E); palm of minor chela brown-orange, with white areas and brown spots; fingers dark brown, paler towards tip; antennular peduncles brown distally, whitish proximally; basicerite and scaphocerite mostly whitish with occasional brown-orange spots, lateral margin and distolateral tooth of scaphocerite brown; antennular and antennal flagella pale brown; corneas dark brown-grey (Fig. 3 E); juveniles similar, but with more contrasting pattern on chelae, carapace, abdomen and tail fan; yellow chromatophores more intense and forming transverse bands fringing brown-grey bands on carapace and abdomen (Fig. 4 B). Size. — The largest specimen from the Pacific coast of Panama is a male from Isla Perico with CL 9.7 mm, TL 24.8 mm. The CL of the Gulf of California specimens ranged from 4.4 to 8.0 mm, and TL from 17.0 to 21.6 mm (Villalobos 2000). Ecology. — Intertidal to about 6 m (Wicksten 1983); in crevices of rocks and under coral rubble on sand or mixed sand-coral bottoms.	en	Anker, Arthur, Hurt, Carla, Knowlton, Nancy (2008): Revision of the Alpheus websteri Kingsley, 1880 species complex (Crustacea: Decapoda: Alpheidae), with revalidation of A. arenensis (Chace, 1937). Zootaxa 1694: 51-68, DOI: 10.5281/zenodo.180675
03A00F6FFFE5FFB8FF7956BDFB39FCCA.taxon	materials_examined	Type locality. — Arena Bank, Gulf of California, Mexico.	en	Anker, Arthur, Hurt, Carla, Knowlton, Nancy (2008): Revision of the Alpheus websteri Kingsley, 1880 species complex (Crustacea: Decapoda: Alpheidae), with revalidation of A. arenensis (Chace, 1937). Zootaxa 1694: 51-68, DOI: 10.5281/zenodo.180675
03A00F6FFFE5FFB8FF7956BDFB39FCCA.taxon	distribution	Distribution. — Eastern Pacific: Mexico: central and southern Gulf of California (e .. g., Arena Bank, Sonora) and Revillagigedo Islands (Socorro) (Chace 1937; Carvacho & Ríos 1982; Wicksten 1983; Wicksten & Hendrickx 2003); Costa Rica: Guanacaste, Puntarenas (Kim & Abele 1988); Panama: Las Perlas (Kim & Abele 1988; present study); Colombia (Wicksten 1983); Galapagos (Wicksten 1993; Wicksten & Hendrickx 2003).	en	Anker, Arthur, Hurt, Carla, Knowlton, Nancy (2008): Revision of the Alpheus websteri Kingsley, 1880 species complex (Crustacea: Decapoda: Alpheidae), with revalidation of A. arenensis (Chace, 1937). Zootaxa 1694: 51-68, DOI: 10.5281/zenodo.180675
03A00F6FFFE5FFB8FF7956BDFB39FCCA.taxon	discussion	Remarks. — The only noticeable morphological difference between A. websteri and A. arenensis lies in the development of the small distal tooth on the ventromesial margin of the major cheliped merus: acute in A. websteri (Fig. 1 H) and blunt or absent in A. arenensis (see Kim & Abele 1988, fig. 11 f). A side by side comparison of two large adults of each species revealed two further, although very subtle differences: the orbital grooves appear to be deeper in A. arenensis compared to those of A. websteri, while the scaphocerite blade is shorter in A. arenensis compared to that of A. websteri, reaching to about 3 / 5 of the scaphocerite length (vs. 3 / 4 in A. websteri). Differences in color between the two species are more numerous both in adults (Fig. 3 D, E) and juveniles (Fig. 4 A, B), including the more concave margins of the abdominal bands in A. arenensis (vs. almost straight in A. websteri); the “ bikini ” - shaped posterior band on the carapace (this band is subrectangular in A. websteri); and the presence of a white dorsomedian patch on the sixth abdominal somite (absent in A. websteri) (see also Table 1).	en	Anker, Arthur, Hurt, Carla, Knowlton, Nancy (2008): Revision of the Alpheus websteri Kingsley, 1880 species complex (Crustacea: Decapoda: Alpheidae), with revalidation of A. arenensis (Chace, 1937). Zootaxa 1694: 51-68, DOI: 10.5281/zenodo.180675
03A00F6FFFE5FFB8FF7956BDFB39FCCA.taxon	description	GenBank number. — EU 339466 [99 - 041, not deposited, Panama]. Feature A. websteri (WA) A. arenensis (EP) A. fagei (EA)	en	Anker, Arthur, Hurt, Carla, Knowlton, Nancy (2008): Revision of the Alpheus websteri Kingsley, 1880 species complex (Crustacea: Decapoda: Alpheidae), with revalidation of A. arenensis (Chace, 1937). Zootaxa 1694: 51-68, DOI: 10.5281/zenodo.180675
03A00F6FFFE6FFBDFF7956CAFED0FD3D.taxon	materials_examined	Material examined. — São Tomé. 1 female (CL 5.0), USNM 1109195, ST 5, Lagoa Azul, shallow bay with rocky shore, from coralline algae and rocks, depth 1 – 3 m, coll. N. Knowlton and A. Anker, 3 Feb 2006 [fcn 06 - 139]; 1 male (CL 7.4), 1 ovig. female (CL 9.6), USNM 1109196, ST 10, Ilha das Cabras, off beach near light house, depth about 2 m, coarse sand and rubble, under rocks and rock crevices, coll. A. Anker, 8 Feb 2006 [fcn 06 - 179]; 1 ovig. female (CL 5.8), USNM 1109197, ST 5 A, Lagoa Azul, shallow bay with rocky shore, from coralline algae and rocks, depth 1 – 3 m, coll. N. Knowlton and A. Anker, 6 Feb 2006 [fcn 06 - 163]; 1 male (CL 5.1) 1 ovig. female (CL 6.6), USNM 1109198, same collection data as previous specimen [fcn 06 - 161, 06 - 162]; 1 ovig. female, CL 10.1, USNM 1109199, ST 3, 200 m west of Lagoa Azul, rocky shore with coralline algae, from holes and under rocks, intertidal, extreme low tide, coll. N. Knowlton and A. Anker, 31 Jan 2006 [fcn 06 - 065]; 1 male (CL 8.7), 1 ovig. female (CL 9.1) MNHN-Na 16708, ST 6 A: São Tome town, near fortress, shallow bay with sand-mud and scattered rocks, under rocks, depth 0.5 – 1 m, coll. N. Knowlton and A. Anker, 4 Feb 2006 [fcn 06 - 168, 06 - 167]; 1 juvenile (CL not measured), USNM 1109200, ST 2, Praia Lagarto, near hospital, sand with Montastraea, zoanthids and rocks embedded in sand, intertidal (extreme low tide), coll. N. Knowlton and A. Anker, 30 Jan 2006 [fcn 06 - 051].	en	Anker, Arthur, Hurt, Carla, Knowlton, Nancy (2008): Revision of the Alpheus websteri Kingsley, 1880 species complex (Crustacea: Decapoda: Alpheidae), with revalidation of A. arenensis (Chace, 1937). Zootaxa 1694: 51-68, DOI: 10.5281/zenodo.180675
03A00F6FFFE6FFBDFF7956CAFED0FD3D.taxon	description	Description. — See Crosnier & Forest (1965, 1966). FIGURE. 4. Color patterns of juveniles of Alpheus websteri Kingsley, 1880 (A); Alpheus arenensis (Chace, 1937) (B); Alpheus fagei Crosnier and Forest, 1966 (C); A, specimen from Utila, Honduras (OUMNH 2007 - 20 - 002); B, specimen from Las Perlas Islands, Pacific coast of Panama (USNM 1109190); C, specimen from São Tomé (USNM 1109200). Color pattern. — Background color cream white or pale grey; carapace with two broad transverse brown bands, one band at about mid-length, somewhat extending anteriorly on mid-dorsal line (Fig. 5 A, B), and one more posterior band, with concave anterior margin and more or less sinuous posterior margin (Fig. 5 A, B), and extending anteriorly dorsolaterally, but not reaching first band (Fig. 5 C), both bands abruptly changing direction on carapace flanks (Fig. 5 C); anterolateral areas of carapace below orbital hoods dark brown (Fig. 5 C); rostral area also brownish, with pale greenish-brown rostral carina; inner organs visible through carapace dorsally as large purple-brown patch; abdomen with six broad dark brown or chocolate brown transverse bands with slightly concave posterior margins, extending to ventral margin of pleurae, abruptly changing direction laterally (towards posterior) but not touching on flanks (Fig. 5 C); second abdominal band bifurcating laterally, reverse V-shaped (Fig. 5 C); sixth abdominal somite almost completely dark brown except for large yellow spot posteriorly (Fig. 5 A, B); white bands between brown bands with intense egg-yellow bands anteriorly, fringing posterior margins of brown bands (Fig. 5 A, B); tail fan mostly purple-brown, with large white areas and spots of yellow chromatophores, proximal portion of telson and uropods whitish, dorsal spines whitish; distolateral spine black; walking legs and second pereiopods pinkish semitransparent speckled with minute red chromatophores except on articulations; cheliped merus and carpus orange-brown laterally; mesial face of palm of major chela orange-brown with paler areas (Fig. 5 B) and sometimes transverse white streaks (Fig. 5 A); pollex darker orange-brown with whitish tip; dactylus brown-grey proximally, pink distally (Fig. 5 A); palm of minor chela brown-orange, white proximally; fingers dark brown, pale grey towards tip; antennular peduncles brown distally, whitish proximally; basicerite and scaphocerite mostly whitish with occasional brown-orange spots, lateral margin and distolateral tooth of scaphocerite brown; antennular and antennal flagella pale brown; corneas grey (Fig. 5 A – C); juveniles with paler bands but more contrasting pattern, especially marked white patches on major chela (Fig. 4 C). Size. — The largest examined specimen is an ovigerous female with CL 10.1 mm and TL 27.9 mm. FIGURE. 5. Color patterns of adults of Alpheus fagei Crosnier & Forest, 1966 (A – C) and Alpheus thomasi Hendrix & Gore, 1973 (D, E); A, female from São Tomé (USNM 1109199); B, C, male from São Tomé (MNHN-Na 16708); D, male from Bocas del Toro, Caribbean coast of Panama (MNHN-Na 16707); E, ovigerous female from Bocas del Toro, Caribbean coast of Panama (USNM 1109188). Notice similarities, especially in the presence of narrow yellow bands fringing brown bands and the bifurcating band on the second abdominal somite. Ecology. — Lower intertidal to about 35 m (Crosnier & Forest 1966); hard bottoms, in crevices of calcareous algal crusts and overgrown rocks, also in coral rubble and under rocks on mixed sand-rock bottoms.	en	Anker, Arthur, Hurt, Carla, Knowlton, Nancy (2008): Revision of the Alpheus websteri Kingsley, 1880 species complex (Crustacea: Decapoda: Alpheidae), with revalidation of A. arenensis (Chace, 1937). Zootaxa 1694: 51-68, DOI: 10.5281/zenodo.180675
03A00F6FFFE6FFBDFF7956CAFED0FD3D.taxon	materials_examined	Type locality. — São Tomé, Gulf of Guinea.	en	Anker, Arthur, Hurt, Carla, Knowlton, Nancy (2008): Revision of the Alpheus websteri Kingsley, 1880 species complex (Crustacea: Decapoda: Alpheidae), with revalidation of A. arenensis (Chace, 1937). Zootaxa 1694: 51-68, DOI: 10.5281/zenodo.180675
03A00F6FFFE6FFBDFF7956CAFED0FD3D.taxon	distribution	Distribution. — Eastern Atlantic: Ivory Coast; Gulf of Guinea: São Tomé and Principe (Crosnier & Forest 1966).	en	Anker, Arthur, Hurt, Carla, Knowlton, Nancy (2008): Revision of the Alpheus websteri Kingsley, 1880 species complex (Crustacea: Decapoda: Alpheidae), with revalidation of A. arenensis (Chace, 1937). Zootaxa 1694: 51-68, DOI: 10.5281/zenodo.180675
03A00F6FFFE6FFBDFF7956CAFED0FD3D.taxon	discussion	Remarks. — Alpheus fagei differs from A. websteri and A. arenensis mainly by the shorter basicerite, not reaching level of stylocerite (vs. reaching to or slightly beyond stylocerite in A. websteri and A. arenensis); the third pereiopod bearing a simple dactylus (vs. biunguiculate in A. websteri and A. arenensis); and in life, by the palm of the major chela lacking dark spots (vs. with spots in A. websteri and A. arenensis) (see also Table 1). The color pattern of A. fagei is remarkably similar to that of the western Atlantic A. thomasi (Fig. 5). The main color differences between the two species are the relatively broader bands on the carapace in A. fagei, and a brighter, more contrasting pattern on the major claw in A. thomasi. In addition to these, A. fagei may be easily separated from A. thomasi by the same morphological criteria as A. websteri (see above). GenBank number. — EU 339464 [06 - 086, not deposited, São Tomé], EU 339465 [06 - 119, not deposited, São Tomé].	en	Anker, Arthur, Hurt, Carla, Knowlton, Nancy (2008): Revision of the Alpheus websteri Kingsley, 1880 species complex (Crustacea: Decapoda: Alpheidae), with revalidation of A. arenensis (Chace, 1937). Zootaxa 1694: 51-68, DOI: 10.5281/zenodo.180675
