Coenobitidae Dana, 1851
|
publication ID |
https://doi.org/ 10.6620/ZS.2023.62-52 |
|
persistent identifier |
https://treatment.plazi.org/id/0E4A87D6-FFD8-8078-FEC1-FDDDFEA11F8C |
|
treatment provided by |
Felipe |
|
scientific name |
Coenobitidae Dana, 1851 |
| status |
|
Family Coenobitidae Dana, 1851 View in CoL Genus Coenobita Latreille, 1829
Coenobita longitarsis De Man, 1902 View in CoL ( Figs. 1–4 View Fig View Fig View Fig View Fig , 5A, C, E View Fig )
Coenobita longitarsis De Man, 1902: 746 View in CoL , pl. 24(47) (type locality: Maluku, Indonesia; see Remarks); Alcock 1905: 193 (list); Gordan 1956: 311 (list); Hartnoll 1988: 14 (list); Nakasone 1988: 165 (list); McLaughlin et al. 2010: 16 (list); Hamasaki et al. 2017: e12369 (p. 2) (list); Sasaki 2019: 5486 (list); Schäfer 2020: 88, 2 images on p. 89 ( Guam).
Material examined: Syntypes: 2 ññ (SL 6.2, 13.1 mm) ( SMF 8170), “Indonesien, Molukken, vermutlich Ternate”, coll. W. Kükenthal, 1894. Others: Maluku, Indonesia: 1 ò (SL 7.4 mm), 1 ovig. ñ (SL 5.1 mm) ( SMF 8199), in the forest, about 1.5 km from the sea, coll. Merton, H., 7 Mar. 1908. Papua New Guinea: 1 ò (SL 10.1 mm) ( UF 11434), 2 ññ (SL 10.3, 11.2 mm) ( UF 11435), 1 ñ (SL 9.1 mm) ( UF 11436), Saukale, New Britain, East New Britain Province, primary hill forest 850 m, about 10 km from the sea (-5.4255, 151.4534), coll. J. D. Slapcinsky, 28 Feb. 2005; 1 ñ (SL 6.1 mm) ( UF 8362), Araetha Village, Sudest Island, Milne Bay Province (-11.4362, 153.4301), coll. J. D. Slapcinsky, 11 Apr. 2004; 1 ñ (SL 9.9 mm) ( UF 11438), Wowo, East New Britain Province, coll. J. D. Slapcinsky, 23 Feb. 2005.
Comparative materials: C. spinosus H. Milne Edwards, 1837: 1 ovig. ñ (SL 35.1 mm) ( UF 7501), Epi Island, Shefa Province, Vanuatu, coll. C. Meyer, 8 Jan. 2005; 1 ò (SL 21.8 mm) ( UF 15602), Moorea, Society Islands, coll. 11 Oct. 2008; 1 ñ (SL 24.8 mm) ( UF 2449), Niue Island, coll. 9 Sep. 1991; 1 ñ (SL 12.1 mm) (NCHUZOOL 17196), Alofi Island, Wallis and Futuna. C. violascens Heller, 1862: 1 ò (SL 20.2 mm) (NCHUZOOL 13631), Dongsha I., Kaohsiung, Taiwan, coll. 17 Nov. 2011; 1 ñ (SL 18.2 mm) ( ZRC), Lobo, Kaimana, West Papua, Indonesia, coll. D. Telnov, 15 Sep. 2010; 1 ñ (SL 18.0 mm) (NCHUZOOL 17221), Shenggang, Changhua, Taiwan, coll. K. Chang, 9 Aug. 2018.
Diagnosis: Shield about 1.4–1.5 times as long as broad (n = 6); lateral projection produced, terminating in blunt spine. Ocular peduncles approximately 0.4–0.5 times length of shield (n = 6), compressed, reaching beyond midpoint of ultimate antennal segments; ocular acicles triangular, terminating acutely. Antennal acicle fused with second peduncular segment. Palm of left cheliped lacking stridulating ridge on upper outer surface; upper half of outer face with more scattered granules, fewer in lower half; lower margin nearly straight or with an obtuse corner in middle portion; inner side near upper margin with brush of setae. Palm of right cheliped with thick brush of long setae on upper margin. P2 and P3 long and slender. Left P3 with propodus 1.1–1.2 times length of shield, length/width 2.1–2.6 (n = 5; excluding one with a small leg probably caused by regeneration); lateral surface of propodus nearly smooth, not separated from dorsal surface by longitudinal crest; ventrolateral margin of propodus concave; inner surface concave; dactyl of left P3 long, 1.0–1.3 times as long as propodus (n = 6). Coxae of male P5 subequal, with left one slightly larger, both thick and short; no sexual tubes developed; sternal protuberance ovate, large.
Size: Largest male SL 10.1 mm ( UF 11434); largest female SL 13.1 mm ( SMF 8170, syntype).
Coloration: Anterior carapace reddish-purple, posterior reddish-yellow. Ground color of cheliped greenish, with bluish areas, outer merus reddish, tips of fingers reddish-white. P2 and P3 steel blue, dactyls with reddish tips. Juvenile with carapace and legs pale yellowish-red, dactyl and propodus of left P3 with bluish tinge (after De Man 1902; cf. Schäfer 2020: 89). Based on fresh color observed in figure 4A–C, carapace, chelipeds and legs brownish gray; ocular peduncles brownish gray, cornea black; antennal and antennular peduncles orangish brown; fingers of chelipeds and dactyls of legs with tips orangish tinge. From residual color of preserved specimens ( Fig. 3 View Fig ), a pale inverted T band on anterior shield; two dark patches, separated by a pale horizontal belt, on outer surface of proximal palm of left cheliped.
Ecological notes: The habitats include the inland forest (ca. 1.5 km from the sea) in Maluku, Indonesia ( De Man 1902); the primary hill forest (850 m a.s.l. (= above sea level) and about 10 km from the sea) in New Britain, East New Britain Province, Papua New Guinea and along creek in disturbed forest (0 m a.s.l.) in Sudest Island, Milne Bay Province, Papua New Guinea (this study); and a forested area far from river and about 3 km from the sea, in San Fernando, Sibuyan Island, central Philippines (Emerson Sy, pers. comm.; Fig. 4A–C View Fig ). The snail shells used by C. longitarsis include Discoconcha isis (Pfeiffer, 1860) ( Camaenidae ) ( Fig. 4D–G View Fig ) and Lamarckiella zeus (Jonas, 1843) ( Chronidae ) ( Fig. 4A–C View Fig ) (see DISCUSSION).
Distribution: Indonesia (type locality: Maluku) ( De Man 1902), Papua New Guinea (this study), Philippines ( Fig. 4A–C View Fig ), and Guam ( Schäfer 2020).
Remarks: The description of C. longitarsis in De Man (1902) was based on one large female (SL 13.1 mm) and one small female (SL 6.2 mm) ( Fig. 1 View Fig ). The description and figures of the coxae of male P5 are provided for the first time in our study ( Figs. 2B View Fig , 5E View Fig ). The morphology of C. longitarsis is unique among the genus in the slender body, as well as the long segments in chelipeds and legs, which was compared with C. spinosus and C. violascens in De Man (1902) . For example, the length/width ratios of shield and left P3 propodus, and length/height ratio of left chela are 1.4–1.5, 2.1–2.6 and 1.3–1.6, respectively (n = 6) (vs. 1.3–1.4, 1.6–1.7 and 1.2–1.5 in C. spinosus [n = 3]; 1.3, 1.7–1.9 and 1.3 in C. violascens [n = 3]). Regarding other characters, C. longitarsis has a smooth outer surface on palm of left cheliped and dactyl and propodus of left P3 ( Figs. 1A, D, G View Fig , 2E, G View Fig , 3 View Fig , 4A– C View Fig , 5A, C View Fig ) (vs. with numerous prominent spinulose tubercles in C. spinosus [ Fig. 5B, D View Fig ; Asakura 2004: fig. 2B, C, D]); with numerous scattered granules in C. violascens [ Nakasone 1988: fig. 7B]); ventrolateral margin of propodus concave ( Figs. 1G View Fig , 2G View Fig , 3 View Fig , 5C View Fig ) (vs. convex in C. spinosus [ Fig. 5D View Fig ]; concave in C. violascens [ McLaughlin et al. 2010: fig. on p. 17]); and ovate sternal protuberance ( Figs. 2B View Fig , 5E View Fig ) (vs. subpentagonal in C. spinosus [ Fig. 5F View Fig ; cf. Asakura 2004: fig. 2E]; elongated in C. violascens [ Nakasone 1988: fig. 7F]). In addition, C. longitarsis is also similar to C. cavipes in the coloration of brownish gray, but no distinct dorsolateral margin on the left P3 propodus in C. longitarsis ( Figs. 1G View Fig , 2G View Fig , 3A, C View Fig , 4A–C View Fig , 5C View Fig ) (vs. dorsolateral margin delimited by row of corneous tipped tubercles in C. cavipes [ Rahayu et al. 2016: figs. 6D–F, 8E]).
Based on the above characters, the species in the images taken from San Fernando, Sibuyan Island, central Philippines ( Fig. 4A–C View Fig ) and the image taken from Guam in Schäfer (2020: 88) are identified as C. longitarsis , especially the long dactylus of left P3 and no distinct dorsolateral margin of left P3 propodus.
Coenobita pseudorugosus Nakasone, 1988 View in CoL ( Figs. 6 View Fig , 7 View Fig , 8A, C, E, 8A, C, E, G View Fig , 9A, C, E, F View Fig , 10A, C, E View Fig , 11A, C, E, G, I, K View Fig )
Coenobita pseudorugosus Nakasone, 1988: 165 View in CoL , figs. 1A–H, 2 (type locality: Cebu, Philippines); McLaughlin et al. 2010: 16 (list); Malay et al. 2018: 53 (list); Hamasaki et al. 2017: e12369 (p. 2) (list); Sasaki 2019: 5490 (list).
? Coenobita pseudorugosus View in CoL : Barnes 2002: 163–172 (SW Madagascar); Barnes and De Grave 2002: 714–726 (SW Madagascar); Barnes 2003: 549–557 (SW Madagascar); Schäfer 2020: 96, 2 images on p. 97.
Not Coenobita pseudorugosus View in CoL : Asakura 2004: 83, fig. 1 ( Maldives) (?= C. perlatus View in CoL ).
Material examined: Philippines: 1 ò (SL 7.9 mm) (NCHUZOOL 17197), Camiguin, coll. H.-T. Shih, 30 Aug. 2003; 1 ñ (SL 6.9 mm) ( UF 7382), Badian, Cebu, coll. 25 Apr. 2005; 2 òò (SL 4.7–5.0 mm), 1 ñ (4.2 mm) ( UF 13164), Panglao, Bohol, coll. 9 Jun. 2004. Central Sulawesi Province, Indonesia: 2 òò (13.0– 14.2 mm) (NCHUZOOL 17199), coll. Oct. 2022; 6 òò (8.0– 13.8 mm) (NCHUZOOL 17200), coll. Dec. 2022; 5 òò (9.1–12.7 mm) (NCHUZOOL 17201), coll. Dec. 2022.
Comparative materials: C. rugosus H. Milne
Edwards, 1837: 1 ò (SL 14.1 mm) (NCHUZOOL 17205), Gangkou R., Pingtung, Taiwan, coll. 10 Sep. 2012; 1 ò (SL 17.8 mm) (NCHUZOOL 17202), 1 ovig. ñ (SL 12.9 mm) (NCHUZOOL 17203), 1 ovig. ñ (SL 16.39 mm) (NCHUZOOL 17206), Taiping (= Itu Aba) I., Kaohsiung, Taiwan, coll. 30 Jun. 2009; 1 ovig. ñ (SL 13.1 mm) (NCHUZOOL 17204), 1 ò (SL 18.6 mm) (NCHUZOOL 15433), Taiping I., Kaohsiung, Taiwan, coll. 12 Aug. 2011; 1 ò (SL 11.2 mm) ( QM W17540), Cartier Reef, Timor Sea, Australia, coll. 5 May 1992. 1 ò (SL 5.1 mm) ( ZRC), Santo, Vanuatu, coll. 14 Sep. 2006; 1 ñ (SL 6.8 mm) (NCHUZOOL 17198), Alofi Island, Wallis and Futuna.
Diagnosis: Shield longer than broad, about 1.2–1.3 times as long as broad (n = 13); lateral projection produced, terminating in blunt spine. Ocular peduncles approximately 0.4–0.5 times length of shield (n = 13), compressed, reaching beyond midpoint of ultimate antennal segments; ocular acicles triangular, terminating acutely. Antennal acicle fused with second peduncular segment. Upper margins of both chelae with brush of setae. Palm of left cheliped with 5–9 oblique laminar tubercles on upper outer surface; outer surface with scattered round granules, numerous on lower portions, lower margins of left palm nearly straight or convex in distal part. Palm of right cheliped with thick brush of long setae on upper margin. Left P3 with propodus 1–1.2 times length of shield, length/width 2.0–2.4 (n = 12); lateral surfaces of propodus smooth and slightly swollen, separated from dorsal surface by distinct crest; ventrolateral margin of propodus concave; dactyl of left P3 long, 1.0–1.2 times as long as propodus (n = 11). Coxae of male P5 subequal, with left thick and short; right coxa produced into an elongate sexual tube, slender and longer than left, its tube turning to left and curved ventrally; sternal protuberance ovate, large.
Size: Largest male SL 14.2 mm (NCHUZOOL 17199); largest female 6.9 mm ( UF 7382).
Coloration ( Fig. 7C–H View Fig ): Color variable, including pink, reddish brown, dark brown and bluish-gray. A broad transverse band or two dark patches at anterior 1/3 of shield and two longitudinal stripes on posterior portion.
Ecological notes: Individuals inhabit sandy beach with vegetation or shrubs and could be found at the lower base of the palm leaves. The sympatric species include C. rugosus (in Panglao, Sumatra and Sulawesi) and C. violascens (in Sumatra).
Distribution: Philippines (Cebu; Bohol; Camiguin) and Indonesia (Sulawesi; Sumatra).
Remarks: In this study, we did not have the opportunity to examine the holotype of Coenobita pseudorugosus Nakasone, 1988 , which should be entrusted to an institution at the University of the Ryukyus. However, Tohru Naruse (University of the Ryukyus) is currently attempting to locate it ( T. Naruse, personal communication). According to the description in Nakasone (1988), this species can be distinguished from C. rugosus by the distal lower margin of the left palm, the coloration of outer surface of left palm, as well as the morphology of sexual tubes of male P5 coxae. However, after examination of the available specimens of C. pseudorugosus and C. rugosus , we note that in both species the distal lower margin of the left palm varies from nearly straight to convex; moreover a large dark-colored patch can either be present or absent on the outer surface of the left palm of both species ( Fig. 8 View Fig ). Nakasone (1988) probably failed to observe the high degree of overlap in these two characters because he only examined smaller specimens: SL 7.3–12.4 mm for males and SL 5.6–10.8 mm for females, with the largest male as the holotype. Our male specimens from Central Sulawesi are larger and the largest one is 14.2 mm (NCHUZOOL 17199). Among the specimens of C. rugosus examined, it is common that the size of C. rugosus is comparatively larger, e.g., the largest is SL 18.6 mm (NCHUZOOL 15433).
Regarding the coloration of C. pseudorugosus, Nakasone (1988) mentioned the basic coloration is dark brown, with a broad transverse band or two dark patches at the anterior 1/3 of shield and two longitudinal stripes on the posterior portion which can be observed in our specimens, but in our specimens the basic coloration included light brown, dark brown and bluish-gray ( Fig. 7C–H View Fig ). Because the basic coloration of C. rugosus is also variable ranging from cream, purplish, bluish-gray to dark blue ( McLaughlin et al. 2007: figs. on pp. 12–14), attempts to separate the two species only by the basic coloration may lead to misidentifications.
The morphology of sexual tubes of male P5 coxae, as emphasized by Nakasone (1988), is reliable in the examined specimens. The right sexual tube is slender and much longer than the left one in C. pseudorugosus from juveniles to adults ( Figs. 6B View Fig , 9A, C View Fig ). However, the right tube is equal to or only slightly longer than the left one in C. rugosus ( Fig. 9B, D View Fig ; Table 2). Variations exist in the length of the right sexual tube in C. pseudorugosus . In the smallest male (SL 4.7 mm; UF 13164), the right one is slender and longer than left one to a lesser degree ( Fig. 9E View Fig ), but it is still longer than those in C. rugosus ( Fig. 9B, D View Fig ). A male (SL 9.9 mm; NCHUZOOL 17200) possesses both long tubes crossed at their midpoints ( Fig. 9F View Fig ), which is suggested to be a malformation during individual development. This suggestion is consistent with the explanation for the significant variation (including either long left or long right sexual tube) in Pteropagurus McLaughlin & Rahayu, 2006 ( McLaughlin and Rahayu 2008) and takes into account the observation of a scar trace in the middle of the right sexual tube ( Fig. 9F View Fig ). The sternal protuberance of male P5 coxae is also different with ovate in C. pseudorugosus ( Figs. 6B View Fig , 9A, C View Fig ) and elongated in C. rugosus ( Fig. 9B, D View Fig ; Table 2).
In addition, the lateral and dorsal surfaces of left P3 propodus are comparatively swollen and wider respectively in C. pseudorugosus , but comparatively flatter and narrower in C. rugosus with similar sizes (ca. SL <15 mm) ( C. pseudorugosus : Figs. 10A, C, E View Fig , 11A, C, E, G; C View Fig . rugosus: Figs. 10B, D, F View Fig , 11B, D, F, H, J View Fig ). However, large specimens of C. rugosus (ca. SL> 15 mm) have swollen lateral surface and wider dorsal surface of left P3 propodus ( Figs. 10G, H View Fig , 11L View Fig ; Table 2).
In conclusion, C. pseudorugosus and C. rugosus can be distinguished reliably by the size of adult, lateral surface of left P3 propodus and sexual tube of male P5 coxae (Table 2).
In this study, we have examined and confirmed s p ecimen s f r o m th e P h ilip p in es (C eb u; B o h o l; Camiguin) and Indonesia (Sulawesi; Sumatra) to be C. pseudorugosus , although several studies reported this species distributed in the Indian Ocean. For the record from Maldives ( Asakura 2004: fig. 1), no morphological characters were provided, and this specimen was likely misidentified as the common Indo-West Pacific C. perlatus based on the picture with red coloration of carapace and pereiopods. Regarding the species identified from Madagascar (Barnes 2002 2003; Barnes and De Grave 2022), neither a description nor pictures were available, so we tend not to consider the East African record until solid information is available. Schäfer (2020: 97) showed two images of specimens from Cebu, Philippines and mentioned additional records from Iriomote, Ryukyus, Japan, but further data is necessary to confirm the identification. The localities recorded of C. pseudorugosus currently are the central Philippines and Indonesia (Sulawesi and Sumatra) which suggest the distribution is at least the Philippines and the Sunda Islands. However, previous studies may have combined this species with C. rugosus , because both species are very similar and even sympatric in some localities. Careful examination of museum specimens and conducting more field surveys may help clarify the distributional ranges.
Molecular analyses
The COI sequences from specimens of ten species of the Indo-West Pacific Coenobita were obtained (or downloaded from GenBank) ( Table 1; Fig. 12 View Fig ). In the NJ tree, each species is well supported, but no apparent grouping is found, except C. brevimanus is sister to nine other species ( Fig. 12 View Fig ). Intra-specific divergences of most species are as small as ≤ 2.01% (K2P) (13 bp difference), except the larger value ≤ 2.65% (17 bp) for C. rugosus . Interspecific divergences range from 10.18% (62 bp) between C. violascens and C. spinosus to 20.28% (115 bp) between C. brevimanus and C. longitarsis ( Table 3).
No known copyright restrictions apply. See Agosti, D., Egloff, W., 2009. Taxonomic information exchange and copyright: the Plazi approach. BMC Research Notes 2009, 2:53 for further explanation.
|
Kingdom |
|
|
Phylum |
|
|
Class |
|
|
Order |
|
|
Family |
Coenobitidae Dana, 1851
| Shih, Hsi-Te, Chang, Kai, Pramono, Félix Adhi, Celia, Maria & Malay, D. 2023 |
Coenobita pseudorugosus
| Asakura A. 2004: 83 |
Coenobita pseudorugosus
| Schafer F. 2020: 96 |
| Barnes DKA 2003: 549 |
| Barnes DKA & De Grave S. 2002: 714 |
Coenobita pseudorugosus Nakasone, 1988: 165
| Sasaki J. 2019: 5490 |
| Malay MCMD & Rahayu DL & Chan TY 2018: 53 |
| McLaughlin PA & Komai T & Lemaitre R & Rahayu DL 2010: 16 |
| Nakasone Y. 1988: 165 |
Coenobita longitarsis De Man, 1902: 746
| Schafer F. 2020: 88 |
| Sasaki J. 2019: 5486 |
| McLaughlin PA & Komai T & Lemaitre R & Rahayu DL 2010: 16 |
| Hartnoll RG 1988: 14 |
| Nakasone Y. 1988: 165 |
| Gordan J. 1956: 311 |
| Alcock A. 1905: 193 |
| De Man JG 1902: 746 |