Clibanarius clibanarius ( Herbst, 1791 )

Komai, Tomoyuki, Reshmi, Rema, Rahayu, Dwi Listyo & Kumar, Appukuttannair Biju, 2019, Redescription of the little known hermit crab, Clibanarius clibanarius (Herbst, 1791) (Decapoda: Anomura: Diogenidae), based on material from southern India and type material, Zootaxa 4555 (3), pp. 372-384 : 373-382

publication ID	https://doi.org/ 10.11646/zootaxa.4555.3.6
publication LSID	lsid:zoobank.org:pub:2D50E43C-9430-49FE-BC1E-0965C574ED8F
DOI	https://doi.org/10.5281/zenodo.5930952
persistent identifier	https://treatment.plazi.org/id/038587D5-FFE9-3414-C2DF-FC06FB8FF807
treatment provided by	Plazi
scientific name	Clibanarius clibanarius ( Herbst, 1791 )
status

Treatment

Clibanarius clibanarius ( Herbst, 1791) View in CoL

( Figs. 1–7 View FIGURE 1 View FIGURE 2 View FIGURE 3 View FIGURE 4 View FIGURE 5 View FIGURE 6 View FIGURE 7 )

Cancer clibanarius Herbst, 1891: 20 , pl. 23, fig. 1 (type locality: not indicated).

Pagurus clibanarius .—Latreille 1803: 167 (no new locality). — H. Milne Edwards1836: 276 (mer d’Asie); 1837: 227 (no new locality); 1848: 62 (no new locality).

Pagurus (Clibanarius) clibanarius .—Hilgendorf 1878: 820 (no new locality).

Clibanarius vulgaris Dana, 1851: 462 View in CoL (invalid replacement name).

Clibanarius clibanarius View in CoL .— Henderson 1893: 423 (Madras).— Rathbun 1900: 306 (no new locality).— Alcock 1905: 43, pl. 4, fig. 1 ( Pondicherry, Mouth of Hooghly).— Khan & Natarajan 1984: 4, fig. 2 (Porto Novo).— Thomas 1989: 62, fig. 2p, r, pl. 1F (Cuddalore; Kakinada).— Sakai 1999: 10, pl. 2C (type material).— Osawa & Fujita 2006: 3, table 1 (list, no new locality).— McLaughlin et al. 2010: 19 (list; no new locality).

Not Pagurus clibanarius .— Quoy & Gaimard 1824: 529, pl. 78, fig. 1. = Clibanarius taeniatus H. Milne Edwards, 1848 View in CoL .

Questionable records

Clibanarius vulgaris View in CoL .— Stimpson 1858: 247 ( Hong Kong); 1907: 209 ( Hong Kong).— Miers 1880: 375 (Borneo).— Walker 1887: 112 ( Singapore).— Stebbing 1908: 21; 1910: 352 (Natal, South Africa).

Clibanarius clibanarius View in CoL .— Krauss 1843: 56 ( South Africa).–– Gee 1925: 159 ( China).–– Estampador 1937: 501 (Taytay, Palawan); 1958: 48 (Taytay, Palawan).— Yap-Chiongco 1938: 190 (Taytay, Palawan).— Barnard 1950: 433 (no new locality).— Rahayu 1996: 382 (list; no new locality).— Sha et al. 2015: 62, fig. 2-23 (no new locality).

Type material. Lectotype (designated by Sakai 1999; photograph): ZMB Herbst 2493, sex not determined (cl ca. 39.0 mm).

Paralectotypes (photographs): ZMB Herbst 2494, 1 specimen (cl 37.5 mm) ; ZMB Herbst 2493, 1 specimen (cl 33.5 mm) ; ZMB Herbst 2495a, 1 specimen (cl 39.0 mm) .

Other material examined. DABFUK/AR-AN 31–32, 2 males (sl 13.1, 12.4 mm), Sakthikulangara (08°55.59’N, 76°32.30’E), Kerala State, southwestern India, 12 July 2012, bycatch of commercial trawler, coll. R. Reshmi; DABFUK/AR-AN 33, 1 male (sl 12.8 mm), Vizhinjam (08 ° 22.33’N, 76 ° 59.28’E.), Kerala State , 23 December 2012, beach seine, coll. A. Biju Kumar GoogleMaps ; DABFUK/AR-AN 34, 1 female (sl 12.4 mm), Neendakara (08 ° 56.15’N, 76 ° 32.20’E), Kerala State , 12 July 2012, bycatch of commercial trawler, coll. R. Reshmi GoogleMaps ; DABFUK/ AR-AN 35, 1 male (sl 11.9 mm), Ponnani (10 ° 46.51’N, 75 ° 54.59’E), Kerala State , 6 April 2011, bycatch of commercial trawler, coll. R. Reshmi GoogleMaps ; DABFUK/AR-AN 106, 1 female (sl 15.6 mm), same data as DABFUK/AR- AN 34 GoogleMaps .

Description. Shield ( Fig. 1A View FIGURE 1 ) as long as broad; anterior margin between rostrum and lateral projections concave; anterolateral angles blunt, but distinctly delineated; lateral margins with few small spines or tubercles; posterior margin roundly truncate; dorsal surface with scattered blister-like tubercles and tuft of short to moderately long setae; paragastric grooves faint. Posterior carapace with scattered tufts of long yellowish setae, most numerous on lateral surface.

Ocular peduncles ( Fig. 1A View FIGURE 1 ) equal in length, slender (more than 8 × as long as corneal width), about 0.8 × as long as shield, cylindrical, inflated basally; cornea small, not dilated, narrower than base of ocular peduncle; dorsal surface with scattered short to moderately long setae. Ocular acicles triangular, with 3 or 4 distal spines; mesial margin slightly convex, lateral margin slightly concave; narrowly separated basally.

Antennular peduncles ( Fig. 1A View FIGURE 1 ) overreaching distal corneal margins by 0.3 length of ultimate article. Ultimate article about half as long as shield, about 1.5 × length of penultimate article, with row of short to moderately long setae on dorsal surface. Penultimate article also with tufts of moderately long setae on dorsal surface. Basal article short, statocyst lobe without spine.

Antennal peduncles ( Fig. 1A View FIGURE 1 ) not reaching corneal bases. Article 5 slightly flattened, with scattered short setae on surfaces. Article 4 with small spine on dorsodistal margin. Article 3 with small ventromesial distal spine. Article 2 with small spines on dorsomesial distal angle and distolateral angle, respectively. Article 1 with prominent spine on ventrodistal margin. Antennal acicle reaching slightly beyond base of article 5, terminating in simple or bifid spine; mesial margin with row of 5 or 6 small spines and tufts of short setae; lateral margin unarmed, with few tufts of short setae. Flagellum as long as carapace, with several minute setae on distal margin of each article.

Maxilliped 3(not figured) typical of genus. Basis and ischium incompletely fused, basis with 3 minute corneous tipped denticles on ventromesial margin. Ischium with crista dentata well-developed, consisting of row of subacute corneous teeth, without accessory tooth. Merus and carpus unarmed on dorsodistal margins. Exopod broad, uncalcified in lateral half, mesial surface with shallow concavity in distal half; flagellum well developed.

Chelipeds ( Fig. 2 View FIGURE 2 ) subequal. Left cheliped with dactylus 1.4 × as long as palm; dorsal surface with 2 longitudinal rows of corneous-tipped spines becoming smaller and corneous distally and row of tufts of short stiff setae flanked by spine rows; dorsomesial margin with row of small corneous-tipped spines becoming smaller and corneous distally and tufts of short setae; mesial surface with 2 longitudinal rows of tufts of short setae and 1 or 2 small spines proximally; ventral surface bluntly carinate; cutting edge with 2 closely spaced calcareous teeth proximal to midlength, distal half bordered by darkly pigmented corneous plate leading to tip. Palm with 5 moderately large spines on dorsomesial margin; laterally sloping dorsal surface with numerous, scattered small to moderately large spines extending onto fixed finger in 2 irregular rows, all spines corneous-tipped, and with tufts of short to long setae arising at bases of spines; nearly perpendicular lateral surface with sparse low protuberances each with setal tuft, becoming spiniform with corneous tip on fixed finger; mesial surface with some low protuberances and tufts of setae; ventral surface with low protuberances and tufts of setae, armature becoming smaller and spiniform with corneous-tip in fixed finger; cutting edge with 4 subacute calcareous teeth in proximal half, bordered by darkly pigmented corneous plate leading to tip. Carpus slightly shorter than palm, about halflength of merus; dorsomesial margin with row of 3 large, corneous-tipped spines; dorsal surface sloping to lateral surface, with scattered small to moderately large spines and tufts of long setae; mesial surface smooth, with few tufts of setae; distal margin with few minute denticles. Merus fairly compressed laterally, subtriangular in dorsal view; dorsal margin carinate, with row of low protuberances and tufts of short to moderately long setae; lateral surface with scattered small tubercles and sparse tufts of short setae, ventrodistal angle with 2 small, corneoustipped spines and ventrolateral margin with row of small tubercles; mesial surface nearly flat, smooth, with subdistal, low transverse ridge, and with tiny, low tubercles dorsally and few tufts of short setae, ventromesial margin with row of small acute or subacute spines; ventral surface with some small tubercles, but without prominent spine or tubercle. Ischium with row of tiny tubercles on ventromesial margin; ventral surface few, small to large, blister-like tubercles.

Right cheliped similar to left cheliped in armature and ornamentation.

Pereopods 2 and 3 ( Figs. 3 View FIGURE 3 , 4 View FIGURE 4 ) relatively stout, second overreaching tip of chelipeds by at least half length of dactyli. Dactyli about 1.3 (second) or 1.4 (third) × as long as propodi, 6.2–6.4 × as long as wide, gently curved ventrally in lateral view, nearly straight in dorsal view, each terminating in darkly pigmented corneous claw; dorsolateral margins bluntly carinate, with row of tufts of moderately long stiff setae mesial to dorsolateral carina; lateral surfaces each with faint longitudinal sulcus medially and 3 rows of tufts of short to moderately long, stiff setae; mesial surfaces each with blunt longitudinal carina just dorsal to shallow median sulcus, and with 3 rows of setal tufts; ventral margins each with row of 7–10 small corneous spines in distal half. Propodus of left pereopod 3 stouter than those of pereopods 2 and right pereopod 3; dorsal surfaces each with 1 small dorsodistal spine followed by 1 subdistal spine and 2 rows of low, setiferous protuberances (second) or only 2 rows of low, setiferous protuberances (third); distal margins each with few minute spinules; lateral surfaces each with 3 longitudinal rows of setal tufts, that of left third bearing some setiferous, low protuberances near ventral margin; mesial faces each with 2 rows (second) or 3 rows (third) of setal tufts; ventral surfaces each with 1 row of tufts of long stiff setae, low protuberances more pronounced in left third. Carpi short, dorsal margins each armed with row of spines increasing in size distally (second) or armed only with dorsodistal spine (third); and with row of tufts of short to moderately long stiff setae; lateral surfaces convex, with setal tufts arranged in longitudinal rows; mesial faces nearly glabrous, only with few short setae; ventral surfaces each with prominent tuft of setae distally and 1 additional tuft of setae at midlength. Meri strongly compressed laterally; dorsal margins each with low, setiferous protuberances; lateral and mesial surfaces with tufts of very short setae adjacent to dorsal and ventral margins, largely glabrous; ventral margins each with 1 distolateral spine and row of minute spines or granules extending onto ventromesial distal margin (second) or followed by row of low, setiferous protuberances (third). Ischia each with large, low protuberances on lateral surface near ventral margin.

Pereopods 4 ( Fig. 1B View FIGURE 1 ) semichelate, articles bearing tufts of long stiff setae particularly on dorsal margins, lateral surfaces with blister-like protuberances of various size. Dactylus nearly straight, overreaching tip of fixed finger, terminating in a small corneous claw; lateral surface with row of minute corneous spines ventrally; ventral surface broad, nearly flat. Propodal rasp well developed, encompassing three-fourths of ventrolateral surface, consisting of numerous small corneous scales. Carpi each with 2 small spines on dorsal surface distally.

Pereopods 5 chelate.

Thoracic sternite 6 ( Fig. 1C View FIGURE 1 ) with anterior lobe subsemicircular, bearing numerous long setae anteriorly.

Pleon dextrally twisted. Pleonite 6 ( Fig. 1D View FIGURE 1 ) approximately twice as long as wide, divided into two parts by distinct transverse groove across slightly posterior to midlength; lateral indentation shallow; posterior part with deep median longitudinal groove, posterolateral margin.

Males with 4 unpaired left pleopods; each exopod well developed. Endopods reduced, about half-length of exopods. Female with 4 biramous unpaired pleopods. Uropods greatly asymmetrical.

Telson ( Fig. 1E, F View FIGURE 1 ) with faint lateral indentations; posterior lobes unequal (left distinctly larger than right), fringed with short to moderately long setae, separated by very small median cleft; terminal margins each with 6–10 corneous or corneous-tipped spines or spinules increasing in size laterally.

Coloration in life. Shield mottled maroon, margins cream; posterior carapace pale brown, branchiostegite maroon; lateral parts of posterior carapace dark brown. Ocular peduncles generally brown, each with dark brown median longitudinal stripe. Antennular and antennal peduncles generally reddish brown. Chelipeds generally maroon. Ambulatory legs also maroon, with cream longitudinal stripes; dactyli each with dorsal and median stripes on lateral surface; propodi also with dorsal and median stripes on lateral surface; carpi each with median stripe on lateral surface; meri each with 1 stripe extending to dorsal surface (second) or 2 submedian stripes on lateral surface. Pleon generally reddish. Figure 5 View FIGURE 5 shows a formalin-preserved specimen with still preserved colour pattern in order to supplement the above description.

Biocoenoses. Found to inhabit in gastropod shells, Rapana rapiformis (Born, 1778) (Muricidae) and Phalium glaucum (Linnaeus, 1758) (Cassidae) .

Remarks. Our specimens from the Kerala State were compared with photographs of the type material of Cancer clibanarius kindly made available through the late Dr. Patsy A. McLaughlin and Dr. Oliver Coleman (ZMB) ( Figs. 6 View FIGURE 6 , 7 View FIGURE 7 ). From the photographs of the type series, the following diagnostic features are detectable: ocular peduncles slender, with corneas not dilated and basal parts inflated; chelae spinose, particularly spines on dorsomesial margins large; cheliped carpus also bearing large spines on dorsomesial margin; merus of cheliped without prominent spine or tubercle on ventral surface; pereopods 2 and 3 relatively stout, dactyli distinctly longer than propodi; carpus of pereopod 2 bearing one or two large dorsodistal spines followed by smaller spines or tubercles. Furthermore, in spite of the rather poor condition, faint longitudinal stripes on the pereopods 2 and 3 can be seen in the lectotype (ZMB Herbst 2493) and at least one of the paralectotypes (ZMB Herbst 2495). Rahayu (2003) reported the presence of faint longitudinal stripes on the ocular peduncles, citing personal communication from the late Dr. P. A. McLaughlin. In these regards, our specimens from the Kerala State agree well with Clibanarius clibanarius , and thus they are identified with the species with little hesitation.

As Rahayu (2003) pointed out, most species of Clibanarius that possess longitudinal stripes on the dorsal surfaces of the ocular peduncles also have longitudinal stripes on the pereopods 2 and 3. These include Clibanarius ambonensis Rahayu & Forest, 1992 , C. antennatus Rahayu & Forest, 1992 , C. bistriatus Rahayu & Forest, 1992 , C. clibanarius , C. eurysternus (Hilgendorf, 1878) , C. fonticola McLaughlin & Murray, 1990 , C. infraspinatus , C. padavensis de Man, 1888 , C. rhabdodactylus Forest, 1953 , C. signatus Heller, 1861 , C. taeniatus ( Milne Edwards, 1848) , and C. zebra Dana, 1852 . Clibanarus clibanarius is morphologically most similar to C. infraspinatus , particularly in the antennular peduncles overreaching the distal corneal margins and the dactyli of the pereopods 2 and 3 being distinctly longer than the propodi and the possession of a dorsal row of spines on the carpi of the pereopods 2. Other diagnostic details and the living coloration are also very similar between the two species.

Nevertheless, C. clibanarius is readily distinguished from C. infraspinatus by the lack of a prominent tubercle or protuberance on the ventral surface of the cheliped merus, which is present in C. infraspinatus (cf. McLaughlin et al. 2007: 114, unnumbered fig.). Furthermore, each ambulatory dactylus possesses a longitudinal groove on the lateral surface adjacent to the ventral margin in C. infraspinatus , whereas in C. clibanarius , such a groove is absent or indistinct.

Clibanarius harisi Rahayu, 2003 View in CoL , known from Indonesian Papua, is also somewhat similar to C. clibanarius View in CoL in the antennular peduncles overreaching the distal corneal margins and in the general stoutness of the pereopods 2 and 3 with dactyli distinctly longer than propodi ( Rahayu 2003). However, the presence of a conspicuous ventral tubercle on the cheliped meri and the generally weak armature of the palms of the chelipeds (cf. Rahayu 2003) distinguish C. harisi View in CoL from C. clibanarius View in CoL . Furthermore, the overall armature of the ambulatory legs is generally weaker in C. harisi View in CoL than in C. clibanarius View in CoL . For example, the propodus of the left pereopod 3 is almost smooth in C. harisi View in CoL , whereas there are several protuberances or blunt tubercles on the lateral to ventral surfaces in C. clibanarius View in CoL ; the carpi of the pereopods 2 are armed only with a dorsodistal and at most one small subdistal spine in C. harisi View in CoL , but there is a row of conspicuous spines in C. clibanarius View in CoL ; the ventral margins of the pereopod 2 meri are smooth in C. harisi View in CoL , rather than having a row of small spines or denticles in C. clibanarius View in CoL . The living coloration is quite different between the two species (cf. Fig. 7 View FIGURE 7 versus Fig. 8B View FIGURE 8 ). In particular, in C. harisi View in CoL , there are no distinct longitudinal stripes on the pereopods 2 and 3.

Clibanarius clibanarius View in CoL has been frequently cited (see synonymy), but only a few references provide sufficient morphological information to assess the specific identity (e.g., Alcock 1905). We assume that references dealing with the Indian material ( Henderson 1893; Khan & Natarajan 1984; Thomas 1989) could refer correctly to C. clibanarius View in CoL . Material from the Kerala State, identified with C. infraspinatus View in CoL by Reshmi (2014), is herein reidentified with C. clibanarius View in CoL . Records from outside of India (H. Milne Edwards 1836; Kraus 1843; Stimpson 1858, 1907; Miers 1880; Walker 1887; Stebbing 1908, 1910; Estampador 1937, 1958; Yap-Chiongco 1938) need to be verified. In spite of the recent scientific expeditions or local samplings carried out by the authors and colleagues in Japan, Taiwan, Philippines, Indonesia, Thailand, Malaysia, and Singapore, no specimens of C. clibanarius View in CoL have been encountered in those areas.