Ciliopagurus strigatus ( Herbst, 1804 )

Poupin, Joseph & Malay, Maria Celia, 2009, Identification of a Ciliopagurus strigatus (Herbst, 1804) species-complex, with description of a new species from French Polynesia (Crustacea, Decapoda, Anomura, Diogenidae), Zoosystema 31 (2), pp. 209-232 : 213-216

Ciliopagurus strigatus ( Herbst, 1804) ( Figs 1A ; 2 ; 3A ; 4A ; 6-9 )

References limited to key references and field guides that illustrate this species in colour. Additional references and full synonymy are in Forest (1995a).

Cancer strigatus Herbst, 1804: 25 , pl. 61, fig. 3 [type locality: East Indies].

Trizopagurus strigatus – Forest 1952a: 256; 1952b: 19, figs 5, 14, 21. — Fielding & Robinson 1987: 78, photo 64. — Haig & Ball 1988: 177. — Nomura et al. 1988: 123. — Allen & Steene 1994: 151. — Debelius & Baensch 1994: 610. — Takeda 1994: 197. — Asakura 1995: 352, pl. 93-3. — Tudge 1995: 30. — Gosliner et al. 1996: 225, fig. 819. — Yu & Foo 1991: 57. — Jones & Morgan 2002: 123. — Laboute & Richer de Forges 1994: 391.

Ciliopagurus strigatus – Forest 1995a: 49, figs 8a, 9, 10a, 12a, 31a-b, 37c (new generic combination plus full synonymy). — Hoover 1998: 252. — McLaughlin 1997: 221. — Sakai 1999: 10, pl. 2b. — Minemizu 2000: 132. — Debelius 2001: 232. — Kato & Okuno 2001: 132. — Poupin 2001: 297, figs 1b, 3a, 4. — Kawamoto & Okuno 2003: 74. — Okuno et al. 2006: 150. — McLaughlin et al. 2007: 70.

Not Ciliopagurus strigatus – Forest 1995a: 50 (in part, specimens from the Marquesas) = C. vakovako Poupin, 2001 .

TYPE MATERIAL. — East Indies, holotype, dry specimen ( ZMB Herbst, 2487).

SHELLS. — Conidae : Conus sp. ; Conus miles Linnaeus, 1758 ; Conus striatus Linnaeus, 1758 ; Conus ? canonicus Hwass in Bruguière, 1792. Columbellidae : Euplica turturina (Lamarck, 1822) . Cypraeidae : Cypraea sp. , Cypraea caputserpentis Linnaeus, 1758 ; Cypraea helvola Linnaeus, 1758 . Muricidae : Drupa morum (Röding, 1798) .

MATERIAL EXAMINED (NB: several depth ranges are indicated for an entire scuba dive operation, sometimes down to 40 m. However, specimens of C. strigatus collected during these dives were always caught between approximately 0-15 m). — Réunion I. “Passe de l’hermitage”, coll. M. Guillaume, I.2007, 2 ♂♂ 2.2, 3.8 mm ( MNHN Pg 7759, ♂ 3.8 mm DNA H275- EF683562 ). — Near Cap Boucan Canot , outer reef slope, overhangs and black sand patches, 8-14 m, coll. G. Paulay, 16.II.2004, 1 ♀ 3.5 mm in Conus , 1 juvenile 2.6 mm in Euplica turturina ( UF Crust 5419). — Coast immediately NE of Cap La Houssaye, 21.01889°S, 55.23806°E, rocky coast with basalt bedrock boulders, 0-4 m, coll. G. Paulay, 14.II.2004, 1 ♂ 3.0 mm, 2 ♀♀ 2.0, 2.4 mm, 1 juvenile 2.0 mm (colour fading, mostly white), all specimens in Conus ( UF Crust 5420, DNA H285- EF683571 , DNA H286- EF683572 ) ; same 1 juvenile 2.3 mm ( UF Crust 5432) .

Mauritius I. Cavern, coll. J. Starmer, 1 ♀ 2.1 mm in Conus ( UF Crust 5237).

Christmas I. N coast, 4 km E of Boat Cave, just outside Thundercliff Cave, dead coral rubble field, under large limestone rocks/rubble, 10 m, coll. L. Kirkendale, 26.XI.1999, 1 ♂ 6.9 mm ( UF Crust 1941).

Taiwan. Northeast coast, Taipei County, LongDong, fringing reef, 0-3 m, coll. K. Netchy & R. Lasley, 16. V.2005, 1 ovig. ♀ 4.9 mm, in Conus ( UF Crust 7366).

Japan. Ryukyu Is, Okinawa I., Ginowan City, off Okinawa Convention Center, 26.28333°N, 127.71667°E, reef front, exposed, 1-5 m, coll. G. Paulay, 2.VII.2004, 1 ♀ 6.2 mm, in Conus miles ( UF Crust 7116). — Cape Zanpa, reef on S side of lighthouse, 26.43858°N, 127.71467°E, inner fringing reef/moat, exposed, 1-2.5 m, coll. G. Paulay, 24.VII.2004, 1 ♀ 4.1 mm, in Cypraea , 1 ♂ 4.4 mm, in Conus ( UF Crust 7235). — Ryukyu Is, Ie I., Kanan-zaki, NW end of island, 26.73917°N, 127.81008°E, outer reef slope, under rocks, 18-22 m, coll. G. Paulay & Kinjo, 8.VII.2004, 2 specimens in Conus , 1 ♂ 3.4 mm, 1 juvenile 1.8 mm ( UF Crust 7001). — Same, area around Crevice & Daidokutsu Cave, 26.724567°S, 127.83165°E, outer reef slope, probably under rock or in cavern, 3-30 m, coll. G. Paulay, 7.VII.2004, 1 ♂ 3.6 mm, in Cypraea , 1 ♀ 4.1 mm, in Conus ( UF Crust 7115, DNA H290- EF683576 ).

Philippines. Luzon I., Pangasinan Province, Bolinao Municipality, Balingasay, 16.37267°N, 119.84717°E, slope and fore reef, 6-8 m, coll. M. Malay, 26.VII.2004, 2 ♂♂ 6.1-6.3 mm, 2 ♀♀ 5.9-6.1 mm, in Conus miles ( UF Crust 6540).

Mariana Is. Supply Reef, 20.13333°N, 144.1°E, 12-20 m, coll. M. Malay et al., 1.IX.2003, 1 ♂ 3.5 mm, in Conus ( UF Crust 5781). — Maug I., Southern part of E island, inner side, 20.01667°N, 145.23333°E, 4-16 m, coll. M. Malay, 3.IX.2003, 1 ovig. ♀ 5.3 mm, in Conus , 1 ♀ 5 mm in Drupa morum , 1 juvenile ( UF Crust 5778). — Agrihan I., SE of island, 18.75°N, 145.7°E, 11-15 m, coll. M. Malay, 27.VIII.2003, 1 ♂ 6 mm in Conus , 1 ovig. ♀ 5.3 mm in Conus , 1 ♂ 4 mm in Conus ( UF Crust 5774). — Pagan I., coll. M. Malay, 1 ♂ 5.1 mm, in Drupa morum ( UF Crust 5779). — Alamagan I. SW of island, 17.58333°N, 145.81667°E, 9.5-10.5 m, coll. M. Malay, 10.IX.2003, 1 ♂ 5.6 mm in Conus miles ( UF Crust 5772). — Anatahan I., 16°N, 145°E, coll. M. Malay et al., 1 juvenile ( UF Crust 5782). — Saipan I., Bat Cave, 27 m, coll. J. Starmer, 18.XI.1993, 1 ♂ 5.6 mm in Conus miles ( UF Crust 5775). — Same, loose rubble on floor, 27 m, 21.XI.1993, 1 ♂ 3.9 mm in Conus ( UF Crust 5780). — Agingan Point, sewage outfall, 15.1185°N, 145.68667°E, exposed on surface of dead limestone rock/ rubble, 21-33 m, coll. L. Kirkendale, 11.VIII.2001, 1 ♀ 4.3 mm ( UF Crust 1871, DNA IP21- EF683559 ). — Arakane Reef, 15.63333°N, 142.76667°E, 14-17 m, coll. M. Malay, 15.IX.2003, 1 ovig. ♀ 5.4 mm in Conus ( UF Crust 5773). — Tinian I., 14.93333°N, 145.63333°E, 9-11 m, coll. M. Malay, 23.VIII.2003, 1 juvenile in Conus ( UF Crust 5776). — Aguijan I., 14.85°N, 145.56667°E, 13-16 m, coll. M. Malay et al., 17.IX.2003, 1 ♂ 2.8 mm, in Conus ( UF Crust 5783). — Rota I., 14.15387°N, 145.25938°E, 9-12 m, coll. M. Malay, 19.IX.2003, 3♂♂ 3.0- 4.3 mm, 1 ♀ 5.3 mm, in Conus sp. , Drupa morum , and Cypraea caputserpentis ( UF Crust 5777). — Guam I., Tanguisson, fore reef, 8-11 m, coll. G. Paulay, 1 juvenile 2 mm, colour completely gone except black tips on chelae and legs ( UF Crust 5920). — Haputo, rubble, 8-10 m, coll. G. Paulay, 8.VII.2003, 2 ♂♂ 2.1, 2.8 mm, in Conus and Cypraea helvola ( UF Crust 5972). — Tanguisson, fore reef, 5-20 m, coll. G. Paulay, 23. VI.2003, 1 juvenile ( UF Crust 6014). — Tepungan channel tunnel, 0-4 m, coll. G. Paulay, 16. VI.2003, 1 juvenile ( UF Crust 6017). — S Orote, under rocks, 18-22 m, coll. G. Paulay, 1 juvenile 1.9 mm, in Cypraea ( UF Crust 8706).

Papua New Guinea. Bismarck Archipelago, Sherburne Reef, 3.32389°S, 148.21528°E, subsided small atoll, outer side, forereef, steep coral slope, 3-40 m, coll.L. Kirkendale, 21. VI.2003, 1 ♂ 9.4 mm in Conus striatus ( UF Crust 4801), 1 ovig. ♀ 5.2 mm in Conus ( UF Crust 4815). — Same, 3.44917°S, 148.08222°E, 3-38 m, 21. VI.2003, 1 ovig. ♀ 6 mm in Conus ( UF Crust 4844). — Sisi Liu I., W side of Manus I., 6 mi offshore, 2.260278°S, 146.67278°E, fore reef, gentle coral slope with abundant Halimeda cover, 1-34 m, coll. L. Kirkendale, 29. VI.2003, 1 ♀ 2.3 mm, in Conus ( UF Crust 6645).

Vanuatu. Sanma Province, Aore I., Aore Point, shore near “Fantastic” dive site 15.5325°S, 167.21417°E, under and on rubble, 4.5 m, coll. C. Meyer, 23.I.2005, 1 ♀ 4.7 mm, in Conus ? canonicus ( UF Crust 8028). — Shefa Province, Epi I., Southern Lamen Bay, 16.59611°S, 168.1675°E, fringing reef, rubble, 1-3 m, coll. C. Meyer, 8.I.2005, 2 ♀♀ 5.5-7.0 mm, in Conus ( UF Crust 8502).

Wallis and Futuna Is. Futuna I., lagoon, Pito area, snorkeling 1-2 m, under rubbles, st CRISP-1, 14.2625°S, 178.17639°E, coll. J. Poupin, 9.X.2007, 1 ♂ 7.0 mm ( MNHN Pg 7762).

Fiji Is. Lau Group, Nayau I. off shore of Salia Village, fringing reef, coll. T. Bell, 8.X.2003, 1 ♀ 6.9 mm ( UF Crust 5510). — Same, 14.X.2003, 1 ovig. ♀ 6 mm ( UF Crust 5517).

American Samoa. Tutuila I. outer reef, 3-15 m, coll. V. Bonito, 16.X.2002, 1 ♂ 6.7 mm in Cypraea ( UF Crust 3247). — Same, 0-2 m, 23.X.2002, 1 ♀ 4.5 mm ( UF Crust 3270). — Same, Au’asi, 14.2715°S, 170.57283°W, under rubble, 9-15 m, 19.X.2002, 2 specimens ( UF Crust 3258).

Niue I. South Alofi, Opaahi, outer reef slope, under rocks and in substrate under rocks, 6-9 m, coll. B. Holthuis & G. Paulay, 15.X.1991, 1 ♀ 5 mm, colour completely gone except for black tips on chelae and legs ( UF Crust 1886).

Hawaiian Is. Oahu I., off Kahe Point Beach Park, outer reef slope, under rocks?, 11-15 m, coll. J. Earle, J. Hoover & G. Paulay, 24.II.2006, 1 juvenile 1.6 mm ( UF Crust 8771, DNA H288- EF683574 ). — Molokai I., Kamiloloa, 21.0697°N, 157.00162°W, fore reef, on dead Pocillopora heads, 5-8 m, coll. V. Bonito, 30.I.2003, 1 individual ( UF Crust 3627).

French Polynesia. Society Is , Tahiti I., Lafayette reef, seashore, coll. J. Poupin, 17.X.1996, 1♂, 2.8 mm ( MNHN Pg 5919). —? Tahiti I., 1 ♂ 5.3 mm ( MNHN Pg 5920).— Papeete reef, coll. “abbé Cullieret”, VIII.1890, 1 ♂ 5.9 mm ( MNHN Pg 1745). — Coll. Bredin Expedition, 8.V.1957, 1 ♀ 5.3 mm ( MNHN Pg 1746) ; 1 ♂ 5.5 mm, 1 ♀ 5.9 mm (MS 5013). — Moorea I. Tiahura reef, coll. M. Monteforte, 26.VI.1982, 1 ♂ 4.1 mm ( MNHN Pg 5425). — Tuamotu Is, Rangiroa I., c. 1 km S of NW point of atoll, off Motu Maeherehonae, 14.92867°S, 147.85783°W, outer reef slope, under rocks, 6-12 m, coll G. Paulay, 10.XI.2001, 2 ♂♂ 3.3, 4.3 mm, 1 ovig. ♀ 4.5 mm, 1 ♂ 3.3 mm, all in Conus ( UF Crust 1818, DNA H287- EF683573 , H288- EF683574 , photo) .

DISTRIBUTION ( Fig. 9 ). — Ciliopagurus strigatus is widely distributed in the Indo-West Pacific, including Red Sea, Réunion, Mauritius, West coast of India, Sri Lanka, Christmas I., Cocos Keeling Is, NW Australian coast ; Vietnam, Indonesia, Taiwan, South Japan (Ryukyu and Izu Is., Izu peninsula), Philippines, Northern Mariana and Guam, Papua New Guinea, East Australia (Queensland), Chesterfields , New Caledonia, Vanuatu, Fiji, American Samoa, Niue, Hawaii , French Polynesia ( Society and Tuamotu Is ). Its northernmost latitude is 34°53’N, at Izu peninsula (water temperature influenced here by warm Kuroshio current) ; its southernmost latitude is about 27°S, Moreton Bay (Queensland). It is collected from low intertidal to about 20 m.

DIAGNOSIS. — Ocular peduncles 0.61-0.74 times as long as shield (average 0.68). Distal segment of antennular peduncle 0.21-0.29 times as long as shield (average 0.25). Ocular acicles with 3 or 4 terminal spines. Chelipeds equal; outer face of chela with 3 complete transverse striae, striae smooth or with minute spinules. Chela 0.70-1.03 times as long as shield (average 0.90); ratio of height to length 0.61-0.75 (average 0.68); fingers 0.48-0.61 times as long as chela (average 0.54). Main stridulating area with 9-11 parallel corneous crests,distally rounded or acute; 6th to 8th (from dorsal margin) crest longest, 0.33-0.50 times as long as stridulating area. Merus of cheliped without prominent tubercle on ventral face. Dactyl of third ambulatory leg 0.88-1.09 times as long as propodus (average 1.03). Posterior lobes of telson subequal, unarmed or with 1 or 2 inconspicuous spines on terminal margins.

COLORATION ( Fig. 1A ). — Antennular and antennal peduncles orangish-red; ocular peduncles red, cornea black. Chelipeds ( Fig. 3A ) and ambulatory legs ( Fig. 4A ) with bright red rings, alternating with narrower yellow rings; red rings present on chelae (including fingers), dactyls, propodi, carpi and meri. Propodi of ambulatory legs each with a distal red ring along anterior margin and 5 or 6 subsequent red rings (sometimes divided by yellow lines). Shield pure white; abdomen red with a pattern of undulating yellow lines.

CLARIFICATION OF THE TAXONOMIC STATUS

OF CANCER STRIGATUS HERBST, 1804 ( Fig. 2 )

The type specimen of Cancer strigatus is part of J. F. W. Herbst’s collection housed in ZMB. This is a dry specimen in its shell ( Conus ), photographed in Sakai (1999: pl. 2b) and also in this work ( Fig. 2A ). It is totally discoloured but the colour pattern is available in Herbst’s original description (pl. 61, fig.3) reproduced here on Figure 2B .When preparing his work Sakai (1999) corresponded with Prof. J. Forest and inserted this comment for the holotype of C. strigatus (p. 10): “The present holotype differs from specimens identified as this species by Forest (1995).A revision of the identity of this new material will be undertaken by J. Forest (pers. comm.), so that the whole case is left here at it is.”

According to J. Forest (pers. comm.), the transverse striae on the cheliped appear tuberculated in the photograph published by Sakai (1999). This is an important character used in his work ( Forest 1995a: 46, key) to split the genus Ciliopagurus into two subequal groups of species: group 1, with eight species that have no tubercles on the transverse striae of the chelipeds, including C. strigatus ; group 2, with seven species that have tubercles on these transverse striae. Based on the impression that tubercles are present in Sakai’s (1999) photograph, J. Forest considered that the type specimen of C. strigatus could not be included in group 1 and was probably conspecific with one species of group 2, possibly C. alcocki Forest, 1995 or C. hawaiiensis .

To clarify the taxonomic status of C. strigatus , detailed digital photographs of the type specimen were obtained for this work via the curator of ZMB collections. They show that the tuberculated aspect of the striae on the chelipeds is mostly an artifact due to the presence of crystals and/or dry pieces of varnish on the type specimen ( Fig. 2C, D ). In addition they confirm that the ocular acicle is truncated with 3 or 4 terminal spines, a characteristic already indicated in 1952 by Prof.Schellenberg, at that time curator of ZMB collections (see Forest 1952b: 22). This kind of ocular acicle is only observed in a few Ciliopagurus species of group 1 whereas group 2 species all have an elongated ocular acicle terminated by 1 or rarely 2 spines. Therefore the type specimens of C. strigatus cannot be conspecific with one species of group 2, as hypothesized by J. Forest.

In conclusion it appears that no tubercules are present on the transverse striae of the chelipeds and the ocular acicles are truncated, each with 3 or 4 terminal spines. Therefore, Forest’s (1995a) interpretation of the taxonomic status of C. strigatus is considered correct, and is the same as commonly recognized in field guides on the Indo-West Pacific fauna (e.g., Hoover 1998; Debelius 2001; Kato & Okuno 2001; Jones & Morgan 2002). This conclusion is reinforced by the fact that species collected at Herbst’s time were common littoral species, whereas all Ciliopagurus species of group 2 in Forest (1995a) are living deeper than 100 m. The possibility that Herbst’s type specimen belongs to one of the other species of the “ strigatus complex” ( C. tricolor , C. vakovako , C. galzini n. sp.) does not seem plausible because none are known from the type locality of C. strigatus , “East Indies”, and all have a different colour pattern (compare Fig. 2B with Fig. 1 B-D).

Figures

FIG. 1. — Live colorations: A, Ciliopagurus strigatus (Herbst, 1804), Futuna I., ♂ 7 mm (MNHN Pg 7762); B, C. tricolor (Forest, 1995), Réunion I., ovig.♀ 2.9 mm (UF Crust 5433);C, C. vakovako Poupin,2001, Marquesas Is, holotype ♂ 4.3 mm (MNHN 5896); D, C. galzini n. sp., Tuamotus, holotype ♂ 3.8 mm (UF Crust 10901).

FIG. 2. — Type specimen of Ciliopagurus strigatus (Herbst, 1804): A, C, D, type specimen in ZMB collection; A, general view; C, detail of frontal region; D, detail of right chela, outer view (photographs courtesy of O. Coleman); B, colour pattern as illustrated in Herbst (1804: pl. 61, fig. 3) (courtesy of M. Türkay).

FIG. 3. — Left cheliped colour pattern, outer view: A, Ciliopagurus strigatus (Herbst, 1804), ♂ 3.8 mm, Réunion I. (MNHN Pg 7759); B, C. tricolor (Forest,1995),ovig.♀ 3.3 mm,Réunion I. (MNHN Pg 7760);C, C. vakovako Poupin,2001, holotype ♂ 4.3 mm,Marquesas Is (MNHN Pg 5896); D, C. galzini n. sp., holotype ♂ 3.8 mm (UF Crust 10901). Scale bars: 1 mm.

FIG. 4. — Left third pereiopod colour pattern, outer view: A, Ciliopagurus strigatus (Herbst, 1804), ♂ 3.8 mm, Réunion I. (MNHN Pg 7759); B, C. tricolor (Forest, 1995), ovig. ♀ 3.3 mm, Réunion I. (MNHN Pg 7760); C, C. vakovako Poupin, 2001, holotype ♂ 4.3 mm, Marquesas Is (MNHN Pg 5896); D, C. galzini n. sp., holotype ♂ 3.8 mm (UF Crust 10901). Scale bars: 1 mm.

FIG. 6. — Length of ocular peduncle/shield length. Data from Table 1, means ± 1.00 or 1.96 standard deviation, calculated for species of the strigatus species complex and Ciliopagurus krempfi (Forest, 1952): * indicates a significant difference (ANOVA and paired tests, P<0.05).

FIG. 7. — Maximum likelihood phylogram based on COI mitochondrial DNA sequences. Values above the branches correspond to parsimony bootstraps (left), maximum likelihood bootstraps (middle), and Bayesian posterior probabilities (right).

FIG. 9. — Geographic distributions of the Ciliopagurus Forest, 1995 species included in the “strigatus complex” based on the material examined for this work and literature. Light gray area is the distribution of C. strigatus (Herbst, 1804) (●); dark gray areas are the distribution of C. tricolor (Forest, 1995) (▲) and C. vakovako Poupin, 2001 (★); asterisk (Å) is the distribution of C. galzini n. sp. A doubtful locality for C. tricolor is indicated by “▲?”.