Diogenes curvimanus
Autor: | Almón, Bruno, Cuesta, Jose A., Schubart, Christoph D., Armenia, Lisa, García-Raso, J. Enrique |
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Rok vydání: | 2021 |
Předmět: | |
DOI: | 10.5281/zenodo.6993829 |
Popis: | DIOGENES CURVIMANUS (CLÉMENT, 1874) (FIGS 5A–E, 6A–G, 7C, F, I, L) Pagurus curvimanus Clément, 1874: pl. III, fig. 1. Diogenes pugilator - Forest & Guinot, 1956: 32, fig. 3. Type material: Neotype: 1 ♂ 3.4 mm, Spain: Guadalquivir, Doñana National Park, Huelva, (MNHN-IU-2019-3214), 36°47 ′ 59.5 ′′ N, 6°23 ′ 27.3 ′′ W, sand, shallow sublitoral, 17 July 2018. To p o t y p e s: 1 ♂, 1 ♀, s a m e d a t a a s n e o t y p e, (IEOCD-BR/2581-2582); 5 ♂ (IEOCD-BR/2585). Other studied material: Belgium: Nieuwpoort-bad, 5 ♂ and 5♀ (IEOCD-BR/2611-2620), 51°09 ′ 08 ′′ N, 2°43 ′ 00 ′′ E, sand, intertidal, 29 September 2018; French Atlantic: Arcachon, 1 ♂ (IEOCD-BR/2621), 44°40 ′ 0.04 ′′ N, 1°10 ′ 34.5 ′′ E, sand, shallow sublitoral, 02 October 2018; Spain: Ribeira, Galicia, 1 ♂, 1 ♀ (ZSMA2019 0398-0399); 1 ♀ (IEOCD-BR/2603), 42°33 ′ 46.36 ′′ N, 8°59 ′ 15.07 ′′ W, sand, 4–5 m depth, 07 October 2017; Doñana N.P., Huelva, 2 ♂ (IEOCD-BR/2586), 36°48 ′ 13.6 ′′ N, 6°23 ′ 46.2 ′′ W, sand, intertidal, 09 September 2018; 2 ♂ (IEOCD-BR/2589,2595), 37°12 ′ 06.5 ′′ N, 7°01 ′ 37.0 ′′ W, sand, intertidal, 01 October 2020; 1 ♂, 1 ♀ (IEOCD-BR/2593-2594), 37°10 ′ 44.6 ′′ N, 7°20 ′ 37.3 ′′ W, sand, intertidal, 13 January 2020; 1 ♂ (IEOCD-BR/2592), 36°58 ′ 02.6 ′′ N, 6°30 ′ 32.2 ′′ W, sand, intertidal, 14 January 2020; 4 ♂ (IEOCD-BR/2587-88, 2591-91), 36°48 ′ 11.6 ′′ N, 6°23 ′ 39.3 ′′ W, sand, intertidal, 11 February 2020; Isla Canela, Huelva, 2 ♂, 1 ♀ (IEOCD-BR/2598-2600), 37°10 ′ 51.8 ′′ N, 7°20 ′ 15.3 ′′ W, sand, intertidal, 25 July 2014; Santibáñez, Cádiz, 3 ♂ (IEOCD-BR/2609-2610), 36°27 ′ 52.3 ′′ N, 6°15 ′ 21.4 ′′ W, sand, intertidal, 5 October 2015; 2 ♂ (IEOCD-BR/2605-2606), 15 October 2015; 2 ♂ (IEOCD-BR/2607-2608), 30 May 2018; Fuengirola, Málaga, 15 ♂, 7 ♀ (IEOCD-BR/2583-2584), 36°32 ′ 57.92 ′′ N, 4°36 ′ 30.89 ′′ W, sand, 4 m, 04 March 2014; La Carihuela beach, Torremolinos, Málaga, 3 ♂ (IEOCD-BR/2601-2602), 36°36 ′ 28.2 ′′ N, 4°30 ′ 13.6 ′′ W, sand, 2–4 m, 11 December 2019; Guadarranque, Algeciras, 1 ♂, 1 ♀ (IEOCD-BR/2696-2597), 36°10 ′ 49.7 ′′ N, 5°24 ′ 42.1 ′′ W, sand, 4m depth, 27 October 1995; San García, Algeciras, 1 ♂ (IEOCD-BR/2604), 36°06 ′ 17.5 ′′ N, 5°25 ′ 56.7 ′′ W, sand, 4m depth, 25 July 1996; Cabo Pino, Marbella, 1 ♂ (IEOCD-BR/2622), 36°29 ′ 05.6 ′′ N 4°45 ′ 00.4 ′′ W, intertidal, 28/04/2005. Redescription: Shield (Figs 5A, 7F) slightly broader than long; rostral lobe broadly rounded, exceeded by lateral projections that are triangular, acutely pointed, with single spine at apex; anterior margins of shield between rostral lobe and lateral projections slightly concave; anterolateral margins sloping, slightly concave, smooth; anterolateral angles rounded, with a small spine and usually another smaller below; lateral margin straight or slightly convex; posterior margin truncate; dorsal surface not vaulted, smooth, with groups of paired setae, and lateral margins with few faint transverse tuberculate ridges extending on to lateral surface of shield. Branchiostegites with dorsal margin bearing row of small spines (up to 12). Posterolateral plates not well calcified, unarmed. Ocular peduncles (including corneas) about 0.6 times as long as shield, moderately stout, slightly inflated in distal half; corneas not dilated, corneal diameter about 0.3 peduncular length; row of short, plumose setae over inner surface of the peduncles. Ocular acicles (Figs 5A, C) broad, subtriangular, concealing basal part of ocular peduncles, with slightly concave mesial margin; anterior margin sloping outwards, bearing 14–16 acute spines, decreasing in size towards outer margin, covering the entire length of the anterior margin; innermost distal spine distinctly larger. Intercalary rostriform process simple, shorter than ocular acicles (including spines), tapering acutely. Antennular peduncles (Figs 5A, B) overreaching distal corneal margin by about 0.4–0.6 length of ultimate segment, not reaching distal margin of antennal peduncle; third segment short, unarmed, broadened distally, maximum length slightly more than three times distal width, subequal in length to penultimate segment, with a tuft of setae on distodorsal margin; second segment unarmed, with tufts of plumose setae on dorsodistal and ventrodistal margins; basal segment moderately broadened distally, unarmed, except for tiny spinules on rounded ventrodistal border. Antennal peduncles (Fig. 5A) overreaching distal corneal margin by almost the entire fifth segment length (0.8–0.9); fifth segment with row of moderately short, stiff setae on ventral surface, and tuft of short setae dorsodistally, unarmed; fourth segment unarmed, with tuft of long setae on distomesial margin; third segment unarmed; second segment with distolateral outer process stout and acute, with subdistal smaller spine and sparse, short setae on lateral outer margin; distomesial spine usually small and obtuse but evident with associated tuft of long setae; mesial margin bearing some sparse, short setae; first segment unarmed. Antennal acicle short and broad, subtriangular, slightly overreaching the proximal half of fourth peduncular segment, not reaching distal margin, bearing a simple strong terminal spine plus usually six to seven spines almost equal in size along mesial margin, and tufts of setae on both mesial and lateral margins. Antennal flagellum short and robust, noticeably setose, with rows of paired setae in dorsolateral (short), ventral (medium) and ventrolateral (long) surfaces. Third maxilliped (Fig. 5D) basis unarmed; ischium with crista dentata bearing one prominent distal spine and two smaller on distal half, with additional one to two spinules on proximal half; ischium and merus with rows of scarce setae on lateral margins; carpus, propodus and dactylus with dense tufts of thick, long, simple and plumose setae at distolateral and distoventral margins, with less numerous long setae at distodorsal margin, concealing part of the segments; exopod peduncle reaching half carpal length of endopod. Male left cheliped (Figs 6A, B, 7C, F, I) much larger than right (Fig. 5E); length and shape variable in males. Dactylus shorter than palm dorsal surface, slightly arched, ending in small but strong calcareous claw, crossing tip of fixed finger; upper, inner margins defined rows of small, obtuse tubercles, upper, outer by larger acute tubercles with associated sparse setae; dorsal surface between upper, outer and inner rows slightly concave; outer surface convex, not flattened, smooth; lower margin with tufts of stout, short setae; cutting edge almost straight, with row of teeth of similar sizes; inner surface with two rows of tubercles running parallel to upper margin on upper half, reaching distal part of dactylus; proximal surface with small subacute to obtuse tubercles. Fixed finger triangular, not proximally broadened and almost smooth, except for some sparse, small tubercles on outer surface and irregular row of low tubercles on lower margin; tufts of stout, short setae on upper margin; inner surface smooth, with tufts of short setae near cutting edge and lower, inner margin. Palm narrow, about 1.5 times as long as high (max. medial length – max. height); upper margin shorter than carpus; dorsal surface of palm slightly convex, covered with evenly spaced obtuse tubercles and rows of short setae; upper, inner margin not well defined; upper, outer delineated by obtuse tubercles; outer surface glabrous, with upper part slightly concave, medially strongly and evenly convex, with numerous small, obtuse tubercles arranged as discernible longitudinal rows; lower palmar margin slightly concave in distal half, defined by row of large obtuse tubercles, and tufts of short setae; inner surface covered with low, rounded tubercles and sparse, short setae. Carpus of males long, about as long as merus and twice as long as high (Fig. 6A, B); usually longer and higher than palm; dorsal surface broadened, with upper, inner margin indistinctly delineated by row of tubercles; upper, outer well defined by curved row of subacute to obtuse tubercles, increasing in size distally, with associated medium-sized setae; outer surface with upper part slightly concave, medially strongly and evenly convex with small, obtuse evenly distributed tubercles, largest near distomesial margin; distal margin serrated; lower margin defined by row of small, obtuse tubercles; lower margin straight in distal part, short and slightly concave proximally, with tufts of short setae; inner surface covered with low, rounded tubercles and tufts of short simple setae. Merus longer than high (Fig. 6A, B, G); subovate in dorsal view; distal margin spinose throughout, with short setae; dorsal surface with small subacute tubercles of similar sizes and tufts of long, simple and plumose setae; lateral surface with evenly distributed rows of rounded tubercles, except on distal lower area; small and shallow transversal furrow subdistally on lower half, with sparse, short setae; ventrolateral margin denticulate accompanied by sparse mediumsized simple and plumose setae, slightly concave in proximal half, with larger subacute tubercle on proximal margin; mesial face with weakly calcified, u-shaped patch; subdistal transversal furrow short and faint, with small tubercles and short setae (Fig. 6G); dorsodistal margin with spines of similar size; ventrodistal margin defined by slightly larger spines with associated plumose setae. Ischium with row of small tubercles on distal margin (Fig. 6A) and row of slightly bigger tubercles on ventromesial margin (Fig. 6G). Variability. Propodus of male varies from form described to one where lower margin is strongly concave, giving a sinuous appearance. Carpus can also show on occasion a row of well-developed rounded tubercles at distomedial part. This is especially common in females and young males with short chelipeds (the reduction in size of the tubercles seems to be quicker because of the faster grow rate of male cheliped). Female left cheliped differs from male in the following features (Fig. 6C). Dactyl higher and shorter in proportion, with upper margin delineated by row of acute to subacute tubercles. Palm short and broad, slightly longer than high; outer surface covered in small, acute tubercles, with two prominent, acute tubercles distodorsally. Carpus relatively much shorter; upper, outer margin with two to three strong spines distally; outer medial surface with large, rounded tubercles increasing in size distally. Merus dorsal margin with larger, acute tubercles; lower margin defined by row of acute tubercles; lateral surface with subdistal transversal furrow more evident; mesial face with acute tubercles on ventral area and subdistal transversal furrow well defined. Right cheliped (Fig. 5E). Much shorter than left, robust (reaching one-third of carpus in neotype); dactylus and fixed finger with prominent hiatus, both terminating in small but strong calcareous claws. Dactylus (measured along mesial margins) about 2.5 times as long as palm upper margin, noticeably arched; upper, inner and outer margins defined by row of subacute tubercles with associated long setae; dorsal surface convex, broad and tuberculate; outer surface strongly convex with row of subacute tubercles parallel to the upper, outer margin; rest of outer surface with sparse, small tubercles and short, simple setae; inner surface with row of tubercles below upper, inner margin, and associated long, plumose setae; lower, inner margin with sparse setae. Palm with dorsal surface broad, short and convex; upper, outer and inner margins defined by row of subacute tubercles; outer surface broad, convex medially, with evenly spaced, subacute tubercles grading distally to acute tubercles; lower palmar margin defined by flat, setosed tubercles with long setae; fixed finger proximally broadened, outer surface slightly concave, with flat, setosed tubercles grading to acute near cutting edge; lower margin defined by flat, setosed tubercles; inner surface with low, rounded tubercles and long setae especially dense on fixed finger inner surface. Carpus dorsal surface narrow throughout; upper, outer margin defined by longitudinal row of spines and short setae; upper, inner margin defined by row of subacute tubercles with long, simple and plumose setae; additional row of small tubercles below the upper, outer margin, delimiting a shallow concave area; rest of outer surface slightly convex with low tubercles and numerous tufts of long setae; lower surface nearly smooth with sparse, short setae; inner surface weakly tuberculated, with sparse, long, plumose setae. Merus distodorsal margin with spinules and long setae; dorsal margin defined by row of small, obtuse spines and tufts of long, plumose setae; lateral surface covered with low, rounded tubercles and sparse, short setae, giving an almost smooth appearance; shallow tuberculate furrow subdistally, bearing short setae; ventrolateral margin delimited by row of small spines decreasing in size proximally, and tufts of long, plumose setae; mesial face with small, weakly calcified, u-shaped patch proximally, smooth, with ventromesial margin defined by row of minute spines and long, plumose setae. Ischium crenulated on distolateral and ventromesial margins. Second and third pereiopods slender (Figs 6D, 7L) subequal in length. Dactyl about 1.3 times as long as propodus, weakly curved; terminating in small corneous claw; upper and lower, outer margins unarmed, with rows of long thin setae, more numerous and longer on lower margin; outer surface with shallow, longitudinal sulcus medially, without setae in the proximal one-fifth, then continuing with sparse, short setae associated with sulcus; inner surface with two longitudinal rows of thin-spaced, long, stout setae adjacent to upper and lower margin, the proximal onethird with additional irregular rows of short, plumose setae placed over the sulcus. Propodus slightly longer than merus in second and third pereiopods, with upper margin defined by row of small acute tubercles (second), or faintly dentate (third), covered with long, simple setae; two rows of small tubercles running parallel to upper and lower margins, with tufts of short setae associated with tubercles; lower margin smooth, with sparse, short setae. Carpus upper margins defined by row of obtuse spines (second) or spinules (third), with dense, medium-sized, simple and plumose setae, ending in two (second) or one (third) conspicuous distal spines; rows of small, obtuse tubercles near outer upper, medial and lower surfaces with tufts of short setae in outer medial and lower surfaces; lower margin smooth with scarce short setae. Meral length about three times the maximum width; upper margin defined by small tubercles (second and third), with sparse, long, plumose setae; lower margin defined by row of low, small, subacute tubercles (second and third), with poorly developed spinules on lower distal margin (second) or without distal spines (third), and dense rows of long, plumose setae. Ischium unarmed, with long, plumose setae on distal margin. Fourth pereiopod (Fig. 6E). Dactyl with row of eight minute, spiniform setae on distal part of ventral margin. Propodus suboval, much longer than broad, with plumose, long setae along unarmed dorsal margin; propodal rasp consisting of five to six rows of corneous scales, covering distoventral part including fixed finger; rest of segments unarmed, with clumps of long, plumose setae. Fifth pereiopods. Propodus slightly shorter than merus, and about 1.7 times longer than carpus; group of subacute corneous scales in distodorsal surface of propodus and smaller ones in dactylus and fixed finger; long clumps of strong simple setae. Male unpaired left pleopods 2–5, uniramous, marginally setose. Female gonopores, paired; two to four unpaired pleopods well-developed, biramous; fifth pleopods without exopod, as in male. Telson (Fig. 6F) with shallow, median cleft, slightly asymmetrical; left posterior lobe slightly larger than right, with small spines on lateral margin, becoming blunt tubercles anteriorly; oblique terminal margin with larger spines; right posterior lobe with row of spines on less oblique terminal margin, extending on to posterior half of lateral margin. Coloration (Fig. 7C, F, I, L). Greyish white eye peduncles, with two diffuse bluish rings and a broader proximal brown triangular stain, projecting towards apex as a narrow line. Bluish ophthalmic scales. Whitish antennules, with triangular brown patch on second segment, orange in proximal two-thirds of third segment and flagellum. Antenna with greenish white or reddish peduncle, with scattered reddish brown streaks and conspicuous greenish blue spot on dorsal face of fourth segment. Carpus of the left cheliped with deep reddish orange background and an incomplete medial reddish brown ring. Orange coloured white hand with chestnut brown spot proximally, extending in two narrow lines towards middle outer surface; additional, more apparent, stain in proximal part of dactylus. Right cheliped with white background on merus and carpus, with some brown spots, and orange proximal carpal area; white palm with some brown spots in proximal upper area, which extends following central rows of tubercles. Second and third pereiopods with reddish white background colour; merus with reddish brown rings on middle (incomplete) and proximal areas. Carpus with an incomplete brown ring on medial area and small spot in the upper, proximal area. Propodus with large stain covering almost the entire proximal half, shorter in dorsal surface. Dactylus with a brown base that extends along the sulcus, sometimes faint and almost imperceptible. Habitat: Sandy beaches, in tide pools and subtidal areas up to 15 m depth, with maximum abundances in the shallow subtidal, between 4– Published as part of Almón, Bruno, Cuesta, Jose A., Schubart, Christoph D., Armenia, Lisa & García-Raso, J. Enrique, 2022, Redescription of the hermit crab Diogenes pugilator (Decapoda: Anomura) reveals the existence of a species complex in the Atlanto-Mediterranean transition zone, resulting in the resurrection of D. curvimanus and the description of a new species, pp. 1116-1146 in Zoological Journal of the Linnean Society 195 on pages 1132-1142, DOI: 10.1093/zoolinnean/zlab093, http://zenodo.org/record/6993813 {"references":["Clement MC. 1874. Description d'un pagure nouveau. Bulletin de la Societe d'Etude des Sciences Naturelles de Nimes 2: 155 - 157.","Forest J, Guinot D. 1956. Sur une collection de Crustaces Decapodes et Stomatopodes des mers tunisiennes. Bulletin Station Oceanographique de Salammbo 53: 24 - 43.","Noel P. 2016. 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