Alpheus carlae, 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 : 61-67

Alpheus carlae sp. nov.

( Figures 41–48 , 64H , 65H , 66H , 67G , 68B )

Alpheus armillatus (not H. Milne Edwards, 1837)—(?) Holthuis 1956: 173; Hendrix 1971: 59, 60 (part., banded colour pattern); Christoffersen & Ramos 1988: 63, 65 (part., banded-spotted type); Mossolin et al. 2006: 48; Pavanelli et al. 2008:

124. Alpheus armillatus sensu stricto (not H. Milne Edwards, 1837) — Rodrigues et al. 2009: 336. Alpheus cf. armillatus F— Mathews & Anker 2009: 277. Alpheus sp. —Osório e Castro 1964: 1 (part.), photos 1, 2, figs. 1–3.

Type material. USA: holotype: 1 male (cl 9.4), MNHN-IU-2010-4089, Florida Keys, Bahia Honda , bay side, near mangroves, 0.5 m, fine sand with rocks, algae, detritus etc., under rocks, leg. A. Anker, C. Hurt, A.L. Rhyne, 25.09.2007 [fcn 07-297, dissected] ; paratypes: 1 male (cl 9.4), OUMNH. ZC. 2011-06-034, same collection data as for the holotype [fcn 07-298]; 1 female (cl 10.9), MNHN-IU-2010-4090, Florida Keys, Bahia Honda , ocean side, north side of channel, 0 m (low tide), under rocks, leg. A. Anker, C. Hurt, A.L. Rhyne, 25.09.2007 [fcn 07-286]; 1 female (cl 9.8), MNHN-IU-2010-4091, Florida Keys, Bahia Honda, ocean side, north side of channel, 0 m (low tide), under rocks, leg. A. Anker, C. Hurt, A.L. Rhyne, 25.09.2007 [fcn 07-288]; 1 ov. female (cl 11.5), RMNH D54821 , Florida, Key Biscayne, shallow flat near mangroves, in muddy sand under rocks, 0 m (low tide), leg. A. Anker et al., 07.09.2008 [fcn 08-270] .

Additional material. USA: 1 male (cl 8.5), RMNH D54822, Florida, south of Fort Pierce , shallow subtidal flat with rocks, 3 m, under rocks, leg. J.A. Baeza, 22.06.2007 [fcn 07-237]. Belize: 1 male (cl 9.7), OUMNH. ZC. 2009-01-0031, Carrie Bow Cay, Twin Cays , 16°50.107’N 88°06.042’W, near mangrove, 0.5 m, from burrows, yabby pump, leg. S. De Grave, D.L. Felder, 23.02.2009 [fcn CBC-163]. Panama: 1 male (cl 8.5), 1 ov. female (cl 8.7), OUMNH. ZC. 2005-10-0098, Bocas del Toro, off STRI station pier, 09°14.983’N 082°08.239’W, 1 m, leg. T. Haney, 03.08.2005 . Venezuela: 1 ov. female (cl 7.2), OUMNH. ZC. 2011-06-035, Isla Margarita, Guamache , shallow sand-mudflat near mangroves, suction pump, <0.5 m, leg. A. Anker, J.A. Vera Caripe, 13.12.2003 [fcn 03- 026, dissected]. St. Martin: 1 ov. female (cl 8.6), FLMNH UF Arthropoda 30331, Réserve Naturelle de Saint-Martin, sta. 37, south side of Baie de l'Embouchure , near drainage of Étang aux Poissons , seagrass with mangroves, leg. Z. Whitener, A. Anker, 16.04.2012 [fcn BSTM-0737]. French Guyana : 1 male (cl 13.2), 2 ov. females (cl 12.0, 14.1), MNHN-Na 14202, locality not specified, Mission F. Geay, N1722, 1900; 2 males, 1 female (cl not measured), RMNH D14329 , Îles du Salut , 01.1957, leg. J. Durand (identification tentative, specimens without major chelipeds and most pereiopods). Brazil : 1 male (cl 11.0), OUMNH. ZC. 2011-06-036, Ceará, Praia de dois Coquieros , rocky intertidal, under large rocks at low tide, leg. A. Anker, 24.05.2011 [fcn 11-011]; 1 ov. female (cl 12.6), MZUSP 25235 , same collection data [fcn 11-012] ; 2 males (cl 5.5, 6.7), 1 female (8.6), MZUESC 1446 , Pernambuco, Paulista, Rio Timbó , 07°51’38.3”S 34°50’21.4”W, on mud under rocks, leg. A.O. de Almeida, R. J.C. Paiva, 09.04.2008 ; 1 male (cl 10.6), 2 ov. females (cl 8.4, 11.2), RMNH D10978, São Paulo, Cananeia (about 200 km south of Santos ), mangroves, leg. S. Gerlach, 06.1954 ; 1 male (cl 8.3), 1 ov. female (cl 9.8), MZUSP 25339 , Alagoas, Barra de Camaragibe, Rio Camaragibe estuary, hand collecting at night, leg. M. Tavares, J.B. Mendonça, 28.10.2011 ; 1 male (cl 12.5), MZUSP 5329 , Rio de Janeiro, Ilha Grande, Praia do Furado , leg. G.S. de Melo, 22.07.1966 ;> 20 specimens (males and females, cl not measured), MZUSP 13720 , São Paulo, Ilha Bela, Praia Engenho d'Água , leg. E.C. Mossolin, 01-04.2001 ;> 20 specimens (males and females, cl not measured), MZUSP 13719 , São Paulo, São Sebastião, Praia do São Francisco , leg. E.C. Mossolin, 01-04.2001 ; 2 males (cl 11.5, 12.7), 1 ov. female (cl 10.6), MZUSP 18008 , São Paulo, São Sebastião, Praia do São Francisco , 23°44'53.6''S 045°24'33.6''W, leg. E.C. Mossolin, 19.11.2006 .

Diagnosis. Rostrum straight, usually overreaching half-length of first article of antennular peduncle; area posterior to rostral carina flattened, markedly but not abruptly delimited from adjacent, shallow to moderately deep rostro-orbital furrows, forming V-shaped post-rostral plate continuous with narrow, dorsally blunt rostral carina, margins of post-rostral plate not overhanging rostro-orbital furrows; post-rostral plate situated at about the same level as rostral carina or slightly above it, in latter case gently sloping. Antennule with stylocerite acute distally, reaching or slightly overreaching distal margin of first article; ventromesial carina of first article with tooth bearing small subacute point above concave anterior margin; second article about twice as long as wide. Antenna with distolateral tooth, reaching slightly past end of antennular peduncles. Third maxilliped with ultimate article as broad as penultimate, tapering distally. Major cheliped edwardsii - type (see under A. armillatus ). Male major cheliped with merus stout, distodorsal margin ending bluntly, ventromesial margin smooth, without stout spiniform setae, with strong, acute distomesial tooth; palm with dorsal shoulder rounded, sloping obliquely into adjacent transverse groove, not overhanging groove; ventral shoulder broadly rounded, not projecting, smooth laterally; fingers more than half-length of palm; pollex without ridge mesially; dactylus plunger large, stout, proximal height about 0.7 length of distolateral margin, anterior angle about 90°. Female major cheliped usually smaller than male major cheliped, with chela smaller and with more or less different proportions. Male minor cheliped with merus moderately stout, distodorsal margin blunt, ventromesial margin without spiniform setae, with sharp distomesial tooth; chela relatively slender, palm with length-height ratio about 2.0; palm without longitudinal depressions, with very slight sinus on ventral margin, not distinctly delimited by ventral shoulder; fingers about as long as palm, slen- der, simple, non-balaeniceps, with sharp cutting edges. Female minor cheliped generally similar to male minor cheliped, more slender; palm without or with very slight ventral sinus. Second pereiopod slender, with first two carpal articles longest, first about 1.3 length of second. Third and fourth pereiopods similar, rather slender; ischium with stout spiniform seta ventrolaterally; merus about five times as long as wide; propodus with stout spiniform setae, sometimes inserted in pairs, along ventral margin, incuding one pair adjacent to dactylus; dactylus about 0.4 length of propodus, simple, conical. Fifth pereiopod much more slender than third and fourth; ischium armed with small spiniform seta, latter sometimes present only as a trace or lacking, in both sexes. Sternum at the base of fourth pereiopods without conspicuously projecting, slender, subacute processes. First and second abdominal sternites each with strong, blunt or subacute median process in males, unarmed in females (sometimes with very small process on second sternite). First to fourth pleopods without spiniform setae on ventrolateral margin of protopod in both sexes; male second pleopod with appendix masculina somewhat shorter than appendix interna, densely covered with stiff setae apically and mesially. 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 stouter spiniform setae, mesial much longer than lateral ( Figs. 41–43 ).

Variation. In addition to sexual dimorphism affecting the shape and proportions of the major and minor chelae, there is some variation in the shape of the post-rostral plate, which is more elevated and sloping in some larger specimens ( Fig. 43L, M ). The presence of a small spiniform seta on the ischium of the fifth pereiopod appears to be extremely variable: it is usually present in younger individuals and absent or reduced to a shallow pit in larger individuals (see also under A. amarillo sp. nov.). In the female specimens from Ceará, a small blunt median process may be present on the second abdominal sternite. In two males from São Paulo ( MZUSP 18008 ), the rostral carina is aberrant by either being strongly arched or absent .

Size range. The type specimens from southern Florida range from 9.4 to 11.5 mm cl (male holotype: 9.4 mm, females: 9.8–11.5 mm); Brazilian specimens range from 5.5 to 12.7 mm; the single Venezuelan specimen is an ovigerous female at cl 7.2 mm cl; the largest specimens examined are a male and an ovigerous female from French Guyana, at cl 13.2 mm and 14.1 mm, respectively.

Colour pattern. Body with red-brown and bluish chromatophores forming broad transverse bands of pale-

brown, red-brown, blue-green or olive-brown colour, on each abdominal somites; many chromatophores interconnecting or forming irregular or branched chains, latter sometimes extending into large white areas between the bands; colourless areas between red-brown bands broadening posteriorly, widely V-shaped on fifth somite, and with yellow chromatophores (much more conspicuous on dark background); second somite with conspicuous black spot laterally on pleuron; third somite with more or less distinct small black spot dorsally; fourth somite with conspicuous black spot dorsolaterally on pleuron; post-rostral plate with brown-red chromatophores; antennular and antennal flagella pale brownish or olive-yellow; chelipeds with ischium, merus and carpus whitish, distally with patches of red-brown and yellow chromatophores; mesial face of major chela red-brown, orange-brown, or olivebrown with numerous pale or whitish patches and spots, including some on dactylus, their size, shape and number extremely variable; distodorsal margin of pollex and dorsal margin of dactylus sometimes brown or blackish; distal portion of dactylus and pollex pinkish; mesial face of minor chela with general pattern similar to that of major chela; second to fifth pereiopods reddish; telson brownish with pale area; uropodal endopod dark purplish-brown; exopod pale yellow with some red-brown chromatophores, more deep-blue distally; general colour and width of abdominal bands and size of black spots on the abdomen rather variable; eggs green or olive-greenish ( Figs. 44–48 ). Compare with “banded colour pattern” in Hendrix (1971, p. 64).

Etymology. This species is named after Dr. Carla Hurt (University of Miami), whose molecular expertise contributed to the taxonomic revision of several transisthmian species complexes of Alpheus .

Type locality. Southern Florida: Florida Keys and Key Biscayne .

Distribution. Southern Florida: Florida Keys, Key Biscayne, Fort Pierce; Caribbean Sea: Venezuela ( Isla Margarita), Panama (Bocas del Toro), Jamaica (Runaway Bay), Puerto Rico (Ceiba), Belize (Carrie Bow Cay); French Guyana; Brazil: Ceará, Pernambuco, Paraíba, Bahia, Rio de Janeiro, São Paulo (see map in Fig. 70 and remarks below).

Ecology. Various intertidal and shallow subtidal habitats; known depth range 0–3 m, possibly deeper; in Florida and Venezuela mostly in shallow protected bays and lagoons, on fine sand and silt, close to mangroves or seagrass beds; in Brazil usually in sandier parts of rocky intertidal shores, in large tide pools or on shallow sand-rock-rubble flats behind rocky reefs; usually under rocks and coral rubble, or in burrows among mangrove roots; in male-female pairs.

Remarks. Alpheus carlae sp. nov. forms a small clade (Clade 2e) within the larger Clade 2 of Mathews & Anker (2009, fig. 4), being most closely related to A. angulosus , from which it is distinguishable morphologically by the first two abdominal sternites each armed with a strong median process in males (vs. only the first sternite with a small process in A. angulosus ). The presence of a spiniform seta on the ischium of the fifth pereiopod seems to be a variable feature in A. carlae sp. nov. (see above); nonetheless, the ischium is always unarmed in A. angulosus . Additional characters, as well as features distinguishing A. carlae sp. nov. from all the other western Atlantic species of the A. armillatus , are summarised in Table 1.

When alive, A. carlae sp. nov. can be easily distinguished from A. angulosus by its diagnostic colour pattern (type BA4 and BA 5 in Mathews & Anker 2009, fig. 2F, N), which, although remarkably variable ( Figs. 44–48 ), is unique in the combination of the transverse banding of the abdomen, the major and minor chelae with whitish spots or blotches, and the second, third and fourth abdominal somites each with a pair of small black spots. With this colour pattern, A. carlae sp. nov. can be immediately separated from all other species of the A. armillatus complex currently known from the western Atlantic.

Many previous records of A. armillatus from Florida may actually refer to A. carlae sp. nov. since A. armillatus appears to be rather uncommon in this area (see above). For instance, Hendrix’s (1971) specimens of A. armillatus with a “banded colour pattern” are most probably A. carlae sp. nov. Similarly, many Brazilian records of A. armillatus may refer to A. carlae sp. nov. Christoffersen & Ramos (1988) commented on two different colour morphs of A. armillatus in Brazil, a uniform colour morph (= A. angulosus , see above), and a banded-spotted colour morph, with greenish or brownish colours concentrated into transversal bands of variable width on the posterior portion of each abdominal somite, and with a pair of conspicuous black spots on the pleura of the second to fourth somites, which closely matches the colour pattern of A. carlae sp. nov. ( Fig. 47D, E , 48 ). The same colour pattern was described for the Paraíba material of “ Alpheus armillatus sensu stricto ” reported by Rodrigues et al. (2009), suggesting that this material is also A. carlae sp. nov. The São Paulo material of “ A. armillatus ” studied by Mossolin et al. (2006) and Pavanelli et al. (2008) is A. carlae sp. nov., based on the colour photographs provided by E. C. Mossolin. Part of the material from Rio de Janeiro reported as Alpheus sp. in the interesting study of the chelipeds by Osório e Castro (1964) is also A. carlae sp. nov., based on two photographs showing the entire shrimp

GenBank accession numbers. Florida: FJ528557 , FJ528558 (MyHC), DQ682869 , FJ528457 (16S), FJ528512 , FJ528515 ( COI) [not deposited]; Puerto Rico: FJ528455 (16S), FJ528511 ( COI) [not deposited]; Vene-

Figures

FIGURE 41. Alpheus carlae sp. nov.: holotype, male from Florida Keys [MNHN-IU-2010-4089]; 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, second pereiopod, lateral view;

FIGURE 42. Alpheus carlae sp. nov.: holotype, male from Bahia Honda, Florida Keys [MNHN-IU-2010-4089]; A, major chela, mesial view; B, same, lateral view; C, same, detail of dorsal shoulder, lateral view; D, major chela dactylus, lateral view; E, minor che-

FIGURE 43. Alpheus carlae sp. nov.: A–E, non-type, ovigerous female from Isla Margarita, Venezuela [OUMNH.ZC. 2011-06-035]; F, G, paratype, ovigerous female from Key Biscane, Florida [RMNH D54821]; H–J, non-type, female from Bahia, Brazil [MZUESC 1446]; K, L, non-type, female from Ceará, Brazil [MZUSP 25235]; M–O, non-type, male from Ceará [OUMNH.ZC. 2011-06-036]; A, frontal region of carapace and frontal appendages, dorsal view; B, rostro-orbital region of carapace, lateral view; C, second pereiopod, lateral view; D, third pereiopod, lateral view; E, fifth pereiopod, lateral view; F, rostro-orbital region of carapace, dorsal view; G, same, lateral view; H, frontal region of carapace and frontal appendages, dorsal view; I, major chela, lateral view; J, diagrammatic representation of first two abdominal sternites, ventroposterior view; K, rostro-orbital region of carapace, dorsal view; L, same, lateral view; M, same, lateral view (different specimen); N, tooth on mesioventral carina of first article of antennular peduncle, lateral view; O, diagrammatic representation of first two abdominal sternites, ventroposterior view. Scale bars as indicated, figs. J, N, O drawn without scale.

FIGURE 44. Alpheus carlae sp. nov.: A, paratype, male from Bahia Honda, Florida Keys [OUMNH.ZC. 2011-06-034], dorsal view; B, C, non-type, male from Fort Pierce, Florida [RMNH D54822], dorsal (B) and lateral (C) views. Photographs: A, by the author; B, C, by J.A. Baeza.

FIGURE 45. Alpheus carlae sp. nov.: holotype, male from Bahia Honda, Florida Keys [MNHN-IU-2010-4089], dorsal (A) and lateral (B) views on white background. Photographs by the author.

FIGURE 46. Alpheus carlae sp. nov.: paratype female from Bahia Honda, Florida Keys [MNHN-IU-2010-4090], dorsal (A) and lateral (B) views on white background. Photographs by the author.

FIGURE 47. Alpheus carlae sp. nov.: A, male from the Caribbean coast of Panama [specimen not deposited], dorsolateral view; B, C,

FIGURE 48. Alpheus carlae sp. nov.: A, B, non-type, male from Ceará, Brazil [OUMNH.ZC. 2011-06-036], dorsal (A) and lateral (B) views; C, D, ovigerous female from Ceará [MZUSP 25235], dorsal (C) and lateral (D) views. Photographs by the author.

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 68. Colour of the plunger of the major chela dactylus in five species of the Alpheus armillatus H. Milne Edwards, 1837 com-

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.