Redescription of Tomopaguroides Valdiviae (Balss, 1911) (Crustacea, Decapoda, Anomura, Paguroidea, Paguridae) with Notes on Variation and Female Morphology

Redescription of Tomopaguroides valdiviae (Balss, 1911) (Crustacea, Decapoda, Anomura, Paguroidea, Paguridae) with notes on variation and female morphology

Patsy A. MCLAUGHLIN Shannon Point Marine Center, Western Washington University, 1900 Shannon Point Road, Anacortes, WA 98221-9081B (USA) [email protected]

McLaughlin P. A. 2004. — Redescription of Tomopaguroides valdiviae (Balss, 1911) (Crustacea, Decapoda, Anomura, Paguroidea, Paguridae) with notes on variation and female morphology. Zoosystema 26 (3) : 469-481.

ABSTRACT A detailed diagnosis of the monotypic genus Tomopaguroides Balss, 1912 is presented for the first time, together with the redescription of its type species, T. valdiviae (Balss, 1911) from the lectotype, herein designated, and supple- KEY WORDS mental specimens collected in New Caledonia and the Philippine, Salomon Crustacea, Decapoda, and Fiji islands. These collection areas represent major range extensions from Anomura, the type locality in the Indian Ocean off Somalia, the only previously Paguroidea, Paguridae, confirmed locality for this species. The genus is one of only two in the family Tomopaguroides valdiviae, Paguridae characterized by the presence of only paired, modified male second lectotype, pleopods. Female attributes, heretofore unknown, are described. Significant redescription, variation, intraspecific morphological variation is reported and an anomaly attributed to anomalous regeneration. regeneration is documented.

RÉSUMÉ Redescription de Tomopaguroides valdiviae (Balss, 1911) (Crustacea, Decapoda, Anomura, Paguroidea, Paguridae) et notes sur la variation et la morphologie femelle. Une diagnose détaillée du genre monotypique Tomopaguroides Balss, 1912 est présentée pour la première fois, avec la redescription de son espèce type, T. valdiviae (Balss, 1911) à partir du lectotype, désigné ici, et de spécimens MOTS CLÉS additionnels récoltés en Nouvelle-Calédonie, aux Philippines et aux îles Crustacea, Decapoda, Salomon et Fidji. Ces zones de collecte permettent d’étendre grandement la Anomura, distribution de cette espèce jusqu’alors uniquement connue de la localité type Paguroidea, Paguridae, au large de la Somalie dans l’océan Indien. Le genre est l’un des deux seuls de Tomopaguroides valdiviae, Paguridae caractérisés par la présence de seconds pléopodes mâles modifiés, lectotype, uniquement pairs. Les attributs femelles, jusqu’ici inconnus, sont décrits. Une redescription, variation, variation morphologique intraspécifique significative est mentionnée ainsi régénération anormale. qu’une anomalie attribuée à la régénération.

ZOOSYSTEMA • 2004 • 26 (3) © Publications Scientifiques du Muséum national d’Histoire naturelle, Paris. www.zoosystema.com 469 McLaughlin P. A.

INTRODUCTION demonstrated by this deep-water taxon. Limited information on reproduction and egg size is In a preliminary report on new species of pagu- provided for the first time. roids collected during the German Tiefsee- Expedition Valdivia and the Japanese Expediton of Professor F. Doflein, Balss (1911) described MATERIALS AND METHODS the new species, Parapagurus valdiviae Balss, 1911, from two male specimens collected at All non-type specimens are from the collections Valdivia’s station 264 in the Indian Ocean off of the MNHN. One of the two syntypes, housed the coast of Somalia. The following year, in his in the ZMB, has been reexamined and, to assure full account of the Valdivia expedition, Balss stability in nomenclature, is herein designated the (1912) established the new genus Tomopaguroides lectotype. With the exception of a pair of speci- and assigned Parapagurus valdiviae as the type mens donated to the USNM, all have been returned species, again listing it as a new species. Neither to their institutions of origin. MUSORSTOM is the genus nor the species was reported again for the acronym for the joint expeditions by the nearly half a century, although Gordan (1956) MNHN and the Institut de Recherche pour le provided bibliographic references to both Para- Développement (IRD, ex-ORSTOM). Station pagurus valdiviae and Tomopaguroides valdiviae. data for the New Caledonia BATHUS 3 cruise However, the dates of publication of Tomopa- are from Richer de Forges & Chevillon (1996); guroides and T. valdiviae were incorrectly cited by data for the MUSORSTOM Philippine cruises Gordan (1956: 342) as 1926. from Forest (1981, 1985, 1989) and data for The only additional morphological information MUSORSTOM 10 and BORDAU 1 from about T. valdiviae was provided by Provenzano Richer de Forges et al. (2000a, b). The classifica- (1968). In a discussion of the phylogenetic position adopted is that of Martin & Davis (2001). tion of his new genus, Lithopagurus Provenzano, Terminology generally follows McLaughlin 1968, Provenzano compared characters of (1997), except for gill structure (cf. McLaughlin L. yucatanicus Provenzano, 1968, with those of & de Saint Laurent 1998), elements of the cara- T. valdiviae, based on Balss’ (1912) description pace (cf. McLaughlin 2003b), and abdominal and on unpublished information provided by segmentation (cf. McLaughlin et al. 2004). Michèle de Saint Laurent, Muséum national Shield length was measured from the tip of the d’Histoire naturelle, Paris (MNHN). Little data rostrum to the midpoint of the posterior margin for de Saint Laurent’s material were provided and of the shield and provided an indication of ani- her specimens have not been available for reexa- mal size. Carapace length was measured from the mination. However, certain characters attributed tip of the rostrum to the midpoint of the poste- to T. valdiviae by Provenzano (1968) do not rior margin of the carapace. agree with those of the specimens examined during the present study. It is uncertain what ABBREVIATIONS taxon de Saint Laurent may have been reporting CP beam trawl; upon. DW Warén dredge; The rediagnosis of the genus includes information ovig. ovigerous; stn station; on the mouthparts and on the morphology of the MNHN Muséum national d’Histoire naturelle, females not available at the time of the original Paris; diagnosis. The redescription of the species is USNM National Museum of Natural History, based upon one of the syntypes, herein designated Smithsonian Institution, Washington, D. C.; as the lectotype and 31 specimens from four ZMB Naturhistorisches Forschungsinstitut other widely separated geographical areas. It Museum für Naturkunde zu Berlin (for- incorporates the surprising amount of variability merly Zoological Museum of Berlin).

470 ZOOSYSTEMA • 2004 • 26 (3) Redescription of Tomopaguroides valdiviae (Crustacea, Decapoda)

SYSTEMATICS delineating anterior and posterior portions; posterior lobes separated by distinct median cleft; Family PAGURIDAE Latreille, 1802 concave terminal margins each with one to several small to moderately large spines. Genus Tomopaguroides Balss, 1912 AFFINITIES Tomopaguroides Balss, 1912: 104. — Gordan 1956: Tomopaguroides is, at least for the present, retained 342 (list). — McLaughlin 2003a: 128. in the Pylopaguropsis group of genera as proposed TYPE AND ONLY SPECIES. — Parapagurus valdiviae by de Saint Laurent-Dechancé (1966), sharing Balss, 1911, by original designation. with the other nine currently assigned genera of the DISTRIBUTION. — Indian Ocean off Somalia, group the presence of 13 pairs of gills. However, Philippine, Salomon and Fiji islands. Tomopaguroides also shares with the unique genus, Pylojacquesia McLaughlin & Lemaitre, DESCRIPTION 2001 of the family Pylojacquesidae McLaughlin Thirteen pairs of quadriserial gills. Carapace lateral & Lemaitre, 2001, a rather distinctive character lobes very narrow, partially to completely calcified. mentioned only for one other member of the Posteromedian and posterolateral carapace plates Pylopaguropsis group, Chanopagurus Lemaitre, often only faintly delineated, membranous to 2003. This is the broad separation of the anterior weakly calcified. Rostrum triangular. Ocular and posterior portions of the sternite of the third acicles well developed, each with subterminal pereopods (thoracic sternite XII of McLaughlin spine or spinule. Mandible (Fig. 1A) with incisor & Lemaitre 2001). Pilgrim (1973) described this process margins calcified, lacking denticles. sternite as being divided by a membranous hinge, Maxillule (Fig. 1B) with external lobe of endo- but this hinge is usually hardly observable. pod moderately well developed, articulated, but Despite familial differences, the overall similari- not recurved, internal lobe with one bristle. ties between Pylojacquesia and certain genera of Maxilla (Fig. 1C) with posterior lobe of scaphogna- the Pylopaguropsis group, as discussed by thite elongate, slender. First maxilliped ( Fig. 1D) McLaughlin & Lemaitre (2001), lends support to with slender exopod lacking setae on distal 0.3 of the proposition by de Saint Laurent-Dechancé outer margin. Second maxilliped (Fig. 1E) with (1966) that this group is primitive within the elongate exopod. Third maxilliped (Fig. 1F) lack- family Paguridae. ing fusion of basis and ischium; basis with two or three teeth; ischium with one accessory tooth on REMARKS well developed crista dentata; merus with dorso- As noted by Provenzano (1968), among genera of distal spine. Chelipeds grossly unequal, right the Pylopaguropsis group, Tomopaguroides appears largest. Sternite of third pereopods (Fig. 1G) most closely allied to Lithopagurus. Only species with anterior and posterior portions separated by of these two genera are characterized as having uncalcified hinge. Fourth pereopods semichelate; paired second male pleopods modified as gono- propodal rasp with two or three irregular rows of pods. This character, which sets the two genera corneous scales; no preungual process. Male with apart not only from other genera of the Pylopa- paired second pleopods modified as gonopods guropsis group but from all other genera of the (Fig. 1H); unpaired, unequally biramous left Paguridae, appears to represent a progressive evo- pleopods 3-5. Female with paired gonopores; no lutionary loss that has occurred independently in paired first pleopods; unpaired left pleopods 2-5. the Diogenidae, Parapaguridae, Pylojacquesidae, Abdomen straight, tergite of sixth pleomere (Fig. 1I) and Paguridae, as well as in the galatheid family thickened, at least weakly calcified, and unequally Porcellanidae. Among paguroids, male paired subdivided by transverse suture; uropods symme- and modified first and second pleopods are cha- trical. Telson with transverse indentations racteristic of all genera of the Pylochelidae, but

ZOOSYSTEMA • 2004 • 26 (3) 471 McLaughlin P. A.

A B C

A-D, F

E G

FIG. 1. — Tomopaguroides valdiviae (Balss, 1911); A-G, I, Philippine Islands, MUSORSTOM 1, stn 44, damaged specimen (MNHN- Pg 7065); A, mandible; B, maxillule; C, maxilla; D, first maxilliped; E, second maxilliped; F, third maxilliped; G, sternite of third pereopods; I, tergite of sixth pleomere; H, off Somalia, Valdivia, stn 264, lectotype 4.4 mm (ZMB 16470), coxae and sternite of fifth pereopods and anterior portion of abdomen (ventral view). Scale bars: 1 mm.

472 ZOOSYSTEMA • 2004 • 26 (3) Redescription of Tomopaguroides valdiviae (Crustacea, Decapoda)

appear to be retained in the Diogenidae only the Philippine Islands. MUSORSTOM 1, stn 44, genera Paguropsis Henderson, 1888, Paguristes 13°46.9’N, 120°29.5’E, 610-592 m, 24.III.1976, 7 4.3-6.3 mm, 2 specimens damaged, not sexed Dana, 1851, and Pseudopaguristes McLaughlin, or measured (MNHN-Pg 7065); stn 47, 13°40.7’N, 2002. A fourth genus, Strigopagurus Forest, 1995, 120°30.0’E, 757-685 m, 25.III.1976, 5 4.6- apparently has already lost the modified, paired 5.7 mm, 1 4.6 mm (MNHN-Pg 7066); 1 first pleopods and is undergoing evolutionary 5.6 mm, 1 5.4 mm (USNM 1016703). MUSORSTOM 2, stn 50, 13°36.7’N, 120°33.7’E, reduction in the second pleopods. This circum- 810-820 m, 27.XI.1980, 1 4.1 mm (MNHN-Pg stance of the loss among the five species assigned 7067); stn 79, 13°44.6’N, 120°31.6’E, 682-770 m, to Strigopagurus was discussed in detail by Forest 1.XII.1980, 1 ovig. 5.1 mm (MNHN-Pg 7068); (1995: 107). Paired and modified first and stn 82, 13°46.1’N, 120°28.4’E, 550 m, 2.XII.1980, 1 5.9 mm (MNHN-Pg 7069). second pleopods also are characteristic of the MUSORSTOM 3, stn CP 106, 13°47.0’N, majority of genera of the Parapaguridae, however, 120°30.3’E, 668-640 m, 2.VI.1985, 1 3.9 mm reduction and/or absence in one or both pairs has (MNHN-Pg 7070). Salomon Islands. SALOMON 1, stn CP 1858, been documented by Lemaitre (1996) for 9°37.9’S, 150°40.7’E, 435-461 m, 7.X.2001, 2 Oncopagurus Lemaitre, 1996 and Paragiopagurus 3.3, 4.0 mm (MNHN-Pg 7071). Lemaitre, 1996. Loss of male paired and modi- Fiji Islands. MUSORSTOM 10, stn CP 1343, fied pleopods is almost complete among pagurid 16°38.95’S, 177°40.84’E, 938-940 m, 10.VIII.1999, 1 ovig. 4.5 mm (MNHN-Pg 7072). genera. Only in species of Xylopagurus A. Milne BORDAU 1, stn CP 1415, 16°31’S, 179°00’W, 670- Edwards, 1880, are both paired and modified 682 m, 27.II.1999, 1 4.2 mm (MNHN-Pg 7073); first and second pleopods found. One or two stn CP 1419, 17°05’S, 178°55’W, 654-656 m, genera reportedly have rare occurrences of male 26.II.1999, 2 4.2, 5.1 mm, 1 4.6 mm (MNHN-Pg 7074); stn CP 1502, 18°21’S, paired first pleopods, and as previously indicated, 178°27’W, 640-660 m, 13.III.1999, 1 4.4 mm, only in Lithopagurus and Tomopaguroides are paired, 1 ovig. 5.3 mm (MNHN-Pg 7075). modified second pleopods observed. No male BORDAU 2, stn DW 1508, 21°02’S, 175°19’W, paired and modified pleopods have been record- 555-581 m, 31.V.2000, 1 3.1 mm (MNHN-Pg 7076). ed for Pylojacquesidae or any Lithodidae. HABITAT. — Appears to be exclusively shells of the scaphopod family Dentalidae. At least specimens of two genera, Fissidentalium Fisher, 1885 and Tomopaguroides valdiviae (Balss, 1911) Compressidentalium Habe, 1963 could be recognized (Figs 1; 2) from the Philippine collections. Parapagurus valdiviae Balss, 1911: 2, fig. 2. REDESCRIPTION Tomopaguroides valdiviae – Balss 1912: 104, figs 14, 15, pl. 9, fig. 1. Shield (Fig. 2A) somewhat vaulted; considerably longer than broad; anterior margin between ?Tomopaguroides valdiviae – Provenzano 1968: 641 (table), 642 ( key), 643 ( key) (see Remarks). rostrum and lateral projections concave; antero- lateral margins sloping; posterior margin roundly TYPE MATERIAL. — Valdivia, stn 264, 6°18’N, 49°32’E, 1079 m, 30.III.1899, 2 syntypes (ZMB 16470). truncate; dorsal surface well calcified, slightly rugose and pitted but glabrous. Rostrum acutely TYPE LOCALITY. — Indian Ocean off coast of Somalia, 6°18’N, 49°32’E, 1079 m. triangular, produced beyond bases of ocular DISTRIBUTION. — Eastern Africa, New Caledonia, acicles but often not beyond level of lateral Philippine, Salomon and Fiji islands. projections, terminating with spinule or spine. MATERIAL EXAMINED.—Indian Ocean. Off Somalia, Lateral projections well developed, acute, subacute Valdivia, stn 264, 6°18’N, 49°32’E, 1079 m, or blunt, each with prominent marginal or sub- 30.III.1899, 1 4.4 mm, lectotype, herein designated marginal spine, occasionally with both marginal (ZMB 16470). and submarginal pair of spines. New Caledonia. BATHUS 3, stn 793, 23°47’S, 169°48’E, 731-751 m, 26.XI.1993, 1 4.7 mm Ocular peduncles very short, 0.3-0.4 shield length, (MNHN-Pg 7064). stout basally and tapering to reduced corneas,

ZOOSYSTEMA • 2004 • 26 (3) 473 McLaughlin P. A.

G B

G, H

A-F, I

E I

FIG. 2. — Tomopaguroides valdiviae (Balss, 1911); A-D, H, Philippine Islands, MUSORSTOM 1, stn 44, 6.2 mm (MNHN-Pg 7065); A, shield and cephalic appendages (aesthetascs omitted); B, chela and carpus of right cheliped; C, chela and carpus of left cheliped; D, left second pereopod (lateral view); H, telson; E, Fiji Islands, BORDAU 1, stn CP 1502, ovig. 5.3 mm (MNHN-Pg 7075), distal three segments of left second pereopod (lateral view); F, I, Salomon Islands, SALOMON 1, stn CP 1858, 4.0 mm (MNHN-Pg 7071); F, distal three segments of left second pereopod (lateral view); I, chela and carpus of regenerated left cheliped; G, off Somalia, Valdivia, stn 264, lectotype 4.4 mm (ZMB 16470), telson. Scale bars: 1 mm.

474 ZOOSYSTEMA • 2004 • 26 (3) Redescription of Tomopaguroides valdiviae (Crustacea, Decapoda)

dorsomesial surfaces each often somewhat flattened generally flat, with scattered tufts of moderately or depressed in distal 0.4-0.5, dorsal or dorsomesial long setae and few small spines proximally, surfaces each with tufts or row of setae; corneas dorsomesial margin with abundance of long setae small, diameter 0.1-0.2 of peduncular length. not concealing two to four irregular rows of Ocular acicles elongate, thick, triangular, dorsally moderate to small spines, decreasing in size flattened in distal 0.3-0.5, each with tiny termi- distally; mesial face with low, sometimes spinulose nal spinule; closely-set basally, divergent distally. protuberances and tufts of moderately short setae Antennular peduncles overreaching ocular frequently forming irregular and incomplete peduncles by 0.1 to entire length of penultimate vertical rows; ventromesial margin not delineated, segment. Ultimate and penultimate segments ventral surface with few sparse tufts of very short each with few long setae dorsally. Basal segment setae. Palm 1.0-1.4 longer than carpus; dorsomesial with slender, acute spine on dorsolateral margin. margin delimited by irregular double to triple Antennal peduncles overreaching distal corneal row of moderate to small, blunt to acute spines, margins by 0.1-0.5 lengths of fourth peduncular dorsal surface with covering of long setae not segments. Fifth and fourth segments each with obscuring armature, usually one row of widely- numerous scattered setae. Third segment with spaced smaller spines adjacent to dorsomesial sparse tuft of long setae at ventrodistal margin, margin, remainder of dorsal surface with irregular occasionally concealing tiny spine or spinule. rows of small spines, most numerous laterally but Second segment with dorsolateral distal angle often forming weak, inverted V in midline, produced, terminating in bifid or simple spine dorsolateral margin with single to double row of and frequently with accessory subdistal spine, small spines, usually larger on fixed finger and dorsal surface commonly with transverse, often extending nearly to tip; flattened dorsal surface of spinulose ridge and long setae, spine, when fixed finger with numerous long setae and often present at lateral margin, usually most promi- with few small spines, tubercles or low protube- nent; dorsomesial distal angle with moderately rances; mesial and lateral faces each often with small to large spine. First segment usually with numerous small spines and long setae, at least small spine at laterodistal margin; ventral margin dorsally; ventromesial and ventrolateral margins with two to five small to moderately large spines not delimited, rounded ventral surface with few distolaterally. Antennal acicle overreaching distal very low protuberances and scattered short setae; margins of corneas by 0.3-0.7 own length and cutting edge of fixed finger with one blunt tooth often reaching to or slightly overreaching distal proximally, one broad tooth medianly and very margin of ultimate peduncular segment, slightly weakly cusped, calcareous margin distally. arcuate, with row of long, moderately dense setae Carpus approximately equal to length of merus, on mesial margin and similar row on distal 0.5 of carpal/meral articulation rotated clockwise 15- lateral margin; terminating in simple spine. 25° from perpendicular; dorsodistal margin un- Antennal flagella moderately long, but usually armed or with one to several small spines or not overreaching outstretched right cheliped; spinules, sometimes extending onto laterodistal each article with several irregularly set, moderate margin, accompanied by several long setae, dorso- to long (four to six article-length) setae. mesial margin with single to double row of Right cheliped (Fig. 2B) much larger than left, moderate to small, acute or subacute spines and operculate or nearly so. Dactyl broad, slightly few long setae, dorsal surface mesiad of midline shorter to slightly longer than dorsomesial with distal 0.3 forming slight to distinct concavity margin of palm; articulation with chela usually or basin, and usually armed with several scattered somewhat oblique; cutting edge with three large, small spines or tubercles, remainder of dorsal sur- prominent to well-worn calcareous teeth and face weakly convex and with scattered small with very short row of tiny corneous teeth adja- spines, tubercles or protuberances; dorsolateral cent to terminal corneous claw; dorsal surface margin not delimited, lateral, mesial and ventral

ZOOSYSTEMA • 2004 • 26 (3) 475 McLaughlin P. A.

surfaces each with few low protuberances and Abdomen elongate, straight; tergite of sixth pleo- sparse short setae. Merus roundly subtriangular; mere (Fig. 1I) divided into unequal halves by dorsal margin unarmed; distal margin with one deep transverse suture; proximal half subtriangular, prominent spine mesially and often one to five moderately well calcified; distal half subquadrate smaller spines dorsally; ventrolateral margin with or subrectangular, well calcified, posterolateral row of widely-spaced minute spinules or gra- angles each somewhat to prominently produced, nules; ventromesial margin with row of very terminally subacute, with or without spine. Males small or small spines, largest in distal 0.3; ventral with two-segmented paired second pleopods surface often with scattered granules, tubercles or modified as gonopods (Fig. 1H); distal segments spinules. Ischium unarmed. Coxa with small each with terminal and/or subterminal dense spine at ventromesial distal angle and frequently tufts of long setae; pleopods 3-5 with exopods one or two spines on ventrolateral distal margin. moderately well developed, endopods rudimentary, Left cheliped (Fig. 2C) short, slender; rotation of both with sparse marginal setae. Females without propodal-carpal articulation 5-30° counterclock- paired, modified first pleopods; pleopods 2-4 wise from perpendicular. Combined length of with both rami well developed, exopod slightly dactyl and palm approximately equal to indivi- longer; pleopod 5 as in males. dual lengths of carpus and merus; all segments Telson (Fig. 2G, H) with broadly rounded, sym- unarmed but with sparse rows of tufts of long metrical posterior lobes separated by narrow, setae, most numerous on dorsal surfaces of chela moderately shallow median cleft; terminal mar- and carpus. gins concave, each with one to three moderately Ambulatory legs (Fig. 2D-F) generally similar. large, often corneous-tipped spines and usually Dactyls 1.2-1.8 length of propodi, maximum three to six smaller spines adjacent to cleft, occa- dorsal-ventral height 0.1-0.2 of total length; dorsal sionally with only two or three small spines or surfaces each with row of tufts of moderately dense, tubercles adjacent to cleft. long setae; lateral and mesial faces each with faint to prominent longitudinal sulcus, flanked on COLOUR (IN LIFE) mesial face dorsally and ventrally by row of sparse Not known. No colour remaining in preserved tufts of setae, lateral face with few tufts of short specimens. setae; ventral margins each with row of 15-26 corneous spines. Propodi 1.2-1.4 longer than carpi, COMPARISON OF LECTOTYPE WITH BALSS’ each unarmed but with tufts of long setae dorsally ILLUSTRATIONS and ventrally, fewer in number ventrally. Carpi Differences in the relative sizes of the corneas and 0.5-0.8 length of meri; dorsodistal angles each with ocular peduncles, armature of the right chela and small spine, dorsal surfaces varying from each with carpus, and the positions of the left cheliped and only sparse tufts of moderately long setae to two ambulatory legs suggest that the syntype illustrated moderate to small spines in proximal half. Meri by Balss (1911: fig. 2; copied in 1912 as fig. 14) with scattered setae on dorsal and ventral margins, is not the same specimen as the syntype illustrated dorsodistal margins unarmed or each with small by the artist W. Engles (Balss 1912: pl. 9, fig. 1), spine. Ischia unarmed but with sparse setae on ven- although, these differences might simply reflect tromesial margins. Coxae each usually with one to “artistic license” in the latter figure. The shapes of five small spines on ventromesial distal margin. the ocular peduncles and corneas of the examined Fourth pereopods with propodal rasp consisting of syntype, herein designated as the lectotype, two or three rows of corneous scales; dactyl with would suggest that it is this specimen that was small terminal claw, no preungual process. Anterior depicted in Balss’ line drawing (1911: fig. 2; lobe of sternite of third pereopods subtriangular to 1912: fig. 14); however, his measurement of 9 mm subquadrate, unarmed or with one to several small for carapace length corresponds well with the spines marginally, usually concealed by long setae. specimen illustrated in plate 9, figure 1, and is

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considerably greater than the actual 7.3 mm mea- but those of the lectotype are longer than those sured carapace length of the lectotype. shown in either of Balss’ figures. The illustrated The supposition that Balss’ (1911: fig. 2; 1912: antennal flagellum (Balss 1911: fig. 2; 1912: fig. 14) line drawing is of one syntype, while the fig. 14) shows each article with a pair of setae of artist’s rendition (Balss 1912: pl. 9, fig. 1) is of equal length; whereas, the specimen illustrated in the other is given additional credence by the fact Balss’ (1912) plate 9, figure 1 shows no setation that only Balss’ (1911: fig. 2) figure of the anterior at all. The articles of the antennal flagella of the portion of the cephalothorax and appendages lectotype each has several irregularly set, moderate appeared in his initial description of Parapagurus to long setae. The armature of the right cheliped valdiviae. It was after that preliminary publication as portrayed by Balss (1911: fig. 2; 1912: fig. 14, that Balss apparently realized that parapagurids pl. 9, fig. 1) does not agree with that of the lecto- did not have 13 pairs of gills and that the gill type. Although a spinose dorsomesial margin of number and other characters seemed to relate this the chela is depicted relatively accurately, the dorsal species to Tomopaguropsis Alcock, 1905. Although surface of the chela of the lectotype has a covering his specific description remained identical with of small spines, but is shown with only setae in that of Parapagurus valdiviae, in Balss’ (1912: 104) the illustrations. In the lectotype, the spines on generic diagnosis he expressed his uncertainty as the dorsolateral margin extend nearly to the to whether the paired gonopods were on the proximal margin of the chela as illustrated in second or first pleonal somite, but suspected the Balss’ (1912) plate 9, figure 1, but not figure 14. second. For Tomopaguroides valdividae, Balss The spines on the dorsomesial margin of the carpus (1912: fig. 15) included the line drawing of the of the lectotype extend the entire length of that ventral view of the thorax and anterior portion of margin, rather than only half the distance as in the abdomen together with his earlier figure of both figures 14 and plate 9, figure 1. On neither the cephalothorax and appendages. In the later pair of ambulatory legs is a dorsodistal carpal publication Balss also added full plates depicting spine indicated on either of Balss’s figures; however, several species drawn by artists. Neither the the carpi of the lectotype each has a small but ventral view of the thorax (fig. 15) nor the illus- distinct dorsodistal spine. trated entire animal (pl. 9, fig. 1) of T. valdiviae correspond well with the lectotype. COMPARISON OF THE LECTOTYPE WITH PACIFIC There are several morphological differences SPECIMENS between the lectotype and the illustrated Although there is variation in rostral development, specimen(s) that may reflect some factual varia- the lectotype agrees well with the Pacific specimens, tions, but also clearly indicate certain incorrect if not with Balss’ (1911: fig. 2; 1912: fig. 14, observations and inaccuracies in the original pl. 9, fig. 1). Similarly, the ratios of ocular pedun- figures. The rostra illustrated in both the line dra- cular length to shield length and corneal diameter wing and the plate figure, for example, are much to peduncular length are within the ranges of the shorter than the tips of the lateral projections and Pacific populations. The antennular and antennal lack a terminal spine. And in his generic diagno- peduncles of the lectotype both overreach the distal sis Balss (1912: 104) specifically noted a similari- corneal margins by approximately 0.5 length of ty between the rostrum of Tomopaguroides the penultimate segments, lengths also with in valdiviae and the short rostra of the two species of the ranges exhibited by the Pacific specimens. Tomopaguropsis known at the time. However, the The produced dorsodistal angle of second segment rostrum of the lectotype is more acutely triangu- of the antennal peduncle lacks an accessory spinule lar than illustrated and has a prominent terminal in the lectotype and the ridge on the dorsal surface spine that extends the rostral length to only of the segment is not spinulose; however, the slightly less than those of the lateral projections. presence of an accessory spine or spinules on the The antennal acicles were not described by Balss, ridge was found to be variable in the Pacific

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populations. The length of the antennal acicles of samples from the Philippines Islands with a ratio the lectotype are well within the range seen in the of four males to every one female. Of those four Pacific specimens. As noted above, the articles of females collected during cruises in Philippine the antennal flagella of the lectotype each have waters in 1976, 1980 and 1985, only one was an several irregularly set, moderate to long setae like ovigerous female taken in March. In contrast, the those of its Pacific counterparts. The spination sex ratio in the Fiji area was evenly balanced, with on the dorsal surface of the right chela common four males and four females recovered. Of the to the lectotype and the Pacific specimens was females, two of the four were ovigerous, with one apparently unnoticed by Balss. collected in February and the second in August. Comparisons of the ratios of dactylar length to The eggs carried by females from both areas were height and dactylar to propodal lengths of the large and relatively few in number, although second left pereopod of the lectotype have shown because of the ease with which eggs could be that both ratios fall well within the ranges found in dislodged from the pleopods, an actual egg count the Pacific populations, i.e. 0.1 and 1.6 respectively. was not reliable. Egg diameters measured Similarly, the longitudinal sulci on the lateral faces between 1.1 and 1.4 mm at early stages of deve- of the dactyls, although weak, are within the range lopment (non-eyed). of variation seen among the Pacific representatives. However, in the number of spines on the ventral VARIATIONS margins of the dactyls, only 15 are present in the As may be ascertained from the character variabi- lectotype, whereas the range is 16-26 in the Pacific lity incorporated into the redescription, specimens. The carpi of the ambulatory legs of the Tomopaguroides valdiviae is another hermit crab lectotype each have the typical small dorsodistal taxon with considerable morphological plasticity. spine as previously noted, but dorsal carpal The differences between specimens collected in surfaces show only low protuberances and sparse the Philippines and those from Fiji initially setae. Carpal armature in Pacific specimens varied suggested two distinct taxa. Fortunately, from only sparse setae to two moderate to small MUSORSTOM collections from a broad area of spines in the proximal half of the segment. In the the western Pacific were available for simulta- lectotype, the coxae of these appendages are armed neous examination and these contained a suffi- with two or three small spinules on the second per- cient number of specimens to make an accurate eopods, but zero to one on the third. The fourth assessment of intraspecific variation possible. pereopods of the lectotype have two rows of cor- Most striking were the variations seen in the neous scales, and the subtriangular anterior lobe of length/depth ratios of the ambulatory dactyls and the sternite of the third pereopods has two small, the armature of the carpi and meri of those subacute terminal spinules. appendages (Fig. 2D-F). The dactyls of the Balss (1912: fig. 15) illustrated the paired second second and third pereopods of the lectotype and male pleopods as being crossed. These appendages Philippine specimens varied from 1.5 to 1.8 times in the lectotype (Fig. 1H) are not crossed, nor were the length of the propodi with maximum dorsal- they in any of the males of the Pacific populations. ventral height of approximately 0.1 of the length, The telson of the lectotype (Fig. 2H) lacks the small whereas, some specimens from Fiji had much spinules seen on telsons of some, but not all Pacific shorter dactyls (1.2-1.3 length of propodi) with specimens. Reexamination of the lectotype has the maximum dorsal-ventral height 0.2 of the confirmed the identifications of the Pacific popula- length. The carpi of the lectotype and most of the tions as belonging to Tomopaguroides valdiviae. Philippine specimens carried only a dorsodistal spine, although setae-bearing protuberances were SEX RATIO AND REPRODUCTION present on the lectotype and occasionally could A single male was collected in New Caledonian be discerned in the posterior halves of the dorsal waters. Males were much more common in the margins of the Philippine specimens. In contrast,

478 ZOOSYSTEMA • 2004 • 26 (3) Redescription of Tomopaguroides valdiviae (Crustacea, Decapoda)

in the majority of specimens from Fiji, the dorsal its position was a fully developed, albeit reversed, margins of the carpi had at least one and more right cheliped (Fig. 2I). On the animal’s right often two well developed spines on the posterior side was a smaller right cheliped, correct in form, half. The ambulatory meri of all of the Philippine but with appreciably weaker spination on the specimens had unarmed dorsodistal margins as chela and carpus than was present on the cheliped did the lectotype, while the dorsodistal margins on the left side. In all other respects the specimen of these segments in some, but not all Fiji speci- appeared normal. mens were each armed with a small spine. Addi- tional significant variations were observed in the AFFINITIES lengths of the antennular peduncles (shorter in There is a very cogent, albeit superficial, resem- the Fiji specimens) and lengths of the ocular blance between females of Bathypaguropsis microps peduncles (longer in the Fiji specimens). That (Balss, 1911) and females of Tomopaguroides these differences reflected within population valdiviae. In addition to sharing the general variations rather than specific differences were characters of the Pylopaguropsis group of genera confirmed, in part, by the two specimens from in terms of gills, both have very short ocular the Salomon Islands and the single specimen peduncles with reduced corneas, thick, dorsally from New Caledonia. Although the larger of the somewhat flattened, triangular ocular acicles, two Salomon Islands males had abnormal cheliped grossly unequal chelipeds, relatively unarmed development, both specimens lacked posterior ambulatory legs with somewhat laterally spines on the dorsal margins of at least one or two compressed dactyls, and broadly rounded poste- of the four pereopodal carpi; the meri all were rior terminal telsonal lobes. And both species unarmed. The New Caledonia male lacked poste- occur sympatrically. Balss (1911) described rior carpal and dorsodistal spines on the segments T. valdiviae (as Parapagurus) and B. microps (as of all pereopods. The lengths of the dactyls and Eupagurus) from the same Valdivia station off the the maximum dorsal-ventral heights were inter- coast of Somalia. In the present study, a single mediate between those of the Philippine and Fiji female of B. microps was collected at MUSORS- specimens. The lengths of the ocular peduncles of TOM Philippine station 44, with nine specimens the New Caledonia and Salomon Islands males of T. valdiviae. Had McLaughlin (2003c) not approximated those of the Philippine specimens, reexamined the male holotype of B. microps and while the lengths of the antennular peduncles of determined that Balss (1911, 1912) had erred in the Salomon Islands males were comparable to his report of the gill number and lamellar type, it those of the Fiji material. Further evidence was is very possible that the female specimen from provided by the examination of one Philippine MUSORSTOM Philippine station 44 might specimen in too poor condition to be included in incorrectly have been described as a new species the species’ evaluation. That specimen had a of Tomopaguroides. small dorsoproximal spine on each pereopodal carpus. Variations in the prominence of the lon- REMARKS gitudinal sulci on the dactyls of the ambulatory As previously pointed out, character states attri- legs were also noted; however, these variations buted to Tomopaguroides valdiviae by Provenzano did not seem to be zoogeographically influenced. (1968) do not agree, in some particulars, with From the information presently available, these those of the specimens included in this investiga- variations do not appear to be correlated to either tion. Most important is the reported presence of size or sex. rudimentary paired first pleopods in females, and The abnormal cheliped development of male referred to by Provenzano as gonopods. None of specimen from the Salomon Islands previously the eight females examined in the present study mentioned involved a presumed regeneration exhibited any development of first pleopods. anomaly. That individual had no left cheliped. In Therefore, if the female specimen(s) that de Saint

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Laurent described through Provenzano (1968) Prof. Philippe Bouchet and Dr Alain Crosnier (both had rudimentary paired first pleopods, it can only Département Systématique et Évolution, MNHN), be assumed that she was incorrect in her assess- for financial assistance and accommodations ment of the species as T. valdiviae. Paired and during her stay at the MNHN and for putting modified female first pleopods do occur in five this important collection at her disposal. genera of the Pylopaguropsis group, Bythiopagurus Identifications of the scaphopod genera were McLaughlin, 2003, Chanopagurus, Munidopa- made by George Holm, Richmond, British gurus A. Milne-Edwards & Bouvier, 1893, Columbia, Canada. This is a scientific contribu- Pylopaguropsis Alcock, 1905, and Xylopagurus tion from the Shannon Point Marine Center, A. Milne Edwards, 1880, but only in Chano- Western Washington University. pagurus atopos Lemaitre, 2003, are the corneas reduced as they are in Tomopaguroides valdiviae. Since females of C. atopos have a single left gono- REFERENCES pore while those of T. valdiviae have paired BALSS H. 1911. — Neue Paguriden aus den gonopores, it is unlikely that de Saint Laurent Ausbeuten der deutschen Tiefsee-Expedition would have confused those two taxa. Valdivia und der japanischen Expedition Prof. Provenzano’s (1968: 641) table 1 indicated that Dofleins. Zoologischer Anzeiger 38 (1): 1-9. dorsal spines were absent from the carpi of both BALSS H. 1912. — Paguriden, in CHUN C. (ed.), Wissenschaftliche Ergebnisse der deutschen Tiefsee- pairs of ambulatory legs of T. valdiviae, but whether Expedition Valdivia 1898-1899 20 (2). Gustav his information was gleaned from de Saint Laurent’s Fischer, Jena: 85-124. unpublished information or Balss’ (1911: fig. 2; FOREST J. 1981. — Report and general comments, in 1912: fig. 14, pl. 9, fig. 1) was not specified. Résultats des campagnes MUSORSTOM. 1. Philippines (18-28 mars 1976). Mémoires However, the lectotype and all of the 31 Pacific ORSTOM 91: 9-50. specimens in the current collection had at least a FOREST J. 1985. — The MUSORSTOM II dorsodistal spine on each carpus. One specimen Expedition (1980) report and list of stations, in from the Philippines and all specimens from Fiji Résultats des campagnes MUSORSTOM I et II. Philippines (1976, 1980), volume 2. Mémoires du and the Salomon Islands also had at least one, and Muséum national d’Histoire naturelle sér. A, 133: often two, dorsoproximal spines on one or more 7-30. carpi. Differences in other attributes, such as ros- FOREST J. 1989. — Compte rendu de la campagne tral length and ocular acicle development, fall well MUSORSTOM 3 aux Philippines (31 mai- 7 juin 1985). Report on the MUSORSTOM within the ranges of variation seen for T. valdiviae. 3 Expedition to the Philippines May 31st-June Provenzano cited only one measurement of speci- 7th 1985), in FOREST J. (ed.), Résultats des cam- men size and that was the 9 mm carapace length pagnes MUSORSTOM, vol. 4. Mémoires du Muséum national d’Histoire naturelle sér. A, 143: given by Balss (1911, 1912). As previously indica- 9-23. ted, the lectotype has a shield length of 4.4 mm FOREST J. 1995. — Crustacea Decapoda Anomura : and a carapace length of 7.3 mm. Specimens in the révision du genre Trizopagurus Forest, 1952 Pacific series ranged in shield length from 3.1 to (Diogenidae), avec l’établissement de deux genres nouveaux, in CROSNIER A. (ed.), Résultalts des 6.3 mm. Shield length in T. valdiviae varies from campagnes MUSORSTOM, vol. 13. Mémoires du 0.6 to 0.7 of carapace length, thus only the largest Muséum national d’Histoire naturelle 163: of these specimens approached the size specified by 9-149. Balss and reflected in his illustration (1912: pl. 9, GORDAN J. 1956. — A bibliography of pagurid crabs, exclusive of Alcock, 1905. Bulletin of the American fig. 1) of the entire animal. Museum of Natural History 108: 253-352. LEMAITRE R. 1996. — Hermit crabs of the family Parapaguridae (Crustacea: Decapoda: Anomura) Acknowledgements from Australia: species of Strobopagurus Lemaitre, 1989, Sympagurus Smith, 1883 and two new Dr Charles Oliver Coleman (ZMB) kindly arranged genera. Records of the Australian Museum 48: the loan of the syntype. The author is indebted to 163-221.

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MARTIN J. W. & DAVIS G. E. 2001. — An updated Fiction? Part II. Evidence from larval, megalopal classification of the Recent Crustacea. Natural and early juvenile development. Contributions to History Museum of Los Angeles County, Science Series Zoology 73: 165-205. 39: 1-124. PILGRIM R. L. C. 1973. — Axial skeleton and muscu- MCLAUGHLIN P. A. 1997. — Crustacea Decapoda: lature in the thorax of the hermit crab, Pagurus Hermit crabs of the family Paguridae from the bernhardus (Anomura: Paguridae). Journal of the KARUBAR cruise in Indonesia, in CROSNIER A. & Marine Biological Association of the United Kingdom BOUCHET P. (eds), Résultats des campagnes 53: 363-396. MUSORSTOM, vol. 16. Mémoires du Muséum PROVENZANO A. J. JR. 1968. — Biological investiga- national d’Histoire naturelle 172: 433-572. tions of the deep sea. 37. Lithopagurus yucatanicus, MCLAUGHLIN P. A. 2003a. — Illustrated keys to the a new genus and species of hermit crab with a families and genera of the superfamily Paguroidea distinctive larva. Bulletin of Marine Science 18: (Crustacea: Decapoda: Anomura), with diagnoses 627-644. of the genera of Paguridae, in LEMAITRE R. & RICHER DE FORGES B. & CHEVILLON C. 1996. — Les TUDGE C. C. (eds), Biology of the Anomura. campagnes d’échantillonnage du benthos bathyal en Proceedings of a symposium at the Fifth Nouvelle-Calédonie en 1993 et 1994 (BATHUS 1 à 4, International Crustacean Congress, Melbourne, SMIB 8 et HALIPRO 1), in CROSNIER A. (ed.), Australia, 9-13 July 2001. Memoirs of Museum Résultats des campagnes MUSORSTOM, vol. 15. Victoria 60: 111-144. Mémoires du Muséum national d’Histoire naturelle 168: MCLAUGHLIN P. A. 2003b. — A new genus and 33-53. species of hermit crab (Decapoda: Anomura: RICHER DE FORGES B., NEWELL P., SCHLACHER- Paguroidea: Paguridae) from the seamount region HOENLINGER M., SCHLACHER T., NATING D., of southeastern Tasmanian. Memoirs of Museum CÉSA F. & BOUCHET P. 2000a. — La campagne Victoria 60: 229-236. MUSORSTOM 10 dans l’archipel des Îles Fidji. MCLAUGHLIN P. A. 2003c. — Reassignment and Compte rendu et liste des stations, in CROSNIER A. redescription of “Eupagurus” microps Balss, 1911 (ed.), Résultats des campagnes MUSORSTOM, (Crustacea, Decapoda, Paguridae), with notes on vol. 21. Mémoires du Muséum national d’Histoire Bathypaguropsis kuroshioensis (Miyake, 1978). naturelle 184: 9-23. Zoosystema 25 (4): 635-642. RICHER DE FORGES B., BOUCHET P., DAYRAT B., MCLAUGHLIN P. A. & LEMAITRE R. 2001. — A new WARÉN A. & PHILIPPE J.-S. 2000b. — La campagne family for a new genus and new species of hermit BORDAU 1 sur la ride de Lau (Îles Fidji). Compte crab of the superfamily Paguroidea (Decapoda: rendu et liste des stations, in CROSNIER A. (ed.), Anomura) and its phylogenetic implications. Résultats des campagnes MUSORSTOM, vol. 21. Journal of Crustacean Biology 21: 1062-1076. Mémoires du Muséum national d’Histoire naturelle MCLAUGHLIN P. A. & SAINT LAURENT M. DE 1998. — 184: 25-38. A new genus for four species of hermit crabs for- SAINT LAURENT-DECHANCÉ M. DE 1966. — merly assigned to the genus Pagurus Fabricius Remarques sur la classification de la famille des (Decapoda: Anomura: Paguridae). Proceedings of the Paguridae et sur la position systématique Biological Society of Washington 111: 158-187. d’Iridopagurus de Saint Laurent. Diagnose MCLAUGHLIN P. A., LEMAITRE R. & TUDGE C. C. d’Anapagrides gen. nov. Bulletin du Muséum natio- 2004. — Carcinization in the Anomura – Fact or nal d’Histoire naturelle (2) 38: 257-265.

Submitted on 29 July 2003; accepted on 13 November 2003.

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