Alpheus bouvieri A. Milne-Edwards, 1878

Anker, Arthur, Hurt, Carla & Knowlton, Nancy, 2009, Description of cryptic taxa within the Alpheus bouvieri A. Milne-Edwards, 1878 and A. hebes Kim and Abele, 1988 species complexes (Crustacea: Decapoda: Alpheidae), Zootaxa 2153, pp. 1-23 : 4-7

Alpheus bouvieri A. Milne-Edwards, 1878

Figs. 1 , 5 A, B

Alpheus Bouvieri Milne-Edwards 1878: 231 ; Coutière 1898: 131 (part., specimens from Congo); Coutière 1899 (part., for page numbers see Chace and Forest 1970).

Alpheus bouvieri— Rathbun 1900: 312 (part.); Balss 1916: 21; Sourie 1954: 253; Forest and Guinot 1958: 9; Ribeiro 1964: 5; Crosnier and Forest 1965: 606; Crosnier and Forest 1966: 273, fig. 22; Ribeiro 1968: 30, fig. 7; Uchakov 1970: 444; Fausto-Filho and Furtado 1970: 286; Chace 1972: 63; Fausto-Filho 1974: 5; Fausto-Filho 1975: 79; Fausto-Filho 1980: 113; Abele 1976: 273; Gore et al. 1978: 225; Christoffersen 1979: 303, figs. 2-5; Christoffersen 1980a: 37, figs. 7–10; Christoffersen 1980b: 39; Freitas-Lins 1985: 11; Abele and Kim 1986: 199, 214–215, figs. 1–d; Manning and Chace 1990: 14; Martínez-Iglesias et al. 1996: 33; Hernández-Aguilera et al. 1996: 30; Martínez-Iglesias et al. 1997: 404, fig. 6; Christoffersen 1998: 357; Santos and Coelho 1998: 74; McClure 2005: 133 (part., not fig. 8); Coelho et al. 2006: 1221; Almeida et al. 2006: 9.

Crangon (Alpheus) bouvieri — Monod 1933: 462, figs. 1A–C.

Alpheus leviusculus bouvieri— Banner and Banner 1982: 250 (part.).

Not Alpheus Bouvieri — Coutière 1898: 131 (part., specimens from Pacific coast of Panama = A. javieri n. sp.); Coutière 1905: 907 (part, other species).

Not Alpheus bouvieri — Holthuis 1951: 81; Holthuis 1952b: 43 (possibly = A. holthuisi Ribeiro, 1964 ; see Crosnier and Forest 1966); Monod 1927: 594 (= A. pontederiae Rochebrune, 1883 ; see Crosnier and Forest 1966); Chace 1962: 610 (possibly = A. longinquus Kim and Abele, 1988 ; see Kim and Abele 1988); Christoffersen 1979: 303 (part., specimens from Eastern Pacific); Kim and Abele 1988: 58, fig. 24; Wicksten and Hendrickx 1992: 4; Ricaurte et al. 1995: 81, 84; Vargas and Cortés 1999: 899; McClure and Wicksten 2000: 966, fig. 2; Wicksten and Hendrickx 2003: 63; McClure 2005: 134 (part., fig. 8 only) (= A. javieri n. sp.); Hickman and Zimmerman 2000: 35 (= Alpheus sp.).

Not Alpheus leviusculus bouvieri — Christoffersen and Ramos 1988: 63 (= A. javieri n. sp.)

Not Alpheus bouvieri var. chilensis (not sensu Coutière 1902)— Schmitt 1924a: 162 (= A. javieri , n. sp.).

Alpheus Edwardsii (not sensu Audouin 1826) — Dana 1852: 542; Dana 1855: 11, pl. 34, fig. 2a–f; Bate 1888: 542, pl. 97, fig. 1; Pocock 1890: 518 (part.) (see Crosnier and Forest 1966).

Alpheus heterochaelis (not sensu Say 1818) — Kingsley 1878: 194 (part.); Rathbun 1900: 152 (part.); Luederwaldt 1919: 429 (part.) (see Christoffersen 1979).

Alpheus nuttingi var.? (not sensu Schmitt 1924b)— Schmitt 1936: 368.

Alpheus sp. — Santos 1974: 7 et seq., figs. 1–16, 18g; Santos 1981: 319 et seq., figs. 1–16, 18g.

Material examined.— São Tomé. 1 male (CL 4.8), 2 ovig. females (CL 5.2, 6.4), USNM 1123843, ST 1, Boca do Inferno, rocky intertidal, under consolidated volcanic rocks and coralline algae, coll. A. Anker and N. Knowlton, 29 Jan 2006 [fcn 06-008 (male), 06-007, 06-009 (females)]; 1 female (CL 4.0), USNM 1123845, ST 2, Praia Lagarto, near São Tomé town hospital, intertidal, sand with Montastraea , zoanthids and rocks embedded in sand, coll. N. Knowlton and A. Anker, 30 Jan 2006 [fcn 06-026]; 1 male (CL 3.9), 1 ovig. female (CL 4.6), USNM 1123846, ST 3, 200 m west of Lagoa Azul, rocky intertidal with coralline algae, from holes and under rocks, coll. A. Anker and N. Knowlton, 31 Jan 2006 [fcn 06-073 (male), 06-072 (female)]; 1 male (CL 4.2), 1 female (CL 5.5), 1 ovig. female (CL 4.5), MNHN-Na 17182, ST 2A, Praia Lagarto, near São Tomé town hospital, intertidal, sand with Montastraea , zoanthids and rocks embedded in sand, coll. N. Knowlton and A. Anker, 2 Feb 2006 [fcn 06-132 (male), 06-131, 06-133 (females)]; 1 ovig. female (CL 5.7), USNM 1123847, ST 11, Kia dive site off Ilha das Cabras, depth about 10 m, under rocks, coll. N. Knowlton, 8 Feb 2006 [fcn 06-189].

Cape Verde. 1 male (CL 5.3), 1 female (CL 7.9), USNM 1123848, São Vicente, Baia das Gatas, coll. N. Knowlton et al., 28 Mar 1998; 7 males (CL 4.2–5.4), 8 females, including 7 ovig. females (CL 3.9–6.9), USNM 1123849, Santiago, Praia, beach near Club Nautics, coll. N. Knowlton et al., 5 Apr 1998; 1 ovig. female (CL not measured), MNHN-Na 13674, Terrafal, in tide pool, coll. P. Wirtz, Feb 2000.

Panama (Caribbean coast). 3 males (CL 5.4, CL 6.1, 7.4), 2 ovig. females (CL 6.1, 8.2), USNM 1123850, Bocas del Toro, Isla Colón, Bocas del Drago, shallow subtidal (less than 0.5 m), from crevices in rocks and rubble, coll. C. Hurt, 16 Sep 2006 [06-474, 06-475, 06-517 (males), fcn 06-473, 06-476 (females)]; 1 male (CL 6.7), 1 ovig. female (CL 7.3), USNM 1123851, Bocas del Toro, Isla Colón, between Big Creek and Playa Bluff, from crevices in rocks and fossilized coral platform, coll. A. Anker, A. Baeza and J.A. Vera-Caripe, 19 Nov 2006 [fcn 06-612 (male, specimen dissected), 06-611 (female)]; 1 male (CL 8.2), 2 ovig. females (CL 7.1, 8.7), UP, same collection data as for previous specimens [fcn 06-616 (male), 06-614, 06-615 (females)].

Mexico (Gulf of Mexico). 1 male (CL not measured), FLMNH UF Arthropoda 16557, Los Alacranes Reef off Yucatan Peninsula, under rocks, coll. N. Simões, 2008 [fcn ALA 068 = 08-253].

Brazil. 1 male (CL 6.1), 9 females (CL 4.6–9.6), 3 juveniles (CL not measured), USNM 1123852, Pernambuco, mouth of Rio Formoso, Carneiro reef, Tomandaré, under rocks, coll. N. Knowlton et al., 17 Sep 1998; 1 ovig. female (CL 10.1), USNM 1123853, Rio de Janeiro, Ilha Grande, coll. N. Knowlton et al., 26 Sep 1998; 1 male (CL 7.4), USNM 1123854, Rio de Janeiro, Cabo Frio, coll. N. Knowlton et al., 18 Sep 1998; 1 ovig. female (CL 5.1), MNRJ 20843, LT 719, Atol das Rocas, Canal da Barretão, coll. P.S. Young, P.C. Paiva and A.A. Aguiar, 10 Oct 2000 [specimen dissected]; 1 ovig. female (CL 7.1), MNRJ 20844, Atol das Rocas, in front of Ilha do Farol, under rocks, coll. P.S. Young, P.C. Paiva and A.A. Aguiar, 14 Oct 2000.

Diagnosis. Species of Alpheus leviusculus group (Clade L). Frontal margin of carapace with distinct, short, narrow, distally acute rostrum; rostral carina sharply delimited, extending to level of eyes, gradually flattening; orbital hoods unarmed, rounded, swollen, gently sloping into shallow orbitorostral grooves. Antennule with second peduncular segment less than twice as long as wide; stylocerite with acute tip, reaching distal margin of first segment. Antenna with basicerite armed with sharp distoventral tooth; scaphocerite with strong distolateral tooth extending well beyond broad blade, reaching end of antennular peduncle; carpocerite overreaching both scaphocerite and end of antennular peduncle. Third maxilliped not particularly broadened, unarmed. Major cheliped with merus lacking sharp distomesial tooth; chela somewhat compressed, dorsal groove broad, deep, extending mesially as contrasting rounded depression and laterally as well defined, posteriorly extending groove; dorsal shoulder rounded, not overhanging; ventral groove broad and deep, adjacent shoulder rounded, slightly protruding; adhesive disks small; fingers about half-length of palm; plunger of dactylus well developed. Minor cheliped sexually dimorphic; palm without grooves or notches in both sexes; fingers somewhat broadened and balaeniceps in males, subconical and non-balaeniceps in females. Second pereiopod with first carpal segment slightly more than twice as long as second. Third pereiopod with unarmed ischium, merus and carpus; propodus with row of stout spines, most paired; dactylus stout, simple, conical. Male second pleopod with typical appendices masculina and interna. Uropod with protopod laterally projecting as acute tooth; exopod with distolateral spine bordered laterally by sharp distolateral tooth; transverse suture sinuous, consisting of two rounded lobes and subtriangular tooth adjacent to distolateral spine; both exopod and endopod furnished with row of spinules distally. Telson broad, tapering distally, with two pairs of dorsal spines; posterior margin slightly convex, with two pairs of spines at each posterolateral angle.

Description. See Crosnier and Forest (1966) for description and figures of eastern Atlantic specimens, and Christoffersen (1979) for description and figures of western Atlantic (Brazilian) specimens. Complementary figures are provided in Fig. 1 .

Size. Eastern Atlantic: largest male from Cape Verde at CL 5.4 mm and TL 13.7 mm; largest female from Cape Verde at CL 7.9 mm and TL 20.1 mm (Crosnier and Forest’s (1966) specimens were comparable, with a TL of 8–22 mm); western Atlantic: largest male from Panama at CL 8.2 mm and TL 23.9 mm; largest female from Brazil at CL 10.1 mm and TL 26.6 mm; egg size: about 0.6 x 0.5 mm (average from 3 eggs from largest ovigerous female).

Color pattern. Opaque-whitish with broad transverse bands varying from brown-green to dark-brown (combination of gray-blue and red chromatophores), two on carapace and one along posterior half of each pleomere; uropods and telson white proximally, brown distally; walking legs whitish with red chromatophores; minor chela pale brown to green-brown; major chela olive-green to pale brown mesially, with more or less marked whitish patch ventroproximally; dorsal and ventral grooves paler brown to almost white; distal half of pollex and dactylus whitish or pink; rostrum and rostral carina region brown or browngreen; antennular peduncles brown; scaphocerite of antenna with brown lateral margin and distolateral tooth; antennal and antennular flagella pale yellow; eggs or ovaries in females yolk-yellow ( Fig. 5 A, B).

Type locality. Cape Verde ( Milne-Edwards 1878); type specimens in MNHN.

Distribution. Amphi-Atlantic. Eastern Atlantic: Cape Verde; Senegal; Guinea; Ghana; São Tomé; Cameroon; Gabon; Congo (e.g., Balss 1916; Sourie 1954; Crosnier and Forest 1966; Ribeiro 1968; present study); Central Atlantic: Ascension Island ( Manning and Chace 1990); Western Atlantic: Florida; Bermuda; Caribbean ( Cuba, Panama, Antigua, Guadeloupe, Grenada, Grenadines, Tobago); Gulf of Mexico; Brazil (Ceará to Rio Grande do Sul, including Atol das Rocas and Fernando do Noronha) (e.g., Chace 1972; Abele 1976; Gore et al. 1978; Christoffersen 1979, 1998; McClure 2005; present study).

Ecology. Intertidal or shallow subtidal, probably to a depth of about 5 m; under rocks or in rock and rubble crevices, on sand and mixed sand-mud substrates; sometimes in crevices of sabellariid reefs ( Gore et al. 1978); usually encountered in male-female pairs.

Remarks. Alpheus bouvieri is most closely related to A. javieri n. sp. (see below). However, the only species, with which it can be confused in the Atlantic, e.g., in Brazil or in West Africa, is A. agilis n. sp. (see below).

GenBank access number. FJ013876 ( Panama, fcn 06-502, 206572334 5'), FJ013875 ( Panama, fcn 06- 495, 206572332 5'), FJ013874 ( Panama, fcn 06-493, 206572330 5'), FJ013873 ( São Tomé, fcn 06-008, 206572328 5'), FJ013873 ( São Tomé, fcn 06-008, 206572328 5').

Figures

FIGURE 1. Alpheus bouvieri A. Milne-Edwards, 1878; male (A – D, F – H) and female (E) from Bocas del Toro, Panama (USNM 1123851): A, frontal region, dorsal view; B, major chela, mesial view; C, same, lateral view; D, male minor chela, mesial view; E, female minor chela, mesial view; F, second pereiopod, lateral view; G, third pereiopod, lateral view; H, telson, dorsal view. Scale bars: 1 mm.

FIGURE 5. Color patterns of Alpheus bouvieri A. Milne-Edwards, 1878 [western Atlantic] (A); Alpheus bouvieri [eastern Atlantic] (B); Alpheus javieri n. sp. (C, D); Alpheus hebes Kim and Abele, 1988 (E); and Alpheus agilis n. sp. [eastern Atlantic] (F): A, male from Bocas del Toro, Panama; B, female from São Tomé; C, male from Punta Morales, Costa Rica; D, female from Las Perlas, Panama; E, male from Las Perlas, Panama; F, male from São Tomé.