Leptalpheus forceps Williams, 1965

Scioli, Justin A., Robles, Rafael & Felder, Darryl L., 2024, New species and records of the symbiotic shrimp genus Leptalpheus Williams, 1965, with notes on Fenneralpheus Felder & Manning, 1986, and preliminary molecular analysis of phylogenetic relationships (Crustacea: Decapoda: Alpheidae), Zootaxa 5466 (1), pp. 1-72 : 47-49

Leptalpheus forceps Williams, 1965

( Figs. 21i–k , 25h )

Leptalpheus forceps Williams, 1965: 194 , figs. 1–2.— Dawson, 1967: 224.— Saloman, 1971: 67.— Chace, 1972: 77.— Williams, 1984: 101, fig. 69.—Abele & Kim, 1986: 194, 228–229, figs. a, b.—Felder & Rodriguez, 1993: 366.— Holthuis, 1993: 203, fig. 197.— Álvarez et al., 1999: table 1.— Hermoso-Salazar, 2001: 1278.— Felder et al., 2003: table 3.— Almeida et al., 2012: figs. 5E–G.— Scioli et al., 2020: 629.

Not Leptalpheus forceps .— Pachelle et al., 2016: 12 , fig. 6C.— Rosário et al., 2020: 63 , figs. 2–6 [= L. degravei n. sp.].

Not Leptalpheus aff. forceps .— Almeida et al., 2012: 15 , figs. 5A–D [= L. degravei n. sp.]. Not Leptalpheus cf. forceps . — Anker, 2008: 788 , figs. 4, 5, 6A, B.— Anker, 2011: 6 , figs. 3A, B.— De Grave & Anker, 2017: 12.— Chow et al., 2021: fig. 3 [= L. degravei n. sp.].

? Leptalpheus forceps .— Christoffersen et al., 1998: 361.—Riul et al., 2007: 7.

Material examined. USA: 1 female (cl 7.4 mm), ULLZ 4250 [ USNM 1540316], North Carolina, Wilmington, coll. C.D. Schubart & J.A. Cuesta, 23.08.2000; 2 females (4.5, 5.5 mm), ULLZ 13541 [ USNM 1547278], Florida, Tampa Bay, Gandy Causeway, sand flat, coll. D.L. Felder et al., 27.10.2011; 1 male (cl 6.9 mm), 1 female (cl 6.3 mm), ULLZ 4492 [ USNM 1540390], Florida, Tampa Bay, Alafia River mouth, coll. D.L. Felder et al., 09.05.2001; 1 non-ovigerous female (cl 7.9 mm), 1 ovigerous female (cl 8.6 mm), ULLZ 4494 [ USNM 1540392], Florida, Tampa Bay, boat launch, shoreline, coll. D.L. Felder et al., 09.05.2001; 1 male (cl 5.3 mm), 1 ovigerous female (cl 7.0 mm), ULLZ 18280 [ USNM 1706516], Florida, Tampa Bay, Old Tampa Bay, coll. C.H. Saloman, 27.04.1968; 2 males (cl 6.1, 6.9 mm), 1 ovigerous female (cl 9.9 mm), ULLZ 7741 [ USNM 1543258], Florida, Tampa Bay, Hillsborough Bay, Ballast Point, with Lepidophthalmus sp. , coll. D.L. Felder, 17.08.2005; 2 non-ovigerous females (cl 5.8, 10.8 mm), 3 ovigerous females (cl 9.0– 9.8 mm), ULLZ 5918 [ USNM 1541196], same locality as previous, coll. D.L. Felder & K. Strasser, 07.08.1999; 4 non-ovigerous females (cl 3.6–8.7 mm), 7 ovigerous females (cl 4.4–8.3 mm), ULLZ 17329 [ USNM 1550322], Florida, Choctawhatchee Bay, with Leptidophthalmus sp., coll. D.L. Felder, 05.06.2002; 2 males (cl 7.4–8.5 mm), 6 females (cl 4.4–9.8 mm), ULLZ 18231 [ USNM 1706497], same locality as previous, with Lepidophthalmus louisianensis , coll. D.L. Felder, 26.04.1990; 3 males (cl 7.1–9.5 mm), 3 females (cl 6.4–9.0 mm), ULLZ 18224 [ USNM 1706490], same locality as previous, coll. D.L. Felder & F.L. Mantelatto, 21.07.2001; 2 females (cl 8.2, 9.4 mm), ULLZ 5904 [ USNM 1541182], same locality as previous, with Lepidophthalmus sp. , coll. D.L. Felder, 26.04.1990; 3 males (cl 5.0– 8.6 mm), 4 ovigerous females (cl 3.9–8.5 mm), ULLZ 5922 [ USNM 1541200], same locality as previous, coll. D.L. Felder, 08.08.1999; 3 males (cl 5.3–5.9 mm), 1 female (cl 9.6 mm), ULLZ 18229 [ USNM 1706495], Florida, Pensacola Bay, coll. F.L. Mantelatto & R. Robles, 21.07.2001; 2 males (cl 2.6, 4.4 mm), 7 females (cl 4.7–8.0 mm), ULLZ 18230 [ USNM 1706496], Florida, Pensacola Bay, north side of Santa Rosa Island, with L. louisianensis , coll. D.L. Felder, 16.12.1999; 8 males (cl 3.0– 8.2 mm), 3 non-ovigerous females (cl 3.2–5.5 mm), 6 ovigerous females (cl 3.4–6.0 mm), ULLZ 5606 [ USNM 1541051], Florida, Pensacola, with L. louisianensis , coll. D.L. Felder & R. Robles, 17.08.2001; 1 male (cl 4.9 mm), 2 females (cl 7.2–7.5 mm), ULLZ 18278 [ USNM 1706515], Florida, near St. Teresa, Alligator Harbor, near FSU Coastal and Marine Laboratory, with Lepidophthalmus sp. , coll. R. Griffis, 28.06.1992; 2 females (cl 4.2, 4.3 mm), ULLZ 6697 [ USNM 1541568], Florida, Ft. Pierce, coll. R. Gulledge & K. Reed, 16.09.2003; 1 male (cl 7.0 mm), 1 non-ovigerous female (cl 7.6 mm), 2 ovigerous females (cl 6.2, 6.4 mm), ULLZ 18710 [ USNM 1706536], same locality as previous, with Upogebia affinis , male specimen also with Rathbunixa sayana , coll. R.B. Manning & D.L. Felder, 11.07.1984; 1 male (cl 5.1 mm), ULLZ 6049 [ USNM 1542551], Florida, Ft. Pierce, A1A causeway, with Upogebia sp. , coll. D.L. Felder, 18.07.2002; 1 ovigerous female (cl 7.3 mm), ULLZ 18276 [ USNM 1706513], Florida, Ft. Pierce, state park north of A1A causeway, with Upogebia sp. , coll. D.L. Felder, 20.07.1994; 1 ovigerous female (cl 5.9 mm), ULLZ 5818 [ USNM 1541154], Florida, Ft. Pierce, Little Jim Island, coll. D.L. Felder, 23.07.2004; 1 male (cl 8.8 mm), 1 non-ovigerous female (cl 7.5 mm), 1 ovigerous female (cl 10.5 mm), ULLZ 17846 [ USNM 1706381], Mississippi, Bay St. Louis, with L. louisianensis , coll. J.A. Scioli, 17.08.2017; 3 males (cl 6.8–8.1 mm), 5 females (cl 5.1–8.4 mm), ULLZ 17847 [ USNM 1706382], same locality as previous, with L. louisianensis , coll. J.A. Scioli, 13.09.2017; 2 males (cl 4.0, 5.8 mm), 1 female (cl 7.7 mm), ULLZ 17848 [ USNM 1706383], same locality as previous, with L. louisianensis , coll. J.A. Scioli, 29.09.2017; 1 male (cl 8.8 mm), 1 female (cl 5.3 mm), ULLZ 18053 [ USNM 1706385], same locality as previous, with L. louisianensis (host catalog no. ULLZ 18055 [ USNM 1706824]), coll. J.A. Scioli, 21.03.2018; 1 female (cl 8.4 mm), ULLZ 18222 [ USNM 1706488], same locality as previous, coll. P. Klerks, S. Pecnik, et al., 02.08.2013; 1 male (cl 7.0 mm), ULLZ 18223 [ USNM 1706489], same locality as previous, with L. louisianensis , coll. D.L. Felder, 01.05.2001; 1 ovigerous female (cl 4.6 mm), same locality as previous, with Lepidophthalmus sp. , coll. D.L. Felder, 20.06.1990; 1 male (cl 7.5 mm), 1 female (cl 6.7 mm), ULLZ 5594 [ USNM 1542623], same locality as previous, with L. louisianensis , coll. L.A. McGraw & R. Robles, 20.07.1999; 1 male (cl 8.6 mm), ULLZ 5603 [ USNM 1542550], same locality as previous, with L. louisianensis , coll. D.L. Felder & R. Robles, 19.03.1999; 1 female (cl 7.8 mm), ULLZ 18277 [ USNM 1706514], Alabama, Ono Island, Old River, with callianassid, coll. R. Bauer & M. Dardeau, 14.03.1986. Mexico: 1 female (cl 6.0 mm), ULLZ 5593 [ USNM 1541048], Campeche, Nuevo Campechito, with Lepidophthalmus statoni (host catalog no. ULLZ 5592 [ USNM 1541047]), coll. D.L. Felder & R. Robles, 07.07.2002.

Description. See Williams (1965).

Color in life. Semitransparent faint bluish tinge; body with sparse red chromatophores throughout; walking legs semitransparent, without chromatophores; ovary green ( Fig. 25h ).

Type locality. Gallants Point , Newport River, Carteret County, North Carolina, USA .

Distribution. Atlantic coast of USA and Gulf of Mexico: Atlantic coasts of North Carolina ( Williams 1965; Felder et al. 2003; present study) and Florida (present study); Gulf coasts of Florida ( Saloman 1971; Felder & Rodrigues 1993; Posey et al. 1998; Felder et al. 2003; present study), Alabama (present study), Mississippi ( Dawson 1967; Felder & Rodrigues 1993; Felder et al. 2003; Scioli et al. 2020; present study), Tamaulipas ( Felder & Rodrigues 1993), Veracruz ( Álvarez et al. 1999; Hermoso-Salazar 2001), and Campeche (Felder 2015; present study).

Ecology. Obligate burrow cohabitant of several species of axiid and gebiid mudshrimps: associated with Lepidophthalmus louisianensis ( Schmitt, 1935) in the gulf coasts of Florida, Mississippi, and Tamaulipas ( Dawson, 1967; Felder & Rodrigues, 1993; present study), Lepidophthalmus statoni Felder, 2015 in Campeche (Felder, 2015; present study), N. grandimana in Veracruz ( Hermoso-Salazar, 2001), and U. affinis in North Carolina and the Atlantic and gulf coasts of Florida ( Williams, 1965; Saloman, 1971; present study).

Remarks. The range of L. forceps is restricted to the Atlantic coast of the USA and the Gulf of Mexico, as records of L. forceps from the Caribbean Sea and Brazil are referable to L. degravei n. sp. (see above). As mentioned above, L. forceps can be distinguished from the morphologically similar L. degravei n. sp. based on the proportions of the antennular peduncle, which are longer relative to their breadth in L. forceps . Although smaller-bodied specimens of L. forceps have less slender antennules than larger-bodied specimens (compare Fig. 21i vs. Fig. 21j ); in all cases the second antennular article is longer relative to its breadth than in individuals of L. degravei n. sp.

Several specimens of L. forceps from Mississippi and Campeche have caudal appendices on the distal margins of the uropodal endopods ( Fig. 21k ). These structures were not reported in the type specimens of L. forceps by Williams (1965). Although these structures have been previously reported from specimens identified as L. forceps or L. cf. forceps ( Anker et al. 2008; Rosário et al. 2020), those records represent t L. degravei n. sp. Therefore, this study represents the first report of these structures in L. forceps sensu stricto.

Figures

FIGURE 21. Leptalpheus azuero Anker, 2011 (a–e), Leptalpheus felderi Anker, Vera Caripe & Lira, 2006 (f–h), and Leptalpheus forceps Williams, 1965 (i–k). Leptalpheus azuero: ovigerous female (cl 4.7 mm), ULLZ 11823 [USNM 1545912], from Las Peñitas, Nicaragua (a, b) and ovigerous female (cl 6.4 mm), ULLZ 5917 [USNM 1541195], from Estero Aserradores, Nicaragua (c–e): a, major chela, lateral; b, same, mesial; c, major chela, dorsolateral; d, same, dactylus and pollex, dorsolateral; e, same, fingers opened, lateral. Leptalpheus felderi: male (cl 8.5 mm), UF 48038, from Wachapreague, Virginia, USA: f, major chela, mesial; g, same, dorsolateral; h, same, dactylus and pollex, dorsolateral. Leptalpheus forceps: male (cl 8.8 mm) (i), female (cl 5.3 mm) (j), ULLZ 18053 [USNM 1706385], from Bay St. Louis, Mississippi, USA and female (cl 6.0 mm) (k), ULLZ 5593 [USNM 1541048], from Nuevo Campechito, Mexico: i, frontal margin, dorsal; j, same; k, uropod with caudal appendix, dorsal.

FIGURE 25. Color patterns of Leptalpheus degravei n. sp. and Leptalpheus forceps Williams, 1965. Leptalpheus degravei n. sp.: a, paratype male (cl 3.6 mm), ULLZ 18511 [USNM 1705361], from Bocas del Toro, Panama, dorsal; b, same, lateral; c, holotype male (cl 5.1 mm), ULLZ 18510 [USNM 1705360], from Bocas del Toro, Panama, dorsal; d, same, lateral; e, paratype female (cl 3.7 mm), ULLZ 18506 [USNM 1705356], from Bocas del Toro, Panama, dorsal; f, same, lateral; g, paratype ovigerous female (cl 4.7 mm), ULLZ 18512 [USNM 1705362], from Bocas del Toro, Panama, lateral. Leptalpheus forceps: h, ovigerous female (cl 5.9 mm), ULLZ 5818 [USNM 1541154], from Ft. Pierce, Florida, USA, lateral. Photographs by the first author (a–g) and third author (h).