Spathapagurus gladius

Spathapagurus gladius

( Benedict, 1892) comb. nov.

( Figs. 1A‒E View Figure 1 , 2A–F View Figure 2 , 3A–G View Figure 3 , 4A–F View Figure 4 )

Eupagurus gladius Benedict, 1892: 7 View in CoL [type locality: USFC Albatross sta 2823, Isla Cerralvo, Gulf of California]. — Nobili, 1901: 22.

Pagurus gladius View in CoL . — Rathbun,1910: 597. — Haig et al., 1970: 21. — Snyder-Conn, 1980: 283. — Brusca, 1980: 283, fig. 18.12. — Hendrickx, 1993: 309. — Hendrickx and Harvey, 1999: 372. — Boschi, 2000: 105. — Lemaitre and Cruz-Castaño, 2004: 78. — Hendrickx et al., 2006: 38. — Vargas and Cortés, 2006: 482. — McLaughlin et al., 2010: 33. — Olguín and Mantelatto, 2013: 439, figs.1, 2.

Type material. Lectotype herein selected, male (5.4 mm), USNM 16723 About USNM , USFC “Albatross” sta 2823, Isla Cerralvo , Gulf of California, 24º18’00”N 110º22’00”W, 48.5 m depth, 30 April 1888 GoogleMaps . Paralectotypes, 2 males (4.0 mm and 4.9mm), USNM 108262 About USNM , same station data as lectotype GoogleMaps .

Other material. Eastern Pacific: 1 ovig female (4.3 mm), USNM 110944, USFC “Albatross” sta 3026, 31º22’00”N 114º07’45”W, Gulf of California, Sonora, west of Adair Bay, 31 m, 25 March 1889; 4 males (3.7–4.8 mm), 1 ovig female (4.5 mm), USNM 16728 About USNM , USFC “Albatross”, same station data as previous lot ; 1 male (4.4 mm), ICML-EMU 4115 C, off Rocas Consag, Gulf of California , Mexico, R / V “El Puma”, CORTES 2 sta 37, 31°16’12”N 114°22’06”W, 32 m, 16 March 1985 GoogleMaps ; 2 males (5.9–6.0 mm), 3 ovig females (4.2–4.8 mm), ICML-EMU 4050 A, off Rocas Consag , Mexico, R / V “El Puma”, CORTES 1 sta 37, 31°16’12”N 114°22’30”W, 35 m, 9 May 1982 GoogleMaps ; 1 female (3.9 mm), 1 ovig female (4.2 mm), ICML-EMU 4051 , off Rocas Consag , Mexico, R / V “El Puma”, CORTES 2 sta 38, 31°09’18”N 114°15’30”W, 65 m, 16 March 1985 GoogleMaps ; 8 males (5.6–6.7 mm), 10 ovig females (4.7– 5.3 mm), ICML-EMU 4050 B, off Rocas Consag , Mexico, R / V “El Puma”, CORTES 1 sta 38, 31°09’N 114°15’18”W, 60 m, 9 May 1982 GoogleMaps ; 3 males (3.3– 4.0 mm), USNM 16727 About USNM , USFC “Albatross” sta 3022, Gulf of California, Sonora, San Jorge Bay , 30º58’30”N 113º17’15”W, 20 m, 24 March 1889 GoogleMaps ; 1 male (5.6 mm), USNM 16726 About USNM , USFC “Albatross” sta 3020, Gulf of California , Sonora, Cape Tepoca, El Desemboque, 30º37’30”N 113º07’00”W, 13 m, 24 March 1889 GoogleMaps ; 2 males (3.7–4.5 mm), ICML-EMU 4115 B, N. of Isla Tiburón, Gulf of California , Mexico, R / V “El Puma”, CORTES 2 sta 27, 29°28’N 112°29’12”W, 41 m, 14 March 1985 GoogleMaps ; 1 male (3.9 mm), 2 ovig females (3.6– 4.5 mm), ICML-EMU 4069 , Tiburón Island, Gulf of California , Mexico, R / V “El Puma”, CORTES 1 sta 27, 29°26’00’’N 112°26’00’’W, 14 March 1985 GoogleMaps ; 4 males (3.7–4.9 mm), 1 female (3.0 mm),1 ovig female (5.4 mm), USNM 16725 About USNM , USFC “Albatross” sta 3014, Gulf of California, south of Tiburón Island , Sonora, 28º28’00”N 112º04’30”W, 53 m, 23 March 1889 GoogleMaps ; 1 male (3.7 mm), 1 ovig female (3.1 mm), USNM 16724 About USNM , USFC “Albatross” sta 3013, Gulf of California, south of Tiburón Island , Sonora, 28º23’45”N 111º58’00”W, 26 m, 23 March 1889 GoogleMaps ; 6 males (4.7–5.7 mm), ICML-EMU 4115 A, off Cabo San Miguel, Gulf of California , Mexico, R / V “ElPuma”,CORTES2sta20, 28°08’12”N 112°45’24”W, 55 m, 13 March1985 GoogleMaps ; 1 ovig female (2.8 mm), ICML-EMU 4116 B, off Punta Arboleda, Gulf of California , Mexico, R / V “El Puma”, CORTES 1 sta 16, 26°55’36’’N 110°05’06’’W, 23 m, 5 May 1982 GoogleMaps ; 1 male (4.5 mm), USNM 16729 About USNM , USFC “Albatross” sta 3037, Gulf of California, Sonora, south of Guaymas , 27º45’00”N 110º45’00”W, 37 m, 31 March 1889 GoogleMaps ; 1female (1.8mm), USNM1289682 About USNM , AHFR / V “Velero III”, sta 520–36, 24º23’N 111º33’W, Baja California Sur, Agua Verde Bay , 9–18 m, 27 February 1936 GoogleMaps ; 3 males (4.2–5.5 mm), 1 female (3.2 mm), ICML-EMU 4116 A, off Santa María Bay, Gulf of California , Mexico, R / V “El Puma”, CORTES 1 sta 3, 25°02’42”N 108°31’30”W, 28 m, 3 May 1985 GoogleMaps ; several disintegrated dried specimens, USNM 16722 About USNM , USFC “Albatross”, sta 2822, Gulf of California, Baja California, Cerralvo Island , 24º16’0”N 110º22’00”W, 28 m, 30 April 1888 GoogleMaps ; 1 male (2.4mm), USNM 1289681 About USNM , USFC “Albatross”, Lower California cruise, [no sta number or other data] ; 2 males (2.8 mm and 4.3 mm), 3 females (2.2– 2.7 mm), USNM 1156044 About USNM , Baja California Sur, Santa Maria Bay , 18.3–36.6 m, 22 January 1938, coll. S.A. Glassell ; 1 male (4.1 mm), ICML-EMU 1347 , Mazatlán Bay, Gulf of California , Mexico, B/E FC1 BBMAZ cruise 13 sta 5, 23°12’06”N 106°28’56”W, 18 m, 26 June 1980 GoogleMaps ; 1 male (4.9 mm), ICML-EMU 0928 , Mazatlán Bay, Gulf of California , Mexico, B/E FC1, BBMAZ cruise 18 sta 18, 23°10’53”N 106°26’11”W, 13 m, 14 March 1981 GoogleMaps ; 1 ovig female (4.4 mm), ICML-EMU 4116 C, off Banco Gordo, Gulf of California , Mexico, R / V “El Puma”, CORTES 1 sta 55, 23°08’30”N 109°27’24”W, 38 m, 13 May 1982 GoogleMaps ; 1 female (1.9 mm), 1 ovig female (2.2 mm), USNM 1289685 About USNM , Mexico, Tenacatita Bay , AHF R / V “Velero III”, sta 485–35, 19º18’N 104º50’W, 9 m, 15 February 1935 GoogleMaps ; 1 male (2.5 mm), 2 ovig females (2.2 mm and 2.3 mm), USNM 1289686 About USNM , Mexico, Tenacatita Bay , AHF R / V “Velero III”, same station data as previous GoogleMaps ; 1 female (2.2 mm), USNM1289677 About USNM , Mexico, Acapulco Bay , 16º50’N 99º53’W, April 1930, coll. H. Lowe GoogleMaps ; 1 male (3.3 mm), 2 females (2.8mm and 3.3 mm), USNM 1289687 About USNM , Mexico, Tangola-Tangola Bay, Santa Cruz Bay , AHF R / V “Velero III”, sta 259–34, 15º45’N 96º06’12”W, 27–37 m, 28 February 1934 GoogleMaps ; 15 males (1.6–3.1 mm), 2 females (2.2 mm and 2.3 mm), 5 ovig females (1.8–3.4 mm), USNM 1289674 About USNM , Costa Rica, Salinas Bay , AHF R / V “Velero III”, sta 480–35, 11º04’10”N 85º44’40”W, 22 m, 11 February 1935 GoogleMaps ; 1 male (1.9 mm), USNM1289688 About USNM , Costa Rica, Salinas Bay , AHF R / V “Velero III”, sta 478–35, 11º03’13”N 85º44’10”W, 2.7 m, 11February 1935 GoogleMaps ; 1male (4.0 mm), USNM 1289684 About USNM , Costa Rica, Port Parker , AHF R / V “Velero III”, sta 472–35, 10º57’50”N 85º48’45”W, 55 m, 9 February 1935 GoogleMaps ; 2 males (2.8 mm and 4.6 mm), 1 female (1.8 mm), 1 ovig female (3.0 mm), USNM 1289676 About USNM , Ecuador, Santa Elena Bay , 02º10’S 80º52’W, [no depth], 9 February1934 GoogleMaps ; 1 male (3.7 mm), 3females (1.9–2.8mm), USNM1289675 About USNM , R / V “Stranger” sta 15, Ecuador, Santa Elena, La Libertad, 02º12’S 80º54’W, 15 m, 21 February 1939, coll. Fred E. Lewis. GoogleMaps

Redescription. Eleven pairs of biserial gills ( Fig. 1A View Figure 1 ) at most weakly divided distally. Shield ( Fig.1B View Figure 1 ) about 1.3 times as broad as long; anterolateral margins sloping; anterior margin between rostrum and lateral projections concave, fringed by short setae; anterolateral angle with slit; posterior margin roundly truncate; dorsal surface flattened, with numerous tufts of short setae. Rostrum obtusely triangular, length slightly exceeding lateral projections in distal extension; lateral projections obtusely triangular, armed with small terminal spine.

Ocular peduncles ( Fig. 1B View Figure 1 ) stout, short, approximately 0.6 length of shield, slightly constricted medially, dorsal surface with few tufts of short setae. Cornea dilated, width about 0.7 length of ocular peduncle. Ocular acicles bluntly subtriangular, weakly concave dorsally, tip subacute, with small, submarginal spine, and moderately long setae, acicles separated basally by about basal width of one acicle.

Antennular peduncle ( Fig. 1B View Figure 1 ), when extended, exceeding ocular peduncles by 0.7 length of ultimate segment; ultimate and penultimate segments both with few short setae on dorsal surface; basal segment with moderately strong spines on laterosubdistal margin.

Antennal peduncle ( Fig. 1B View Figure 1 ) moderately long, when extended, exceeding ocular peduncle by 0.5 length of ultimate segment. Fifth segment unarmed, with scattered setae on dorsal and ventral margins. Fourth segment unarmed, with scattered setae. Third segment with small spine on ventrodistal margin, partially obscured by tufts of setae. Second segment with dorsolateral distal angle reaching about 0.4 of antennal acicle, terminating in strong spine; lateral margin unarmed; dorsomesial distal angle with strong spine; mesial margin with short setae. First segment lateral margin unarmed; ventral margin produced, with 1 small laterodistal spine. Antennal acicles moderately long, exceeding length of ocular peduncles by 0.3 times their length, terminating in spine; dorsal surface and mesial margin with tufts of short setae. Antennal f lagella long, reaching to distal third of right cheliped; each article with few bristles less than 1 article length.

Mandible ( Fig. 2A View Figure 2 ) with upper and lower incisor edges calcareous; palp3-segmented, ultimate segment setose, slightly longer than combined length of penultimate and basal segments. Maxillule ( Fig. 2B View Figure 2 ) with proximal endite subquadrate, distal endite subrectangular, enlarged distally; endopod with external lobe moderately developed, not recurved; internal lobe with long terminal bristle. Maxilla ( Fig. 2C View Figure 2 ) with endopodite inflated basally, exceeding scaphognathite by 0.3 in distal extension. First maxilliped ( Fig. 2D View Figure 2 ) with endopodite approximately 0.6 length of exopodite; basal segment of exopodite inf lated. Second maxilliped ( Fig. 2E View Figure 2 ) without distinguishing characters. Basis of third maxilliped ( Fig. 2F View Figure 2 ) with 1 strong spine; ischium with welldeveloped crista dentata, consisting of 9–12 corneous-tipped teeth and 1 accessory tooth; merus unarmed.

Chelipeds unequal, right ( Fig. 3A–D View Figure 3 ) long and slender, twice as long as left cheliped; chela 2 (female) to 3 (males) times longer than broad; palm, fixed finger and dactyl slender, dorsoventrally compressed. Fingers each with dorsal and ventral row of tufts of setae parallel to and near cutting edges; dorsal surfaces somewhat flattened, with numerous small tubercles or protuberances, each terminating in calcareous claw, often worn. Dactyl moderately long, about0.8 length of propodus; cutting edge with row of 6 or 7 strong calcareous teeth on proximal half, and row of small calcareous teeth on distal half; dorsomesial margin delimited by row of small tubercles decreasing in size distally. Fixed finger broader than dactyl, broader in females ( Fig. 3D View Figure 3 ) than in males ( Fig. 3A View Figure 3 ); lateral margin delimited by row of tubercles decreasing in size distally;cutting edge with moderately prominent calcareous tooth at midlength, 5–7 smaller calcareous teeth on proximal half, and row of small calcareous teeth on distal half.Palm slightly shorter than carpus, dorsomesial margin distinctly delimited by row of small granules or tubercles; mesial face rounded, sloping, covered with numerous small tubercles; dorsolateral margin well delimited by row of spines or tubercles, dorsal surface moderately elevated along midline, covered with numerous small granules or tubercles. Carpus distinctly longer than merus, with dorsal surface convex, and moderately produced ventrally, 1.8–2.1 times longer than broad, broader in females than in males; dorsomesial margin weakly expanded, armed with row of small spines of different size; dorsodistal margin with row of strong or moderately strong spines; dorsal surface with numerous spiny granules or tubercles and scattered setae; dorsolateral margin delimited by row of spiny granules or tubercles; mesial, lateral, and ventral surfaces with small granules or tubercles; mesial face strongly sloping, nearly perpendicular to dorsal face. Merus subtriangular; dorsodistal margin with 3–5 well-spaced, small spines; dorsal and ventrolateral surfaces with transverse, sparsely setose furrows; ventromesial and ventrolateral margins each with row of spines, ventrolateral angle with several spiny teeth, ventral surface usually with few scattered granules and sparse, moderately long setae. Ischium unarmed or with inconspicuous small protuberances on dorsal and ventral surfaces.

Left cheliped( Fig.3E–G View Figure 3 )slender, scarcely reaching base of right cheliped palm; dactyl and fixed finger weakly arched ventrally; fingers with ventral row of setal tufts parallel to and near cutting edges, each finger terminating in inwardly directed corneous claw.Dactyl 1.8–2.0 length of palm; cutting edge with row of small corneous teeth; dorsal surface slightly elevated in midline, with row of small granules or tubercles on proximal half; dorsomesial margin not well defined, with row of small spine-like tubercles extending on proximal half, mesial surface with row of long setal tufts. Fixed finger broader than dactyl, somewhat expanded laterally, dorsal surface with few small tubercles or granules on proximal half; cutting edge with row of small calcareous teeth interspersed with small corneous teeth. Palm 0.6–0.7 length of carpus; dorsomesial margin with row of small granules or tubercles; mesial surface with scattered small tubercles and short setae; dorsal surface with numerous minute granules or tubercles, and distinctly elevated on midline and armed with longitudinal row of small spine-like tubercles; dorsolateral margin with row of small spinules or granules; ventral surface with scattered tufts of long setae. Carpus slightly shorter than merus; dorsal surface somewhat f lattened, with dorsomesial and dorsolateral rows of spine-like tubercles; dorsodistal margin with row of spine-like tubercles; mesial face with numerous small tubercles and short setae; lateral face covered with small spine-like tubercles, ventrolateral margin produced in moderately strong spine. Merus subtriangular;dorsal surface with transverse, sparsely setose furrows; mesial surface with few small tubercles and with tufts near to ventral margin; ventrolateral margin with row of spines largest distally; ventral surface granulose and tufts of short setae. Ischium with row of small calcareous spines on ventromesial margin.

Second and third pereiopods similar left from right, slender, of similar length ( Fig. 4A–D View Figure 4 ). Dactyls curved, twisted, 1.5–1.7 times length of propodi; dorsal margins each with few spines on proximal half, and sparse row of short stiff setae; ventromesial margins each with row of 20–22 small corneous spinules; mesial faces each with row of small corneous spines near dorsal margin, and weak median longitudinal sulcus; lateral faces each with weak longitudinal sulcus. Propodi about 1.3 times length of carpi; dorsal margin each with row of small spines or spine-like tubercles and sparse short setae; dorsomesial and dorsolateral faces each with numerous small spine-like tubercles or tubercles fringed with very short stiff setae; ventral margin with small spines or granules; ventromesial and ventrolateral faces with numerous small spine-like tubercles or tubercles, fringed with very short setae. Carpi each with row of spines and sparse setae on dorsal margin, and 1 or 2 spines on dorsodistal angle; mesial surfaces each with few small granules near dorsal margin; dorsolateral faces each with few spine-like tubercles fringed distally with short stiff setae; ventral margin unarmed. Meri with low protuberances and tufts of setae on dorsal surfaces; lateral and mesial faces glabrous; ventral margin with few minute spines or tubercles, at least distally. Ischia unarmed or with minute granules on ventromesial margin and stiff setae or bristles dorsally. Anterior lobe of thoracic sternite XII (of third pereiopods) ( Fig. 1C View Figure 1 ) subcircular, with long stiff bristles.

Fourth pereiopods ( Fig. 4E View Figure 4 ) without preungual process at base of claw on lateral face of dactyl; propodal rasp with single row of lanceolate scales.

Fifth pereiopod ( Fig. 4F View Figure 4 ) chelate.Dactyls with one row of small ovate scales. Propodal rasp with 8–9 rows of lanceolate scales, extending posteriorly nearly to midlength of propodus. Coxae weakly asymmetrical.

Males with paired, short sexual tubes consisting of slight protrusions of vas deferens partially masked by stiff, long, forwardly directed setae on coxae around margins of gonopores ( Fig. 1D View Figure 1 ); unpaired left pleopods 3–5 with exopods well developed and endopods rudimentary. Females without paired pleopods 1; pleopods 2–4 with endopods stout, exopods long, slender; pleopod 5 weakly developed, uniramous.

Uropods markedly asymmetrical.Telson ( Fig. 1E View Figure 1 ) subquadrate, with transverse indentation; anterior lobes each with tuft of long setae; posterior lobes asymmetrical, left slightly longer than right, separated by small round cleft in inverted U-shaped; posterior half of lateral margins and terminal margins armed with spines (strongest on left lobe) interspersed with small spines, terminal margins slightly convex.

Color. According to Brusca (1980: 283): “In life the carapace is orange to red, lightly speckled with pale green. Abdomen slightly lighter in color. Chelae pinkish-buff, with a light purplish tinge and speckled with pale brown and red small blotches.Eyestalks pale orange, cornea dark green to black. Walking legs pale gray with two orange bands and a bright red band on the merus (not distinct in all specimens), and vague red or purple bands on the propodus. Entire ventral side of the hermit crab pinkish-buff, except on the frontal portion, which is scarlet red”. According to Glassell (1937: 258): “In alcohol the carapace is a dark cream maculated with red and purple. Chelipeds and ambulatories with a purple tinge and red spots; a dark purple stain on hand near dactyl; propodus and dactyli of ambulatories banded with purple. Eye-stalks mottled”.

Habitat. On sandy substrates, at a depth range of 2.7 to 65 m, although mostly in less than 55 m. In waters ranging in temperatures of 13.6–25 °C, and oxygen concentration of 2.56–5.4 ml/l ( Hendrickx et al., 2006). Glassell (1937) mentioned that this species uses shells partially disintegrated by the action of bryozoans or polyps. The unidentified polypoid growth can form a large, f lexible, spiral home for the crab.

Geographic distribution. Eastern Pacific:from Tosca Point, west coast of Baja California Sur, the Gulf of California and further south on the coasts of Mexico, Costa Rica, and Ecuador ( Nobili, 1901; Rathbun, 1910; Haig et al., 1970; Hendrickx and Harvey, 1999; this report).

Morphological variations. The rostrum can be broadly rounded and unarmed or terminate in a small spine. The right cheliped in larger males (shield length> 4 mm) is longer and more slender than in females ( Fig. 3A, D View Figure 3 ).

Remarks. The study of Benedict’s (1892) types of Pagurus gladius has shown that this species clearly exhibits all diagnostic characters of Spathapagurus , and is therefore reassigned to that genus along with the other two congeneric species, the eastern Pacific Spathapagurus collinae Lemaitre and Felder, 2011 , and the western Atlantic Spathapagurus longimanus ( Wass, 1963) . Most striking in species of Spathapagurus are the grossly unequal chelipeds, the right being twice, or more, as long as the left and with a flattened, spatulate chela. Other primary characters of these species include:11 pairs of phyllobranch gills,absence of preungual process on the fourth pereiopod, and in males, paired, short sexual tubes that slightly protrude from the gonopores and are masked by forwardly directed setae.

Spathapagurus gladius comb. nov. can be distinguished from the other congeners by several characters. The relative length of the antennular and antennal peduncles are proportionally shorter in S. gladius comb.nov. than in S. collinae , but longer than in S. longimanus . The antennal acicles are proportionally longer relative to the ocular peduncles in S. gladius comb. nov. than in the two other congenerics. The length/width ratio of the right chela is proportionally greater in S. longimanus than in S. collinae and S. gladius comb. nov., reaching 3 in females and 4 in males of the former, whereas in the latter two species it reaches 2 in females and 3 in males. The armature of the palm of the right and left chelipeds of S. gladius comb. nov. is more granulose than in S. collinae and S. longimanus . The ambulatory legs in S. gladius comb. nov. have dactyls that each bear a longitudinal row of small corneous spines on the mesial surface, and the propodi and carpi have numerous spine-like tubercles on mesial and lateral surfaces, whereas in S. collinae and S. longimanus those segments are unarmed.

Phylogenetic relationships. Benedict (1892) noted, without providing any details, that his Eupagurus gladius (= Pagurus gladius ) “is closely allied to longicarpus [= Pagurus longicarpus ( Say, 1817) , a western Atlantic species] and related species”. Both P. gladius and P. longicarpus have an elongated right cheliped, a similarity that possibly led Benedict to suggest a relationship between these two species. However, the chela of the right cheliped in P. longicarpus is not f lattened as in P.gladius , and other characters exhibited by P. longicarpus also do not conform with the morphology that defines the species of Spathapagurus .

In a recent study using molecular data, Olguín and Mantelatto (2013) evaluated the phylogenetic position of species of Pagurus from the Pacific and Atlantic coasts of South America that several authors had previously placed in three groups (“provenzanoi” group, “comptus” group, and “exilis ” group) of the 11 groups that informally have been proposed for several species of this genus (see Forest and de Saint Laurent, 1968; McLaughlin, 1974; Ingle, 1985). In addition to P. gladius , the “exilis ” group originally included five other species, three of which, P. perlatus , Pagurus exilis ( Benedict, 1892) , and P. longicarpus , occur in South America. Olguín and Mantelatto (2013) added Pagurus pollicaris Say, 1817 , a western Atlantic species known from the eastern coast of the United States and the Gulf of Mexico to the Florida Keys, to the “exilis ” group. Pagurus longimanus , a western Atlantic species originally placed in the “exilis ” group, was removed from that group and reassigned by Lemaitre and Felder (2011) to Spathapagurus . Olguín and Mantelatto’s (2013) analysis of the molecular data showed that P.gladius , P. exilis , P. perlatus ,and P. longicarpus , form a stable and well supported grouping or clade. With P.gladius now reassigned to Spathapagurus based on clear morphological grounds, it seems desirable to expand Olguín and Mantelatto’s (2013) analysis to include more taxa of Pagurus (not just the ones that occur in South America) in order to ascertain more accurately the relationships of the species that have been included in the “exilis ” group.