Paguristes spinipes A. Milne-Edwards, 1880

Lemaitre, Rafael & Tavares, Marcos, 2015, New taxonomic and distributional information on hermit crabs (Crustacea: Anomura: Paguroidea) from the Gulf of Mexico, Caribbean Sea, and Atlantic coast of South America, Zootaxa 3994 (4), pp. 451-506 : 472-480

Paguristes spinipes A. Milne-Edwards, 1880

( Figs. 1 D, 6–11)

Paguristes spinipes A. Milne-Edwards, 1880: 44 (type locality: Grenada, USCSS Blake, sta 253, 11°25’N, 62°04’15”W).— A. Milne-Edwards & Bouvier, 1893: 33, pl. 3, figs 1–13.— Lemaitre & Felder, 2012: 69, tab. 1.—Forest & de Saint Laurent, 1968: 68.— Williams, 1965: 118, fig. 95.—Williams, 1984: 204, fig. 143.— Coelho & Ramos, 1973: 167.— Provenzano, 1978: 512, figs 1–8 (larvae).— Coelho & Santos, 1980: 142.— Abele & Kim, 1986: 346, 347, figs. [unnumbered] f, g.— Coelho & Ramos-Porto, 1987: 48.— Rieger, 1998: 419.— Melo, 1999: 86, figs. 39, 40.— Manjón-Cabeza et al., 2002: 137.— Nizinski, 2003: 118.— Rodríguez-Almaraz & Zavala-Flores, 2005: 293, fig. 20.—Felder et al., 2009: 1069.— McLaughlin et al., 2010: 23.

Paguristes visor Henderson, 1888: 78 , pl. 8, fig. 3.—Forest & de Saint Laurent, 1968: 68.

Paguristes armatus Hay, 1917: 73 .— Hay & Shore, 1918: 409, fig. 15, pl. 30, fig. 7.

Type material. Holotype female 6.6 mm, USCSS Blake, sta 253, Caribbean Sea, Grenada, 11°25’N, 62°04’15”W, 176 m, 27 Feb 1879, coll. A. Agassiz ( MCZ 3985).

Lectotype, herein selected, of Paguristes visor Henderson, 1888 : male 10.0 mm, H.M.S. Challenger, sta 122, off Alagoas, 9°5’S, 34°50’W, 640 m, 10 Sep 1873 (BM 88.33); paralectotype: 1 male 5.0 mm, same sta as lectotype (BM 88.33).

Holotype of Paguristes armatus Hay, 1917 : female 5.1 mm, [off coast of North Carolina], 30 miles due S of Cape Lookout lightship, [no depth], [day, month not recorded] 1915 ( USNM 49699).

New material. Southeastern coast of United States. Off Cape Canaveral, Florida: R/V Combat, sta 235, 27°27’N, 78°58’W, 329 m, 2 Feb 1957: 1 female 8.3 mm ( USNM 101363); R/V Combat, sta 237, 27°28”N, 78°44’W, 393 m, 3 Feb 1957: 1 male 11.6 mm ( USNM 101665).

Bahamas. New Providence Island, off Goulding's Cay, JSL-II-2587, 25°02’30”N, 35°12’W, 518 m, 25 May 1995, coll. M. Harasewych & T. Askew: 1 male 6.6 mm ( USNM 275960).

Caribbean Sea. Cuba, [NE of Havana], USFC Albatross, sta 2327, 23°11’45”N, 82°17’54”W, 333 m, 17 Jan 1885: 1 male 4.3 mm ( USNM 267540); Quintana Roo ( Mexico), Cozumel Island, Yucatan Channel, USFC Albatross, sta 2354, 20°59’30”N, 86°23’45”W, 238 m, 22 Jan 1885: 1 female 8.6 mm ( USNM 9550); St. Lucia [Lesser Antilles], R/V Oregon, sta 5956, 13°40’N, 60°54’W, 229 m, 10 Mar 1966: 4 males 8.6–9.6 mm, 1 female 8.8 mm ( USNM 265305).

Gulf of Mexico. Off Sarasota, Florida, sta B- 166, 179 m, 8 May 1952, coll. R. Stewart: 1 male 10.9 mm, 1 female 9.2 mm ( USNM 103441); off western coast of Florida: sta MMS-SOFLA/ 2/II-32 -TDS, 26°16’40”N, 84°04’05”W, 137 m, 25 Jul 1981: 3 males 3.6–5.1 mm, 1 female 3.0 mm, 1 ov female 6.6 mm ( USNM 1017882); same sta data, 6 Feb 1982: 2 females 3.7, 6.4 mm ( USNM 1021172); same sta data: 2 males 3.0, 3.3 mm, 1 female 9.8 mm ( USNM 1073011); sta MMS-SOFLA/ 2/III-35 -TDS, 25°44’50”N, 84°21’02”W, 159 m, 7 Feb 1982: 1 male 3.0 mm ( USNM 1017913); same sta data: 2 males 3.0, 7.5 mm, 1 female 2.7 mm ( USNM 1017915); sta MMS- SOFLA / 2/III-36 -TDS, 25°16’50”N, 83°57’21”W, 127 m, 9 Feb 1982: 1 female 7.8 mm ( USNM 1072742); sta MMS-SOFLA/ 2/III-38 -TDS, 25°16’30”N, 84°14’46”W, 159 m, 10 Feb 1982: 1 female 5.7 mm ( USNM 1021170); sta MMS-SOFLA/ 2/III-39 -RDS, 24°47’10 N, 83°55’22”W, 151.5 m, 10 Feb 1982: 1 male 9.5 mm ( USNM 1021171).

Curaçao. Curasub 12–03: 28 May 2012, 145– 183 m: male 4.8 mm, CURI 12040 ( USNM 1253258); Curasub 12: 1 male 3.3 mm, CURI 12029, no other data recorded ( USNM 1253266); Curasub 12–11: 220 m, 6 Aug 2012: 1 female 6.7 mm, CURI 12077 ( USNM 1253265), 1 female, 7.1 mm, CURI 12078 ( USNM 1253267), 1 male 6.4 mm, CURI 12083 ( USNM 1253268); Curasub 12–12, 290 m, 7 Aug 2012: 1 ov female 8.5 mm, CURI 12124 ( USNM 1253269), 1 male 7.5 mm, CURI 12125 ( USNM 1253270), 1 male 6.0 mm, CURI 12131 ( USNM 1253271), 1 ov female 6.7 mm, CURI 12132 ( USNM 1253272); Curasub 12–13, 286 m, 8 Aug 2012: 1 male 6.6 mm, CURI 12142 ( USNM 1253273); Curasub 12–02, 201 m, 22 May 2012: 1 female 9.6 mm, CURI 12152 ( USNM 1253274).

Brazil. Bahia, REVIZEE—Comissão Central 2, PS/V Astro Garoupa, sta 6C, 15°54’23”S, 38°02’53”W, 340 m, 25 Oct 1997: 1 female 7.8 mm ( MZUSP 16121).

Diagnosis. Thirteen pairs of quadriserial gills. Shield ( Figs. 6 A, 10A) distinctly longer than broad, ranging 1.1–1.3 as long as broad (narrowness increasing with specimen size); dorsal surface with scattered irregular rows of tufts of setae more numerous dorsolaterally, and irregular rows of small spines or tubercles on dorsolateral surface; anteromedian region subdivided into 2 low, rounded ridges. Rostrum ( Figs. 6 A, 9A, B, 10A) distinctly exceeding lateral projections, acutely triangular, often slender, and pointing slightly ventrally. Lateral projections broadly subtriangular, terminating in small spine. Ocular peduncles long, about 0.8 to as long as shield length, with dorsal row of tufts of short setae; corneas weakly dilated; ocular acicles subtriangular, each terminating in simple or rarely bifid spine (on one or both sides). Antennular peduncles slender, not exceeding distal margins of corneas. Antennal peduncles short, slender, at most reaching to about distal two-thirds of ocular peduncles; second segment with distolateral angle strongly produced, terminating in strong, bifid spine; acicle reaching to about distal threefourths of fifth antennal segment, terminating in bifid spine and with 1–4 distinct spines mesially and laterally; fourth segment with weak lateral dorsodistal spine; flagellum exceeding tip of extended chelipeds, with series of long and short setae every 3 or 4 flagellar articles (long setae 3 or 4 articles in length, short setae ≤ articles in length). Third maxilliped merus armed with 2–5 ventral spines; ischium with crista dentata consisting of row of 18–20 subequal, corneous teeth; basis with mesial row of small calcareous spines. Chelipeds ( Figs. 1 D, 6B–E, 10B–E) slightly unequal in strength, moderately setose, similarly armed on dorsal surfaces of carpi and chelae with dense spines (most corneous tipped and often curved forwardly); carpi, palm and dactyl each with row of strong spines on dorsomesial margin; meri mostly smooth except for scattered small tubercles, and row of small spines on dorsodistal and ventromesial margins. Ambulatory legs ( Figs. 7 , 11 A–H) with moderately dense setae or tufts of setae, denser on propodi and dactyls; carpi with dorsolateral longitudinal groove; carpus and propodus of first ambulatory leg with dorsal row of strong spines; carpus of second ambulatory leg with weak row of small spines, propodus unarmed except for tufts of setae on dorsal and ventral margins; dactyls with numerous tufts of setae, and ventromesial row of 16–33 small corneous spines (spine number increasing with specimen size). Fourth pereopod ( Fig. 8 A) semichelate; dactyl with prominent preungual process at base of claw, and ventrolateral row of corneous spinules; propodal rasp consisting of 1–4 rows of ovate, corneous scales (number of rows increasing distally). Protopod of uropods ( Fig. 6 G) with several rows of small, sharp calcareous or corneous spines on ventral angle. Telson ( Fig. 6 F) with posterior lobes slightly to distinctly asymmetrical, roundly subtriangular, setose, and separated by shallow, median cleft. Male with paired first and second gonopods ( Figs. 9 C–E, 11I, J); first gonopod well developed, inferior lamella with distal margin armed with row of small, hooked corneous spines, external lobe naked, internal lobe with long setae on mesial margin; second gonopod with distal segment slightly twisted, setose distally. Female with paired first gonopods ( Fig. 8 B, C) and left pleopods 2–5; first gonopod with distal lobe slender, with plumose setae; with well developed, subcircular brood pouch ( Fig. 6 H) over left pleopod 4.

Color ( Fig. 1 D). General background color light to dark orange mottled with whitish spots and spines, chelipeds with whitish or dark orange-tipped spines. Shield with darker orange or reddish color anteriorly except for distinct white anterior margin and rostrum. Ocular peduncles white throughout dorsal surfaces, light orange ventrally; corneas light gray to light blue. Meri of chelipeds and ambulatory legs each distinctly white band distally and extending posteriorly on about one-third of dorsal margin, followed by dark orange-red band fading to light orange posteriorly, latter band darker on meri of chelipeds and gradually diminishing in darkness posteriorly on meri of ambulatory legs.

Distribution. Western Atlantic: from off the North Carolina to Florida, Bahamas, Gulf of Mexico, Caribbean Sea, and Brazil (off Alagoas and Bahia). Depth: 70 to 640 m.

Remarks. The single female specimen reported herein from Bahia, Brazil (MZUSP 16121) agrees with the description and types of Paguristes spinipes , originally obtained from further north (Alagoas, Brazil), except for the terminally bifid condition of its ocular acicles, and the slightly less asymmetrical condition of the terminal lobes of its telson. In some Paguristes species, however, the terminal spine of the ocular acicle is known to occasionally vary from simple to bifid ( McLaughlin 2004b), and the subtle difference in telson asymmetry can be attributed to the expected intraspecific variation normally observed in many Paguristes species.

Paguristes visor Henderson, 1888 , a species described using specimens from off Alagoas, Brazil, has been considered a junior synonym of P. spinipes for more than a century ( A. Milne-Edwards & Bouvier 1893; Forest & de Saint Laurent 1968; Williams 1984; McLaughlin et al. 2010), although modern reports of new material or discussions about the morphology and color pattern of P. spinipes are sketchy and scattered in the literature. Forest & de Saint Laurent (1968), without any comments, considered Paguristes armatus Hay, 1917 , a junior synonym of P. spinipes . Hay’s original description was brief, lacking illustrations, and was based on a single female specimen collected off North Carolina on the eastern coast of the United States, although subsequently Hay & Shore (1918) did publish an illustration of the cephalothorax and chelipeds, and a photograph, of the same female. We have studied the types of all these three taxa, and compared them to our Brazilian female specimen as well as other unreported materials of P. spinipes deposited in the USNM, and can confirm based on morphological grounds that they are all conspecific. Nevertheless, given that Henderson (1888) did not select a holotype among the three syntypes of his P. v i s o r, and the morphological complexities as yet incompletely understood of western Atlantic species of Paguristes , we consider wise to select a lectotype in order to fix the name of that taxon, and in case future evidence may show that name to represent a different, valid species.

The diagnosis presented herein of Paguristes spinipes includes observed variations in the listed material examined. The characteristic coloration of the meri of chelipeds and ambulatory legs of this species (i.e., with an orange-red or orange-yellow band) was only briefly mentioned by A. Milne-Edwards & Bouvier (1893), Hay & Shore (1918, as P. armatus ), and Forest & de Saint Laurent (1968). The detailed description presented herein of the coloration of this species is based on live specimens captured in waters off Curaçao, using the manned submersible Curasub, and brought to a laboratory aquarium for photographing ( Fig. 1 D).

Currently, Paguristes spinipes is considered to have a wide western Atlantic distribution ranging from the southeastern coast of the United States to off Bahia, Brazil. Previous to this study there had been no reports of this species in the Caribbean outside of the Antilles ( Cuba and Barbados). Herein we report for the first time this species in the Bahamas, the Caribbean Sea other than the Antilles (off Quintana Roo in the Yucatan Channel, and Curaçao), and provide only the second report for the Lesser Antilles ( St. Lucia). There are still no published records of P. spinipes along the vast stretch of South American coastline from Venezuela to Pernambuco in northern Brazil.

Henderson (1888: 88) described Paguristes visor based on one male and one female specimens from “off Pernambuco”, 09°05’S, 34°50’W. However, these geographic coordinates actually place the locality further south, off the State of Alagoas, which therefore is considered herein the correct type locality for this species.

Figures

FIGURE 6. Paguristes spinipes A. Milne-Edwards, 1880: female 7.8 mm, Brazil, REVIZEE—Comissão Central 2, sta 6 C (MZUSP 16121). A, shield and cephalic appendages, dorsal; B, right cheliped, dorsal; C, same, mesial; D, left cheliped, dorsal; E, same, mesial; F, telson, dorsal; G, left uropod, dorsal; H, female brood pouch on fourth pleopod. Scales: 5 mm (A – E, H), 1 mm (F, G).

FIGURE 7. Paguristes spinipes A. Milne-Edwards, 1880: female 7.8 mm, Brazil, REVIZEE—Comissão Central 2, sta 6 C (MZUSP 16121). A, C, right second pereopod in mesial (A) and lateral (C) view; B, D, right third pereopod in mesial (B) and lateral (D) view; E, G, left second pereopod in mesial (E) and lateral (G) view; F, H, left third pereopod in mesial (F) and lateral (H) view. Scale: 5 mm.

FIGURE 8. Paguristes spinipes A. Milne-Edwards, 1880: female 7.8 mm, Brazil, REVIZEE—Comissão Central 2, sta 6 C (MZUSP 16121). A, propodus and dactyl of right fourth pereopod, lateral; B, C, right first gonopod, anterior (B) and posterior (C). Scales: 1 mm.

FIGURE 9. Paguristes spinipes A. Milne-Edwards, 1880: A, male 10.9 mm, Gulf of Mexico, off Sarasota, Florida (USNM 103441); B, female 7.8 mm (USNM 1072742); C – F, male 4.8 mm, Curaçao, Curasub 12 – 03, CURI 12040 (USNM 1253258). A, B, anterior portion of shield with rostrum and lateral projections; C, left first male gonopod, lateral; D, same, mesial; E, distal margin of inferior lamella, posterior; F, left second male gonopod, lateral. Scales: 1 mm (A, B), 0.25 mm (C, D, F), 0.1 mm (E).

FIGURE 11. Paguristes spinipes A. Milne-Edwards, 1880. Lectotype of Paguristes visor Henderson, 1888: male 10.0 mm, Brazil, off Alagoas, H. M. S. Challenger, sta 122 (BM 88.33). A, C, leftt second pereopod in lateral (A) and mesial (C) view; B, D, right third pereopod in lateral (B) and mesial (D) view; E, G, left second pereopod in lateral (E) and mesial (G) view; F, H, left third pereopod in lateral (F) and mesial (H) view; I, male left first gonopod, posterior; J, male left second gonopod, mesial. Scales: 10 mm (A – H), 1 mm (I, J).

FIGURE 1. A, Coenobita clypeatus (Fabricius, 1787), in gastropod Melongena melongena: dry specimens [not sexed or measured], W. J. Broderip collection (NHM B 2195, NHM B 2196); B, Allodardanus bredini Haig & Provenzano, 1965, female 10.1 mm, in gastropod shell, Curaçao, Curasub 13 – 21 (USNM 1253325); C, Bathynarius anomalus (A. Milne-Edwards & Bouvier, 1893), in gastropod shell, male, 4.0 mm, Curaçao 12 – 11, CURI 12088 (USNM 1253278); D, Paguristes spinipes A. Milne-Edwards, 1880, in gastropod shell: male 4.8 mm, Curaçao, Curasub 12 – 03, CURI 12040 (USNM 1253258); E, Pylopagurus discoidalis (A. Milne-Edwards, 1880), in scaphopod shells: male 6.3 mm (right), 1 female 5.1 mm (center), 1 ov female 5.8 mm (left), Curaçao, Curasub 12 – 09 (right and center specimens: CURI 12030, CURI 12031, USNM 1253259; left specimen USNM 1253260). (Photos: A, H. Taylor, NHM; B – E, B. B. Brown, Curaçao Seaquarium). Scale: 20 mm (A).