Alpheus armillatus H. Milne Edwards, 1837

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 : 8-18

Alpheus armillatus H. Milne Edwards, 1837

( Figures 1–8 , 16H , 64A , 65A , 66A , 68A , 69A, B )

Alpheus armillatus H. Milne Edwards 1837: 354 ; Coutière 1899: 24 et seq. (see Chace & Forest 1970 for all page numbers and figures); Coutière 1910: 485; (?) Zimmer 1913: 401 (part.?, not figs. K 1 -T 1); (?) Chace 1956: 147; Chace 1972: 62 (part.); Hernández Aguilera et al. 1996: 28; (?) McClure & Wicksten 1997: 480; Anker 2001: 69; (?) Tagliafico et al. 2005: 91; (?) Felder et al. 2009: 1056 (part.?).

Crangon armillatus — Schmitt 1924a: 76; (?) Schmitt 1924b: 65; (?) Darby 1934: 349; (?) Darby 1935: 151; (?) Schmitt 1935: 142; (?) Schmitt 1936: 368; (?) Darby 1938: 78; (?) Darby 1939: 61; (?) Hess 1941: 215; (?) Armstrong 1949: 10.

Alpheus cf. armillatus —(?) Román-Contreras & Martínez-Mayén 2010: 224.

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

Not Alpheus armillatus —(?) Zimmer 1913: 401 (part., figs. K 1 -T 1 = A. lancirostris ); Boone 1930: 168, pl. 60; Hazlett 1962: 82; Hazlett & Winn 1962: 26; Sterrer 1986: 325, text-pl. 105, pl. 9, fig. 5 (= A. lancirostris ).

Not Alpheus armillatus — Holthuis 1956: 173; Tabb & Manning 1961: 595; Williams 1965: 67, fig. 55; Forneris 1969: 86; Rouse 1970: 137; Hendrix 1971: 59, pls. 3, 4 (part.?); Chace 1972: 62 (part.); Coelho & Ramos 1972: 149; Corrêa 1972: 2; Fausto-Filho 1978: 66; Fausto-Filho 1979: 49; Christoffersen 1980a: 23, figs. 5, 6; Christoffersen 1980b: 39; Ramos- Porto 1980: 299; Gore et al. 1981: 490; Santos 1981: 342, figs. 17a-e, 18c-e; Williams 1984: 92, fig. 63 (part.); Sampaio Neto & Fausto-Filho 1984: 13; Freitas-Lins 1985: 9; Abele & Kim 1986: 198, 212-213, figs. d-f; Santos Calado 1996: 143; Christoffersen 1998: 356 (part., Brazilian records); Santos & Coelho 1998: 73, fig. 10 (adapted from Williams 1984); Sousa et al. 1998: 119; Coelho et al. 2006: 51; Mossolin et al. 2006: 48; Pavanelli et al. 2008: 124 (all = A. angulosus and/ or below-described new species).

Not Alpheus armillatus — Mathews 2007: 471 (= one of the below-described new species).

Not Alpheus armillatus — Wicksten 1983: 50; Hendrickx et al. 1983: 73; Wicksten & Hendrickx 1992: 4; Alvarez del Castillo et al. 1992: 5 (= one or several Pacific species of the A. armillatus complex).

Not Alpheus armillatus —Christoffersen & Ramos 1988: 63, 65 (= A. angulosus and one of the below-described new species).

Not Alpheus armillatus sensu stricto — Rodrigues et al. 2009: 336 (= one of the below-described new species).

Not Alpheus armillatus - Amaral et al. 2010: 246 (possibly = A. verrilli or other species).

Not Crangon armillatus — Hay & Shore 1918: 386, fig. 9, pl. 27, fig. 1 (at least part. = A. angulosus ).

Not Crangon armillatus — Darby 1934: 349; Darby 1935: 151; Darby 1938: 78; Darby 1939: 61 (at least part. = A. viridari ; see Armstrong 1949 and Williams 1984).

Not Alpheus armillatus or Crangon armillatus — Verrill 1922: 76, pl. 48, figs. 2-2n (= A. lancirostris and A. verrilli ).

Not Alpheus cf. armillatus — Wicksten 1993: 152 (= one or several Pacific species of the A. armillatus complex).

(?) Alpheus heterochaelis (not Say 1818)—(?) Rathbun 1901: 107 (part.?).

Not Alpheus heterochaelis (not Say, 1818) [assigned to A. armillatus by Christoffersen 1980a: 24]—(?) Smith 1871: 23; Kingsley 1878: 194 (part., specimens Bermuda most likely = A. lancirostris ); Rathbun 1900: 152 (part.); Luederwaldt 1919: 429 (part.)

Not Alpheus Edwardsii (not Audouin, 1826) [assigned to A. armillatus by Holthuis 1951: 84]— Pocock 1890: 518 (part.).

Crangon verrilli Schmitt, 1924a: 77 (part., paratype).

Type material. Aruba: neotype: male (cl 10.7), MNHN-IU-2010-4062, Baby Beach, exposed flat with coral rocks and rubble, in crevices of coral rocks, 1 m, leg. A. Anker, 06.12.2003 [fcn 03-027].

Additional material. Aruba: 2 males (cl 6.0, 6.9), 1 female (cl 5.2), OUMNH. ZC . 2011-06-024, same collection data as for the neotype [fcn 03-022] . Venezuela: 1 male (cl 9.7), 1 ov. female (cl 9.7), MNHN-IU-2010-4063, Morrocoy , degraded reef and rubble fields, in coral rocks, 0.5–1.5 m, leg. A. Anker, 09.12.2003 [fcn 03-023, male dissected] . Panama: 2 males (cl 10.0, 11.2), MNHN-IU-2010-4064, Bocas del Toro , Isla Carenero, near Bucaneer resort, sand flat with abundant coral rubble and seagrass, under rocks, 0.5–1 m, leg. A. Anker, 25.10.2005 [fcn 05- 178, 1 male dissected]; 1 male (cl 6.2), UP , Bocas del Toro , Isla Colón, Boca del Drago , under rubble, 0.5–1 m, leg. A. Anker, J.A. Vera Caripe, 11.11.2006 [fcn 06-559]; 1 ov. female (cl 6.5), UP , same collection data [fcn 06-560]; 1 male (cl 11.4), UP , same collection data [fcn 06-558]; 1 male (cl 11.6), OUMNH. ZC . 2011-06-025, Bocas del Toro , Isla Colón, STRI point, sand flat with abundant rubble and some seagrass, near mangroves, less than 1 m, leg. A. Anker, 17.11.2006 [fcn 06-596]; 1 male (cl 13.9), MNHN-IU-2010-4065, Bocas del Toro , Isla Bastimentos, off Salt Creek, degraded reef, under rocks and rubble, 2–3 m, leg. A. Anker, 31.03.2008 [fcn 08-019]; 1 ov. female (cl 10.5), MNHN-IU-2010-4066, Bocas del Toro , San Cristobal, Punta Coco, sans flat with seagrass and rubble, under coral rocks and rubble, 0.5–1 m, leg. A. Anker, 29.10.2005 [fcn 05-186]; 1 ov. female (cl 9.6), RMNH D54809 , Isla Grande, southern coast near Playa de la Punta, seagrass, rubble, under rubble and coral rocks, less than 1 m, leg. A. Anker, 11.06.2006 [fcn 06-428]; 1 male (cl 8.6), RMNH D54808 , Isla Grande, southern coast near Playa de la Punta, seagrass, rubble, under rubble and coral rocks, less than 1 m, leg. A. Anker, 04.09.2006 [fcn 06-431]; 1 ov. female (cl 9.6), RMNH D54807 , Isla Grande, southern coast near Playa de la Punta, 1 m, rubble, leg. A. Anker, 17.09.2005 [fcn 05-181] . Belize: 1 male (cl 7.3), OUMNH. ZC . 2009-01-0022, Carrie Bow Cay, off dock, 16°48.171’N 88°04.933’W, breaking of coral rubble, leg. S. De Grave, 19.02.2009 [fcn CBC-016]; 1 ov. female (cl

9.6), OUMNH. ZC. 2009-01-0028, same collection data [fcn CBC-027]; 2 ov. females (cl 12.5, 12.8), OUMNH. ZC. 2009-01-0024, Carrie Bow Cay, north-east side, under rocks, 0.5–1.0 m, leg. S. De Grave, 20.02.2009 [fcn CBC-075]; 1 male (cl 12.5), OUMNH. ZC. 2009-01-0026, Carrie Bow Cay, 16°48.141’N 88°04.905’W, intertidal flat with Porites , inside dead conch, leg. S. De Grave, 23.02.2009 [fcn CBC-146]. Mexico: 1 male (cl 11.7), OUMNH. ZC. 2012-03-0001, Quintana Roo, Isla Cozumel, 1 km north of Playa Corona, nearshore fossilised coral platform, in dead conch, 2–3 m, leg. A. Anker, J. Duarte-Gutiérrez, 08.07.2010 [fcn COZ 2- 014]; 1 male (cl 8.1), OUMNH. ZC. 2012-03-0002, Yucatan, Quintana Roo, Mahahual, shallow flat with seagrass and rubble, 1–1.5 m, leg. A. Anker, J. Duarte-Gutiérrez, 06.07.2010 [fcn MAH-005]; 1 ov. female (cl 11.7), OUMNH. ZC. 2011-06-026, southern Gulf of Mexico off Yucatan, Arrecife Alacranes, Anegada Channel, 22º 27' 36.7'' N, 89º 36´42.4'' W, under large boulders, 1–2 m, leg. N. Simões et al., 14.06.2008 [fcn ALA-108A = 08-245]; 1 ov. female (cl 12.1), MNHN-IU-2010-4067, same collection data [fcn ALA-108C = 08-248]; 1 ov. female (cl 11.2), RMNH D54806 , same collection data [fcn ALA-108B = 08-247]; 1 ov. female (cl 10.4), OUMNH. ZC. 2012- 03-0003, southern Gulf of Mexico off Yucatan, Arrecife Alacranes, 22°23'2.18"N, 89°37'55.60", under rocks/rubble, 0.5 m, leg. J. Duarte-Gutiérrez, 09.08.2009 [fcn JD-048]. St. Martin: 1 female (cl 9.6), FLMNH UF Arthropoda 30324, Réserve Naturelle de Saint-Martin, sta. 33, Plage du Galion, under rocks in sand, 0.5–1 m, leg. A. Anker et al., 14.04.2012 [fcn BSTM-0522]; 1 ov. female (cl 11.0), FLMNH UF Arthropoda 30325, Réserve Naturelle de Saint-Martin, sta. 39, off Rocher Créole, 3–5 m, in silt and sand under large rocks, leg. A. Anker, J.F. Maréchal, 18.04.2012 [fcn BSTM-1001]. British Virgin Islands: 1 male (cl 11.0), 1 ov. female (cl 10.8), LACM- MBPC 11726, Guana Island, White Bay, sta. 33 C, shallow ARMS rubble, leg. J.W. Martin et al., 10.07.2000; 1 male (cl 9.8), LACM-MBPC 11727, Guana Island, North Beach, south of dock, sta. BVI-01-067, 18.482°N 64.575°W, coral rubble, less than 1 m, leg. J.W. Martin, T. Zimmerman, T. Haney, K. Fitzhugh, 25.07.2001 [fcn Vd0725]; 1 female (cl not measured), LACM-MBPC 11728, Guana Island, Grand Ghut, ~ 18.48°N ~ 64.563°W, sta. BVI-00-075C, very open flat dominated by soft corals growing on a coraline pavement, widely scattered mediumsized eroded coral heads present, rubble and sand rare, ARMS-07 rubble basket (mostly pieces of old Acropora palmata ), 9 m, leg. T. Zimmerman, T. Haney, J.W. Martin, R. Ware, 20.07.2000 [fcn vc1283]. USA: 1 ov. female (cl 7.2), RMNH D16987, Florida, Soldier Key near Miami, coral reef with Halimeda , 0–1 m, leg. L.B. Holthuis, 07.09.1960. Barbados: female (cl 11.0), USNM 57938, “Barbados-Antigua Expedition from the University of Iowa”, in empty shell of Strombus , 29.05.1918 [paratype of Crangon verrilli Schmitt. 1924 , see below].

Diagnosis. Rostrum straight, exceeding half-length of first article of antennular peduncle; area posterior to rostral carina flattened, abruptly delimited from adjacent deep rostro-orbital furrows, forming somewhat between V and U-shaped post-rostral plate, its margins usually overhanging rostro-orbital furrows; post-rostral plate situated slightly above and gently sloping into narrow rostral carina. Antennule with stylocerite acute distally, not overreaching distal margin of first article; ventromesial carina of first article with large tooth bearing acute point on anterior margin; second article about twice as long as wide. Antenna with basicerite armed with strong distolateral tooth; scaphocerite with slightly concave lateral margin, strong distolateral tooth reaching well beyond relatively narrow blade; carpocerite exceeding scaphocerite blade but not distolateral tooth, at about the same level as end of antennular peduncles. Epistomial sclerite with sharp process. Mouthparts typical for genus; mandible with eight or nine teeth on distal margin of incisor process. Third maxilliped with ultimate article about as broad as penultimate article, tapering distally. Major cheliped edwardsii - type: palm laterally compressed, dorsal margin with transversal groove delimited by dorsal shoulder, and extending ventrally to both lateral and mesial faces, then extending posteriorly as longitudinal depression, ventral margin with transversal groove delimited by ventral shoulder and extending to lateral face; adhesive disks on pam and dactylus small. Male major cheliped with merus very stout, distodorsal margin ending bluntly, ventromesial margin smooth to slightly rugose, with row of minute spiniform setae distally and strong distomesial tooth; palm with dorsal shoulder rounded, sloping into adjacent transverse groove almost vertically, not overhanging groove; ventral shoulder broadly rounded, not projecting, smooth or with a few minute tubercles ventrally and/or laterally; fingers more than half-length of palm; pollex without oblique ridge mesially; dactylus plunger large, stout, proximal height about 0.7 length of distolateral margin, anterior angle about 90°. Female major cheliped somewhat smaller than male major cheliped, chela usually weaker and with more or less different proportions. Male minor cheliped with merus stout, distodorsal margin blunt, ventromesial margin usually with small spiniform setae, distomesial tooth small; chela moderately stout, palm with lengthheight ratio about 1.8; palm without longitudinal depressions, with deep sinus on ventral margin; fingers about as long as palm, simple, non-balaeniceps, with sharp cutting edges. Female minor cheliped generally similar to male minor cheliped, usually more slender; palm with faint ventral sinus. Second pereiopod slender, with first two carpal times 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 spiniform seta in both sexes, exceptionally unarmed. Sternum at the base of fourth pereiopods without conspicuously projecting, slender processes. First to fourth abdominal sternites in males each with median process between bases of pleopods, processes usually larger and sharper on first and second sternites, smaller and blunter on third and fourth sternites; first to fourth sternites in females each with strong, subacute, median process. First to fourth pleopods with ventrolateral margin of protopod bearing row of widely spaced, short spiniform setae in adult males, with dense row, often two or three rows, of longer, stronger spiniform setae in larger females; male second pleopod with appendix masculina slightly shorter than 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 at some distance 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; anal tubercles well developed ( Figs. 2–5 ). Gill-exopod formula typical for genus.

Variation. Alpheus armillatus shows some sexual dimorphism in the size and shape (proportions) of the major sloping rather abruptly compared to the lower and more gently sloping post-rostral plate of smaller individuals. The ischium of the fifth pereiopod is typically armed with a small spiniform seta; however, in one relatively small specimen from Belize, this seta was absent. The spiniform setae in the female first to fourth pleopods vary from very small and widely spaced in younger individuals to strong and elongate, forming dense double or triple rows in larger ovigerous females ( Fig. 2U ).

Size range. The present specimens range from 6.0 to 13.9 mm cl in males (neotype: 10.7 mm), and from 5.2 to 12.8 mm cl in females.

Colour pattern. Background opaque greysh-white; carapace with two broad, transverse, usually brown, sometimes grey-brown or bluish-grey bands; each abdominal somite with a broad transverse bands of brown, greybrown, bluish-grey or greenish-brown colour; post-rostral plate not particularly marked, anterior rostral carina dark brown; antennular and antennal flagella more or less intensely blue; chelipeds with ischium, merus and carpus marbled with brown or grey-brown-green; mesial face of major chela marbled with brown or grey and white or pale yellow, with numerous interconnecting white spots and patches, pale yellowish area usually present on proximal half; dactylus and pollex brown and white-spotted except for pale orange or pinkish distal areas; mesial face of minor chela brown with some white marbling and spotting, finger tips conspicuously orange; second to fifth pereiopods yellow; telson and uropods mostly brown or dark-grey distally, mostly white proximally, with large area of golden-yellow chromatophores laterally on uropodal exopod; colour of eggs variable from dark yellow to olivegreen ( Figs. 6–8 ); juveniles strikingly banded and with conspicuously blue antennal flagella ( Fig. 69A, B ).

Type locality. Neotype locality: Aruba [original typ locality: “Les Antilles” (H. Milne Edwards 1837)].

Distribution. Caribbean Sea: Guadeloupe, St. Martin, Barbados, Aruba, Venezuela (Morrocoy), Panama (Isla Grande, Bocas del Toro), Costa Rica (Cahuita), Belize (Carrie Bow Cay), Honduras (Utila), Mexico (Cozumel); southern Gulf of Mexico (Alacranes Reef off northern Yucatan); southern Florida: Dry Tortugas, Florida Keys, Soldier Key (off Miami) (present study; see also Coutière 1910); presently not known from the Guyanas, Brazil, northern Gulf of Mexico, and the Atlantic coast of the USA north of southern Florida (see map in Fig. 70 and remarks below).

[ OUMNH. ZC. 2009-01-0024, 2009-01-0026]; A, thoracic sternum, ventral view; B, same, ventrolateral view; E, abdominal sternum, ventrolateral view. Abbreviations: P1–P5, first to fifth pereiopods; PL1 – PL5 , first to fifth pleopod; S1–S6, first to sixth abdominal somites; U, uropod. See also Figure 16H [paratype of A. verrilli (Schmitt, 1924) ] .

Ecology. Shallow coral reefs and adjacent habitats at a depth range of 0.5–9 m; usually on mixed sand-rubble bottom types, especially on sand bottoms mixed with coral rubble, Thalassia or Halimeda , sometimes near mangroves; typically under rocks or in coral rubble crevices, also in dead conch shells; in male-female pairs.

closely related species by a unique combination of morphological characters (see Table 1 and below), as well as a unique and diagnostic colour pattern (type BA 2 in Mathews & Anker 2009, fig. 2H), which combines the dark transverse banding of the body with the conspicuously blue antennular and antennal flagella ( Figs. 6–8 ).

The presently known distribution range of A. armillatus includes the Caribbean Sea with the adjacent areas of southern tip of Florida and southern-most Gulf of Mexico ( Fig. 70 ). Despite numerous literature records, A. armillatus seems to be absent from Brazil, the northern Gulf of Mexico and the Atlantic coast of the USA north of Florida. As pointed out by McClure (1995), all previous records of A. armillatus north of southern Florida, as well as records from the northern Gulf of Mexico from the Everglades to Texas, most probably refer to A. angulosus (see below). Similarly, A. armillatus has not yet been found south of the Lesser Antilles, including the coast of the Guyanas and Brazil. The abundant records of A. armillatus from Brazil (e.g., Holthuis 1956; Coelho & Ramos 1972; Christoffersen 1998 and references therein; Coelho et al. 2006; Mossolin et al. 2006; Rodrigues et al. 2009) refer either to A. angulosus (which is present in Brazil, see below) or to one of the below-described new species, two of them being present in Brazil (see below). However, more sampling is needed in the scattered rocky reef habitats of the northern Brazilian states of Amapá and Pará, and especially in Atol das Rocas and Fernando do Noronha, to confirm the herein presumed absence of A. armillatus from Brazil.

In most females of A. armillatus , especially in larger ovigerous females, the ventrolateral margin of the protopods of first to fourth pleopods are furnished with a row or double row of stout spiniform setae ( Fig. 2U ). These spiniform setae are also present on male pleopods, where however they are shorter and more widely spaced ( Fig. 2O ). The prickly pleopodal protopods represent an additional diagnostic character of A. armillatus for most of the closely related species don’t have them (see below).

GenBank accession numbers. Aruba: FJ528564 (MyHC) , FJ528477 , FJ528472 (16S) , FJ528543 , FJ528538 ( COI) [MNHN-IU-2010-4062, neotype] ; Panama: FJ528565 (MyHC) , FJ528471 , FJ528468 (16S) , FJ528537 , FJ528534 ( COI) [? MNHN-IU-2010-4065] ; Venezuela: FJ528473 (16S) , FJ528539 ( COI) [MNHN-IU-2010- 4063] .

Figures

FIGURE 1. Alpheus armillatus H. Milne Edwards, 1837 species complex, early illustrations (1899–1924): A—A. armillatus, probably syntype, rostro-orbital region of carapace, lateral and dorsal views [from Coutière 1899, p. 97, figs. 66, 67, abbreviations: exc, extra-corneal areas of orbital hoods; r, rostrum; ror, rostro-orbital furrows]; B—type of A. lancirostris Rankin, 1900, frontal region, dorsal view [from Rankin (1900), pl. XVIII, fig. 5a]; C—specimen of A. armillatus sensu Zimmer (1913), frontal region, dorsal view, antenna, lateral view, and major chela, mesial view [from Zimmer 1913, p. 402, figs. K–M]; D—specimen of A. armillatus sensu Hay & Shore (1918), rostro-orbital region of carapace, dorsal view [from Hay & Shore 1918, p. 386, fig. 9]; E—specimen of A. armillatus sensu Verrill (1922), habitus, dorsal view [from Verrill 1922, fig. 6b]; F—type of A. verrilli (Schmitt, 1924), frontal region, dorsal and lateral views, and major cheliped, lateral (above) and ventromesial (below) views [from Schmitt 1924, pl. II, figs. 7–10].

FIGURE 2. Alpheus armillatus H. Milne Edwards, 1837: A–S, male from Bocas del Toro, Panama [MNHN-IU-2010-4064]; T, U, ovigerous female from Isla Grande, Panama [RMNH D54809]; A, frontal region of carapace and frontal appendages, dorsal view; B, same, lateral view; C, rostro-orbital region of carapace, dorsolateral view; D, tooth on mesioventral carina of first article of antennular peduncle, lateral view; E, antennal scaphocerite, dorsal view; F, epistome, ventral view; G, mandible, mesial view; H, third maxilliped, lateral view; I, second pereiopod, lateral view; J, third pereiopod, lateral view; K, same, distal propodus and dactylus, lateral view; L, fifth pereiopod, lateral view; M, first two abdominal sternites, lateral view; N, diagrammatic representation of first two abdominal ster-

FIGURE 3. Alpheus armillatus H. Milne Edwards, 1837: A–J, male from Bocas del Toro, Panama [MNHN-IU-2010-4064]; K, ovigerous female from Isla Grande, Panama [RMNH D54809]; A, major chela, mesial view; B, same, lateral view; C, same, detail of ventral shoulder; D, same, detail of dorsal shoulder; E, major cheliped, ischium, merus and carpus, mesial view; F, same, detail of ventromesial margin; G, major chela dactylus, lateral view; H, minor cheliped, mesial view; I, minor chela, mesial view; J, same, lateral view; K, major chela, lateral view. Scale bars as indicated.

FIGURE 4. Alpheus armillatus H. Milne Edwards, 1837: male from Morrocoy, Venezuela [MNHN-IU-2010-4063]: A, frontal region of carapace and frontal appendages, dorsal view; B, anterior carapace, lateral view; C, tooth on mesioventral carina of first article of antennular peduncle, lateral view; D, major chela with dactylus open, lateral view; E, major chela with dactylus closed, mesial view; F,

FIGURE 5. Alpheus armillatus H. Milne Edwards, 1837: ovigerous female (A, B) and male (C) from Carrie Bow Cay, Belize

FIGURE 6. Alpheus armillatus H. Milne Edwards, 1837: male from Bocas del Toro, Panama [MNHN-IU-2010-4064], dorsal (A) and lateral (B) views. Photographs by the author.

FIGURE 8. Alpheus armillatus H. Milne Edwards, 1837: A, male from Cozumel, Mexico [OUMNH.ZC. 2012-03-0001], dorsal view; B, C, male and female from Guadeloupe, French Antilles [aquarium specimens, not deposited]; D, in situ photograph of an ovigerous female, southern Florida [specimen not collected]. Photographs: A, by the author; B, C, by F. Fasquel; D, by G. Paulay.

FIGURE 16. Alpheus verrilli (Schmitt, 1924) [A–G] and (?) Alpheus armillatus H. Milne Edwards, 1837 [H]: A, B, ovigerous female from Carrie Bow Cay, Belize [OUMNH.ZC. 2009-01-0034]; C–E, male from Bocas del Too, Panama [OUMNH.ZC. 2011-06-028]; F, G, holotype, ovigerous female from Barbados [USNM 68786]; H, paratype of A. verrilli, probably A. armillatus, female from Barbados [USNM 57938]; A, thoracic sternum, ventral view; B, same, ventrolateral view; C, thoracic sternum, posterior part (third to fifth pereiopods), ventral view; D, same, ventrolateral view; E, abdominal sternum, ventrolateral view; F, pleopods, lateral view; G, thoracic

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

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.