Alpheus punctatus, Anker, 2012

Anker, Arthur, 2012, Revision of the western Atlantic members of the Alpheus armillatus H. Milne Edwards, 1837 species complex (Decapoda, Alpheidae), with description of seven new species, Zootaxa 3386 (1), pp. 1-109 : 50-56

Alpheus punctatus sp. nov.

( Figures 33–37 , 64F , 65F , 66F , 67F , 69E )

Alpheus cf. armillatus D— Mathews & Anker 2009: 277.

Type material. Panama: holotype: 1 male (cl 10.5), MNHN-IU-2010-4085, Bocas del Toro , Isla Colón, Punta Caracol, sand flat with rubble, sponges, clumps of Halimeda , 1–2 m, leg. A. Anker, 30.03.2008 [fcn 08-005a, dissected]; paratypes: 1 ov. female (cl 10.7), MNHN-IU-2010-4086, same collection data (mated pair) [fcn 08-005b]; 1 male (cl 11.6), MNHN-IU-2010-4087, same collection data [fcn 08-003]; 1 male (cl 12.6), 1 ov. female (cl 11.7), RMNH D54819, Bocas del Toro , Isla San Cristobal, Punta Coco, shallow subtidal sand flat with turtle grass and abundant rubble, under large pieces of rubble, 0.5–1 m, leg. A. Anker, 29.10.2005 [fcn 05-187]; 1 ov. female (cl 9.9), UP, Bocas del Toro , Isla Colón, Punta STRI, sand flat with rubble, seagrass, sponges, clumps of Halimeda , 1–2 m, leg. A. Anker, J.A. Vera Caripe, J.A. Baeza, 17.11.2006 [fcn 06-595]; 1 ov. female (cl 10.4), UP, same collection data [fcn 06-594]; 1 male (cl 9.8), OUMNH. ZC. 2008-14-0101, Bocas del Toro , Isla Bastimentos, Crawl Cay (Cayo Coral), 09°14.502'N 082°08.288'W, sand flat with some seagrass and sponges, under coral rocks, 1 m, 11.08.2008, leg. A. Anker [fcn BSC-017] .

Additional material. Costa Rica: 1 male (cl 11.3), MNHN-IU-2010-4088, Cahuita, shallow subtidal sand flat with turtle grass, large pieces of rubble, clumps of Halimeda , under large pieces of rubble, ~ 1 m, leg. A. Anker, 26– 27.11.2005 [fcn 05-179]. British Virgin Islands: 1 male (cl 9.5), LACM-MBPC 11730, Guana Island, Monkey Point, ~ 18.463°N ~ 64.57°W, sta. BVI-00-046C, very open reef flat with scattered small-sized eroded coral heads with sponges and soft corals, gorgonians very common, benthos consisting mostly of sand and Acropora cervicornis rubble, ARMS-04, rubble basket (mostly pieces of old Acropora palmata ), 9 m, leg. T. Zimmerman, R. Ware, T. Haney, J.W. Martin, 22.07.2000 [fcn vc1561]. USA: 1 male (cl 8.6), RMNH D54820, Florida Keys, Seven Mile Bridge, ocean side, Halimeda -rubble bottom, 0.5 m (low tide), in rubble, leg. A. Anker, A.L. Rhyne, 19.04.2004 [fcn 04-023].

Diagnosis. Rostrum straight or slightly descendent, barely or not reaching half-length of first article of antennular peduncle; area posterior to rostral carina flattened, markedly and abruptly delimited from adjacent, deep rostro-orbital furrows, forming more or less V or lance-shaped post-rostral plate continuous with usually narrow, dorsally somewhat flattened rostral carina, margins of post-rostral plate distinctly overhanging rostro-orbital furrows; post-rostral plate situated slightly above and gently sloping into rostral carina. Antennule with sctylocerite acute distally, reaching distal margin of first article; ventromesial carina of first article with large tooth bearing small point and shallowly concave anterior margin; second article about 2.2 times as long as wide. Antenna with basicerite armed with stout distolateral tooth; scaphocerite with lateral margin shallowly concave; strong, sharp distolateral tooth reaching slightly beyond relatively broad blade; carpocerite exceeding scaphocerite blade and (very slightly) distolateral tooth, reaching beyond end of antennular peduncles. Third maxilliped with ultimate article markedly narrower than penultimate article, tapering distally. Major cheliped edwardsii - type (see under A. armillatus ). Male major cheliped with merus stout, distodorsal margin blunt, ventromesial margin somewhat rugose, with short protuberances each carrying a small spiniform seta, distomesial tooth strong; palm with dorsal shoulder rounded, descending almost vertically into adjacent transverse groove, not or very slightly overhanging groove; ventral shoulder broadly rounded, not projecting, with field of minute tubercles laterally; fingers about 0.7 length of palm; pollex with short oblique ridge mesially; dactylus plunger short, truncate distally, proximal height about 0.4 length of distolateral margin, anterior angle very slight, superior to 90°. Female major cheliped smaller than male major cheliped, with chela smaller, less stout, and with different proportions. Male minor cheliped with merus rather slender, distodorsal margin blunt, ventromesial margin with small spiniform setae and strong distomesial tooth; chela relatively slender, palm with length-height about 2.5; palm with trace of longitudinal groove on lateral face, shallowly concave ventrally, but without distinct ventral sinus or shoulder; fingers about as long palm, slender, simple, non-balaeniceps, with sharp cutting edges. Female minor cheliped generally similar to male minor cheliped, often more slender. Second pereiopod slender, with first two carpal articles longest, first about 1.5 length of second. Third and fourth pereiopods similar, moderately slender; ischium with stout spiniform seta; merus about six times as long as wide; propodus with stout spiniform setae, some inserted in pairs, along ventral margin, incud- tylus, mesial view; L, diagrammatic representation of first two abdominal sternites, ventroposterior view; M, second pleopod, detail of appendix masculina and appendix interna, lateral view; N, telson, dorsal view; O, uropod, dorsal view; P, rostro-orbital region of carapace, dorsal view; Q, same, dorsolateral view. Scale bars as indicated, figs. D, L drawn without scale. pereiopods without conspicuously projecting, slender, subacute processes. First and second abdominal sternites unarmed in both sexes. First to fourth pleopods with ventrolateral margin of protopod furnished with spiniform setae in both sexes, longer and more dense in females; male second pleopod with appendix masculina very slightly exceeding appendix interna, densely covered with stiff setae, especially on apex. Uropod with exopod and endopod broadly rounded; exopod with sinuous diaeresis and stout distolateral spiniform seta; endopod with row of small spiniform setae on distal margin. Telson broad, slightly tapering posteriorly; dorsal surface with two pairs of spiniform setae inserted far from lateral margins; posterior margin broadly rounded, with row of small spiniform setae; posterolateral angles each with two spiniform setae, mesial much longer than lateral ( Figs. 33–35 ).

Variation. As in the other species of the A. armillatus complex, the major chela of A. punctatus sp. nov. shows some sexual dimorphism, especially in size ( Fig. 37A, B ). There is some variation in the shape of the post-rostral

Size range. The Panamanian and Costa Rican specimens range from 9.8 to 12.6 mm cl in males (holotype: 10.5 mm), and from 9.9 to 10.7 mm cl in females; the specimens from Guana Island and the Florida Keys are some what smaller, at cl 9.5 mm cl and cl 8.6 mm, respectively .

Colour pattern. Body background colour pale greyish-green, dotted with small bluish-grey to brownish-grey spots, some interconnecting and forming straight, curving, or multi-branched chains, others grouped together to and carpus, darker on margins; mesial face of palm of major chela olive-grey, with bluish tinge in some areas, dorsal and ventral transversal grooves marked with pale yellow or pale orange; fingers usually darker grey, dactylus tip pink; mesial face of minor chela pale grey-yellow, with darker olive-grey finger tips; second to fifth pereiopods pale pink-yellow to reddish, whitish near articulations; pattern on telson and uropods similar to that of the abdomen ( Fig. 36 , 37 ); conspicuous dark dotting also present in juveniles ( Fig. 69E ).

Etymology. Refers to the fine dark dotting of the body (punctatus = speckled, dotted in Latin); used as an adjective.

Type locality. Caribbean coast of Panama: Bocas del Toro .

Distribution. Caribbean Sea: Panama (Bocas del Toro), Costa Rica (Cahuita), British Virgin Islands (Guana); Florida (Florida Keys) (see map in Fig. 70 ).

Ecology. Shallow subtidal rubble flats at a depth range of 0.5–9 m; most frequently found on near-shore flats with an abundance of large coral debris and piles of loose coral rubble, patches of Thalassia , and Halimeda clumps, often near mangroves; usually in male-female pairs.

Remarks. Alpheus punctatus sp. nov. belongs to Clade 1 of Mathews & Anker (2009, fig. 4), together with with the above-described A. amarillo sp. nov., and can be distinguished from all other species of the A. armillatus complex by the unique combination of morphological characters (see Table 1 and below) and a diagnostic colour pattern ( Figs. 34 , 35 ), which corresponds to the colour type NB 2 in Mathews & Anker (2009, fig. 2B).

Alpheus punctatus sp. nov. shares with A. amarillo sp. nov. the post-rostral plate V-shaped, abruptly delimited, situated slightly above the rostral carina, and with margins overhanging the adjacent rostro-orbital furrows; the ventral shoulder of the major chela with a small field of minute tubercles laterally; the major chela dactylus with a short plunger; as well as several other characters. However, A. punctatus sp. nov. can be separated from A. amarillo sp. nov. by the sternites of the first and second abdominal somites being unarmed (vs. armed with a median process in A. amarillo sp. nov.); the male minor chela without a trace of ventral sinus (more or less distinct in A. amarillo sp. nov.); and the noticeably more pronounced dactylus plunger of the major chela (cf. Figs. 28E , 34E ; see also Fig. 65 ). Alpheus punctatus sp. nov. can be separated from the more distantly related A. armillatus , A. verrilli , A. lancirostris , and A. angulosus , by essentially the same criteria as A. amarillo sp. nov. (see above and Table 1).

GenBank accession numbers. Costa Rica: FJ528570 (MyHC) , FJ528445 (16S) , FJ528489 ( COI) [MNHN- IU-2010-4088] .

Figures

FIGURE 28. Alpheus amarillo sp. nov.: A–J, holotype, male from Portobelo, Panama [MNHN-IU-2010-4081]; K, paratype, male from San Blas Islands, Panama [RMNH D54800]; L–N, paratype, ovigerous female from San Blas Islands [MNHN-IU-2010-4083]; O, P, non-type, male from Alacranes Reef, Gulf of Mexico [UNAM-CNCR 26483]; A, major chela, mesial view; B, same, lateral view; C, same, detail of ventral shoulder, lateral view; D, same, detail of dorsal shoulder, lateral view; E, major chela dactylus, lateral view; F, major cheliped, ischium, merus and carpus, mesial view; G, same, detail of ventromesial margin, mesial view; H, minor chela and carpus, mesial view; I, minor chela, lateral view; J, minor cheliped, ischium, merus and carpus, mesial view; K, major chela, mesial view; L, major chela, lateral view; M, same, detail of dorsal shoulder, lateral view; N, same, detail of ventral shoulder, lateral view; O, major chela, lateral view; P, same, detail of dorsal shoulder, lateral view. Scale bars as indicated.

FIGURE 33. Alpheus punctatus sp. nov.: A–O, holotype, male from Bocas del Toro, Panama [MNHN-IU-2010-4085]; P, Q, paratype, ovigerous female from Bocas del Toro [MNHN-IU-2010-4086]; A, frontal region of carapace and frontal appendages, dorsal view; B, rostro-orbital region of carapace, lateral view; C, same, dorsolateral view; D, tooth on mesioventral carina of first article of antennular peduncle, lateral view; E, antennal scaphocerite, dorsal view; F, third maxilliped, lateral view; G, second pereiopod, lateral view; H, third pereiopod, lateral view; I, same, propodus and dactylus, lateral view; J, fifth pereiopod, lateral view; K, same, propodus and dac-

FIGURE 34. Alpheus punctatus sp. nov.: A–J, holotype, male from Bocas del Toro, Panama [MNHN-IU-2010-4085]; K, paratype, ovigerous female from Bocas del Toro [MNHN-IU-2010-4086]; A, major chela, mesial view; B, same, lateral view; C, same, detail of ventral shoulder, lateral view; D, same, detail of dorsal shoulder, lateral view; E, major chela dactylus, lateral view; F, major cheliped, ischium, merus and carpus, mesial view; G, same, detail of ventromesial margin, mesial view; H, minor chela, mesial view; I, same, lateral view; J, minor cheliped, ischium, merus and carpus, mesial view; K, major chela, lateral view. Scale bars as indicated.

FIGURE 35. Alpheus punctatus sp. nov.: non-type, male from off Seven Mile Bridge, Florida Keys [RMNH D54820]; A, frontal region of carapace and frontal appendages, dorsal view; B, rostr-orbital region of carapace, lateral view; C, tooth on mesioventral carina of first article of antennular peduncle, lateral view; D, major chela, lateral view; E, same, mesial view; F, major cheliped, ischium, merus and carpus, mesial view; G, major chela dactylus, lateral view; H, minor cheliped, lateral view (linea impressa omitted); I, minor chela, lateral view (linea impressa omitted); J, minor cheliped, ischium, merus and caprus, mesial view; K, second pleopod, lateral view; L, third pereiopod, lateral view; M, fifth pereiopod, lateral view. Scale bars as indicated.

FIGURE 36. Alpheus punctatus sp. nov.: A, paratype, male from Bocas del Toro, Panama [RMNH D54819], dorsal view; B, paratype, male from Bocas del Toro [MNHN-IU-2010-4087], dorsal view. Photographs by the author.

FIGURE 37. Alpheus punctatus sp. nov.: A, paratype, ovigerous female from Bocas del Toro, Panama [RMNH D54819], dorsal view; B, non-type male from Guana Island, British Virgin Islands [LACM-MBPC 11730], dorsal view; C, non-type male from Florida Keys. Photographs: A, by the author; B, by T.L. Zimmerman / J.W. Martin; C, by the author and A.L. Rhyne.

FIGURE 64. Comparison of proportions of the major chela (in lateral view) among 10 western Atlantic species of the Alpheus armillatus H. Milne Edwards, 1837 complex; A–I, males; J, female; A, A. armillatus H. Milne Edwards, 1837; B, A. lancirostris Rankin, 1900; C, A. verrilli (Schmitt, 1924); D, A. angulosus McClure, 2002; E, A. amarillo sp. nov.; F, A. punctatus sp. nov.; G, A. tampensis sp. nov.; H, A. carlae sp. nov.; I, A. brasileiro sp. nov.; J, A. nuno sp. nov. Abbreviations: FB, finger base height; FH, fixed finger (pollex) height; FL, fixed finger (pollex) length; PL, palm length. Not to scale (for scales and specimen details see Figs. 3, 10, 15, 20, 28, 34, 39, 42, 54, 58); C, F, G mirrored horizontally from the originals for better visual comparison.

FIGURE 65. Comparison of dactylus plungers of the major chela (in lateral view) among 12 western Atlantic species of the Alpheus armillatus H. Milne Edwards, 1837 complex and the related A. viridari (Armstrong, 1949); A–J, L, males; K, female; A, A. armillatus H. Milne Edwards, 1837; B, A. lancirostris Rankin, 1900; C, A. verrilli (Schmitt, 1924); D, A. angulosus McClure, 2002; E, A. amarillo sp. nov.; F, A. punctatus sp. nov.; G, A. tampensis sp. nov.; H, A. carlae sp. nov.; I, A. mathewsae sp. nov.; J, A. brasileiro sp. nov.; K, A. nuno sp. nov.; A. viridari (Armstrong, 1949). C, E, F, G, I, J mirrored horizontally from the originals for better visual comparison.

FIGURE 66. Colour patterns of 11 western Atlantic species of the Alpheus armillatus H. Milne Edwards, 1837 complex and A. viridari (Armstrong, 1949): A, A. armillatus H. Milne Edwards, 1837; B, A. lancirostris Rankin, 1900; C, A. verrilli (Schmitt, 1924); D, A. angulosus McClure, 2002; E, A. amarillo sp. nov.; F, A. punctatus sp. nov.; G, A. tampensis sp. nov.; H, A. carlae sp. nov.; I, A. mathewsae sp. nov.; J, A. brasileiro sp. nov.; K, A. nuno sp. nov.; L. A. viridari (Armstrong, 1949). All photographs by the author

FIGURE 67. Frontal regions of nine western Atlantic species of the Alpheus armillatus H. Milne Edwards, 1837 complex, showing

FIGURE 69. Juvenile colour patterns of five western Atlantic species of the Alpheus armillatus H. Milne Edwards, 1837 complex [specimens not deposited]: A, B, A. armillatus H. Milne Edwards, 1837; C, A. amarillo sp. nov.; D, A. mathewsae sp. nov.; E, A. punctatus sp. nov.; F, A. angulosus McClure, 2002. All photographs by the author.

FIGURE 70. Preliminary distribution maps for 11 western Atlantic species of the Alpheus armillatus H. Milne Edwards, 1837 complex (A. armillatus H. Milne Edwards, 1837; A. lancirostris Rankin, 1900; A. verrilli (Schmitt, 1924); A. angulosus McClure, 2002; A. amarillo sp. nov.; A. punctatus sp. nov.; A. tampensis sp. nov.; A. carlae sp. nov.; A. mathewsae sp. nov.; A. brasileiro sp. nov.; A. nuno sp. nov.) and A. viridari (Armstrong, 1949); individual locations refer to material reported in this study, verified literature records and in situ colour photographs.