DORIPPOIDINAE, Guinot, 2023

Guinot, Danièle, 2023, A new subfamily classification of the highly diversified Dorippidae H. Milne Edwards, 1837 (Crustacea, Decapoda, Brachyura, Dorippoidea), using morphological, molecular and palaeotonlogical data, with special emphasis on its unique female reproductive system, Zoosystema 45 (9), pp. 225-372 : 261-266

Subfamily DORIPPOIDINAE n. subfam.

TYPE GENUS. — Dorippoides Serène & Romimohtarto, 1969 (type species by original designation and monotypy: Cancer facchino Herbst, 1785 ). Other included species: Dorippoides nudipes Manning & Holthuis, 1986 .

DESCRIPTION

Carapace ( Fig. 16 )

Carapace wider than long, convex posteriorly, flattened anteriorly, appearing flatter in large individuals. Dorsal surface weakly sculptured, rather smooth, covered with pubescence; ‘human face’ distinctly delineated. Only a few grooves: precervical groove continuous, distinct in small specimens, with extreme lateral part behind orbit vaguely distinct, or not indicated at all medially in adult males; indistinct, medially interrupted in females; cervical and branchiocardiac grooves deep. Median area (urogastric region) circular or oval, flanked by large, rounded, convex branchial lobes. Meso-metagastric region with two small, oblique submedian gastric pits at the base. Front consisting of two distinct triangular teeth directed forward, or of blunter teeth directed outward, separated by rather deep emargination. Inner orbital angle triangular or blunt. Outer orbital tooth sharp, pointed, reaching as far forwards as or slightly beyond frontal teeth. Antero- and posterolateral margins devoid of spine or tooth, only demarcated by very faint angle. Exposed pleurites 5-7 as granulous sclerites; exposed pleurite 6 rather narrow, granular, separated from P3 coxa by thick, whitish membrane. Carapace posterior rim not extending laterally at all along inflated posterolateral margin and lined posteriorly by nearly rectangular strip, thinner medially in male Dorippoides facchino ( Fig. 16A ) and D. nudipes ( Fig. 16C ); strip with two developed lateral extensions and thus appearing much more hollowed medially in female D. facchino ( Fig. 16B ) and D. nudipes ( Fig. 16D ).

Illustrations: Dorippoides facchino: Herbst 1785 : pl. 11, fig. 68, as Cancer facchino (reproduced by Holthuis & Manning 1990: fig. 23); H. Milne Edwards 1837: pl. 20, fig. 11, as Dorippe sima (reproduced by Holthuis & Manning 1990: fig. 24); Verrill 1869a: pl. 2, fig. 1, as Dorippe facchino (reproduced by Holthuis & Manning 1990: fig. 25a); Holthuis & Manning 1990: figs 19a-d, 22 (male syntype of Dorippe astuta Fabricius, 1785 ), 19b (reproduced by Sin et al. 2009: fig. 3B), 25b; Chen & Sun 2002: fig. 92.1; Naruse et al. 2014: fig. 2d; Wong et al. 2021: fig. 10a, pl. 2D. Dorippoides nudipes: Chen 1988 : fig. 1a, pl. 1B, C; Holthuis & Manning 1990: fig. 26a-c.

Cephalic structures ( Fig. 17A )

Eyestalk inclined, well protected in rather deep orbital hollow and along outer orbital tooth. Antennule almost entirely folded into fossa. Antenna: basal article exposed; articles 2+3 very developed, moveable, with salient setose external part; other articles directed forward; flagellum curved outwards.

Oxystomatous disposition

Opening of exhalant channels perceptible in dorsal view between rostral teeth, hardly visible in Dorippoides nudipes . Pereiopods ( Figs 16 ; 17A ; 18C )

Chelipeds of females and most males of same size and shape; heterochely only in large adult males: major chela swollen, short, with short fingers; minor chela with slender and downwardly curved fingers, forming slight angle with axis of palm.

Illustrations: Dorippoides facchino: Holthuis & Manning 1990 : fig. 19e; Chen & Sun 2002: fig. 92.2, 3; Wong et al. 2021: fig. 10b. Dorippoides nudipes: Holthuis & Manning 1990 : fig. 26d.

P2 and P3 not very long, P3 longest, all articles rather short and wide, flattened, unarmed. Meri without spines on dorsal margins; sexual dimorphism of setation in D. facchino ( Fig. 16A, B ): males with dense setae only on posterior margins and females with entirely naked margins; dimorphism not marked in D. nudipes , with naked margins in both sexes ( Fig. 16C, D ). Dactyli flattened, twisted, without fringes of hair. P3 ischium of females with spur-like process on anterior margin; process may be present on ischium of P2 but smaller ( Figs 9A ; 18A, B ); these processes completely absent in males.

Illustrations: Dorippoides facchino: Holthuis & Manning 1990 : fig. 20. Dorippoides nudipes: Manning & Holthuis 1986 : fig. 1c; Chen 1988: fig. 1d, e; Holthuis & Manning 1990: figs 26e, f, 27f; Chen & Sun 2002: fig. 92.2, 3; Wong et al. 2021: fig. 10c.

Thoracic sternum ( Figs 17 ; 18 )

Thoracic sternum almost smooth in males, minutely granular and eroded in females.Sternite 1 pointed; sternite 2 pentagonal, high, separated from sternite 3 by broad depression; sternite 3 large, extended laterally; sternite 4 with two raised submedian prominences protruding from pubescence; sternites 4 and 5 lacking sharp ridges, only with blunt, naked transverse ridge. Suture 3/4 rather long, ending in deep depression in males. In males, interruption points of sutures very close to each other at bottom of deep sterno-pleonal cavity, which may give the false impression of a median line. Female thoracic sternum extremely tilted backwards at level of ridge running through entire sternite 6.

Pleon and telson ( Figs 17B ; 18D )

Sterno-pleonal cavity very narrow and deep. On pleon no tubercles or spines, only low, blunt transverse elevations or indistinct grooves. Male pleon with articular membranes situated between all somites and across entire breadth of each somite. In males, somite 1 long, trapezoidal, widening posteriorly, with posterior margin deeply concave in middle; somite 2 long, widening posteriorly, lacking erect tubercles or spines; somite 4 short, wide; somite 5 distinctly narrowing; somite 6 elongate, very narrow, with concave lateral margins and produced posterolateral angles; telson triangular, with constricted base, its tip far exceeding level of suture 5/6.

Illustrations: Dorippoides facchino: Holthuis & Manning 1990 : fig. 21g, f. Dorippoides nudipes: Chen 1988 : fig. 1b, c.

Female pleon ( Fig. 18A, E ) wide and rounded, without tubercles or teeth; somites 2-5 with blunt but distinct transverse carina; somite 5 widest; somites 5 and 6 longest; telson small, with semicircular posterior margin.

Pleonal-locking mechanism by press-button ( Figs 17C, D ; 18B, C, F ),

Press-button in deep curve of thoracic sternal suture 5/6.

Additional female pleonal-retention mechanism

In females of D. facchino ( Fig. 16B ), dorsally exposed part of sternite 8 with very small prominence that does not overhang pleonal somite 2 and seems not functional; in females of D. nudipes ( Fig. 16C ), the process of sternite 8 is quite developed but seems too distant to be able to overhang pleonal somite 2, thus also nonfunctional. Small telson engaged between raised slopes of sterno-pleonal cavity at level of sternite 5 ( Fig. 18A, E ).

Male gonopore and penis

Gonopore coxal; coxo-sternal condition with penial tube consisting of inclined portion, then vertical portion, without visible membrane between the two (sternites 7 and 8 in contact over very short distance); bulb more or less long.

Illustrations: Dorippoides facchino: Guinot et al. 2013 : fig.18A, B. Dorippoides nudipes: Guinot et al. 2013 : fig. 19A, B.

Gonopods ( Figs 17C ; 31B )

G1 short, stubby, with very elongated coxa and well-developed basis encircling most of endopodite. Apical process twisted, ending in slender simple point, rolled-up in a spiral, triangular ( D. facchino ) or produced into long, thin whip-like appendage ( D. nudipes ); subdistal cluster of setae. Basal lobe rounded, covered with several setae.

Illustrations: Dorippoides facchino: Chen 1986a : fig. 4a, b; Holthuis & Manning 1990: fig. 21a-e (reproduced by Sin et al. 2009: fig. 4B); Dai & Yang 1991: fig. 22, as Dorippe (Dorippoides) facchino ; Chen & Sun 2002: fig. 92.4; Guinot et al. 2013: fig. 18C, D; Vehof 2020: fig.12. Dorippoides nudipes: Stephensen 1946 : fig. 4C, as Dorippe facchino ?; Chen 1988: fig. 1f-h; Holthuis & Manning 1990: fig. 26g, h.

G2 shorter than G1, straight, with indistinct flagellum.

Illustrations: Dorippoides nudipes: Stephensen 1946 : fig. 4D, as Dorippe facchino ?; Chen 1988: fig. 1i; Dorippoides facchino: Vehof 2020 : fig. 12A.

Vulvae ( Figs 17D ; 18B, C, F ; 32B )

Vulvae on clearly raised, well delimited, globose, papillaelike whitish prominences on sternite 6, each very close to the other, almost joining on median axis, in prolongation of raised oblique setose ridge; opening relatively large, not recessed, near external margin of prominence. In the diagnosis of Dorippoides by Holthuis & Manning (1990: 48) the statement that the ‘female gonopore’ is on the third sternite is erroneous (a confusion with sternite of P3).

Illustrations: Dorippoides facchino: Holthuis & Manning 1990 : fig. 21h, i; Vehof 2020: fig. 7A-C.

Female reproductive system

Studied in Dorippoides facchino by Vehof (2020: 55, figs 7E-G, 17, 20, 22). See Figure 37 and below, The female reproductive system in Brachyura , its evolution and unique disposition in Dorippidae .

Callosities

In both sexes of Dorippoides facchino ( Fig. 33A ), on dorsal part of P3, between the coxa and the widely exposed pleurite 6, a large thick, movable, whitish membrane lined on pleural side by narrow calcified strip: structure considered here as a ‘simple’ callosity. A similar structure observed in females of D. nudipes but seemingly lacking in males ( Fig. 33B ).

DISTRIBUTION AND HABITAT

Dorippoides facchino is known from Sri Lanka and India (? Pillai & Nair 1970, 1976, as Dorippe astuta ; Dev Roy 2008; Dev Roy & Nandi 2001, 2008; Venkataraman et al. 2004; Ravichandran & Kannupandi 2007; Vidhya et al. 2017; Trivedi et al. 2018: table 1; Bhat et al. 2021: table 1) eastward to Malaysia, Indonesia, Vietnam ( André 1931: 639), Thailand, southern China, Hong Kong ( Chen & Sun 2002; Wong et al. 2021), at depths between 2 and 80 m, most often between 10 and 30 m, usually in sandy mud, but also in soft clay, soft gray mud, and in stones and sand (Holthuis & Manning 1990; Ng & Davie 2002); there are no definite records from either Japan or the Philippines.

Dorippoides nudipes is known from various parts of the eastern Indian Ocean, including the southern Red Sea, the Persian Gulf ( Apel 2001), the Gulf of Oman ( Naderloo et al. 2015: table 2), Iran ( Stephensen 1946), southern Madagascar, and probably reaches South Africa (Holthuis & Manning 1990; Chen 1988). And also from the western Indian Ocean: India ( Devi & Kumar 2017: fig. 1E, F; Trivedi et al. 2018: table 1; Gosavi et al. 2021: table 3) and western Thailand ( Davie et al. 2002; Ng & Davie 2002). Recent records of Dorippoides nudipes in the Middle East need to be reviewed. In Naderloo’s 2017 Atlas of crabs of the Persian Gulf, although the keys to Dorippe quadridens and Dorippoides nudipes , their figures of the G1 and distribution maps ( Naderloo 2017: 47, fig. 4.2.e and 4.2.f, respectively) are correct, the carapace assumed to be that of Dorippoides nudipes ( Naderloo 2017: fig. 7.3) is actually that of Dorippe quadridens . In our opinion, the figures for the two species have been accidentally mixed. The correct representation of the carapace of Dorippoides nudipes is shown under Dorippe quadridens ( Naderloo 2017: fig. 7.2). Subsequently, crabs from the northwestern Persian Gulf, Iraq, identified as Dorippe quadridens by the records of Yasser & Naser (2019: fig. 2) and Al-Khafaji et al. (2019: fig. 2a, table 2) are likewise Dorippoides nudipes and not Dorippe quadridens (see under D. quadridens ).

CARRYING BEHAVIOUR

The association of Dorippoides facchino , the ‘porter crab’ or ‘anemone-carrying crab’, with another organism, in fact with a sea anemone, has long been recognised (Herbst 1796: 215, as Cancer facchino ; Stimpson 1855: 37, as Dorippe facchino ; Verrill 1869a: 58-60; 1869b: 249-250, pl. 2, fig. 1, as D. facchino [reproduced by Holthuis & Manning 1990: fig. 25a]; Henderson 1893: 405, as D. facchino ; Alcock 1896: 279, as D. granulata ; Lanchester 1900: 769; 1902: 55, as D. facchino ; Shelford 1916: 299-300, as D. facchino ; Hornell 1922: 934- 935, fig. 5, as D. dorsipes ; Verrill 1928: 16, as D. facchino ; Hose 1929: 31, as D. facchino ; Shen 1931: 101, as D. facchino ; Gravely 1941: 81, as D. facchino ; Chopra 1935, as D. facchino ; Serène & Romimohtarto 1969: 11; Morton & Morton 1983: 187, fig. 10.4, 7, as D. granulata ; Tan & Ng 1988: 149, unnumbered fig.; 1992: 149; Holthuis & Manning 1990: 60, 63, fig. 25a, b; Manning 1993: 114, fig. 3a, b; Guinot et al. 1995: fig. 4A, pl. 1B; Guinot & Wicksten 2015: 599, fig. 71-11.8B; Fautin et al. 2015: 47, fig. 5). The symbiosis between Dorippoides facchino and the actinid Cancrisocia expansa Stimpson, 1856 is very close and obligatory, with the young crab carrying a very small shell and holding with its P4 and P5 dactyli an ovoid to kidney-shaped platform on which the actinian is sitting and will grow. Most platforms consist of an eccentrically positioned bivalve shell around which the anemone had secreted chitinous material as it grows; in some cases, the shell extends the edge of the platform and gives the crab greater coverage ( Fautin et al. 2015).

The carrying behaviour of D. nudipes , a species closely related to D. facchino , has not been documented ( Manning & Holthuis 1986: 364, fig. 1c; Holthuis & Manning 1990: 70; Davie et al. 2002: 315; Devi & Kumar 2017: fig. 1E, F).

REMARKS

The species known as Medorippe lanata in the Adriatic was actually described by James Plancus (Latinisation of Simon Giovanni Bianchi, 1693-1775), who gave it the Italian vernacular name ‘facchino’, which means ‘porter’, to the inhabitants of his hometown, Rimini, a famous city on the Adriatic coast of northern Italy. According to Holthuis & Manning (1990: 52), Plancus only referred to the fact that the carapace of this crab resembles an ugly human face, as supposedly often found among members of the porter profession, and without mentioning the possibility that the crabs can carry objects with their last two pairs of legs. The point is that, in originally establishing and figuring Cancer facchino for material from the East India, Herbst (1785) erroneously included in the description a reference to the description and figure by Plancus (1739, 1760) of the Mediterranean crab now known as Medorippe lanata , thus incorrectly assuming that Cancer facchino occurred in the Mediterranean. The specific name facchino is thus the result of the misidentification by Herbst (1785) of the pre-Linnean ‘ Cancer hirsutus personatus maris Superi, vulgo Facchino Ariminensibus dictus’ by Plancus (1739; 1760: 36-38, pl. 5, fig. 1) with Dorippoides facchino . Plancus’ specimen thus becomes a syntype of the Herbst’s Cancer facchino . As Herbst’s Cancer facchino was a composite species, Serène & Romimohtarto (1969: 4) selected as the lectotype the specimen represented by Herbst (1785: pl. 11, fig. 68) (reproduced by Holthuis & Manning 1990: fig. 23), thereby fixing the identity of Herbst’s species and making facchino its valid name. Through this lectotype selection, the specific name facchino is now definitively linked to the Indo-West Pacific species of Dorippoides rather than to the Mediterranean Medorippe lanata (Holthuis & Manning 1990: 51-52, 63-64).

Dorippe astuta Fabricius, 1798 , is actually a junior synonym of Dorippoides facchino ( Ng et al. 2008: 59) .

Dorippoides nudipes was established by Manning & Holthuis (1986: 364, fig. 1c) on a single male specimen, the holotype 17.0 × 19.0 mm, from Massawa, Ethiopia, Red Sea, and on the basis of the more granular carapace and the dactyli of P2 and P3 broadest in the distal fourth rather than at midlength, in contrast to the only other species of the genus, D. facchino . Later, Holthuis & Manning (1990: 66-68) designated as paratypes several specimens from the Red Sea and various samples from the Gulf of Aden, Gulf of Oman, Persian Gulf and Madagascar, deposited in various institutions. This was an unjustified act, and these paratypes are invalid (see above, Subsequent erroneous designation of paratypes by Holthuis & Manning 1990). Although Holthuis & Manning (1990: 49 key, 69, 70, fig. 26) have listed the characters distinguishing D. nudipes from D. facchino , it is useful to restate them here. In D. nudipes , the general granulation is more marked, as for example on the anterolateral margin behind the outer orbital tooth; the fingers of chelae are shorter in young males as well in males with inflated palm; the P2 and P3 are cylindrical, with longer and slender articles with naked margins, and with paddle-like dactyli (in D. facchino merus and propodus stout and wider, with a fringe of setae on posterior border in adult males, dactylus tapering distally.

When Chen (1988: 678, fig. 1, pl. 1B, C) assigned some specimens from Madagascar to D. nudipes , she found that the main differences between the two were related to the length and setation of P2 and P3. According to Devi & Kumar (2017: 626, fig. 1E, F), their specimen from southern India fits well with the characters previously mentioned for D. nudipes , and some of the older records of D. facchino may well refer to D. nudipes due to their close similarity. The sexual dimorphism of the setation of P2 and P3 meri is marked in D. facchino , with thick fringe of setae on the posterior margins in males instead of margins naked in females ( Fig. 16A, B , respectively), but not marked in D. nudipes with naked margins in both sexes ( Fig. 16C, D ).

The Dorippoidinae n. subfam. is monotypic, with the genus Dorippoides , only known from two species.

Figures

FIG. 16. — Dorippoidinae n. subfam.: A, B, Dorippoides facchino (Herbst, 1785): A, ♂ 25.3 × 32 mm, Malaysia, Johore, Pontian, ZRC 1991.66-72; B, ovigerous ♀ 20.3 × 26.2 mm, South China Sea, Hee Huat, ZRC 1984.5347; C, D, Dorippoides nudipes Manning & Holthuis,1986: C, ♂ 16.2 × 19.4 mm, Iran,ZRC 2017.1227; D, ♀ 21.6 × 29.5 mm, South Africa, vicinity of Tugela River mouth, ZRC 2009.0885.

FIG. 17. — Dorippoidinae n. subfam. Dorippoides facchino (Herbst, 1785): A-C, ♂ 25.3 × 32 mm, Malaysia, Johore, Pontian, ZRC 1991.66-72: A, anterior ventral view; B, thoracic sternum with pleon; C, without pleon. D, ovigerous ♀ 21 × 25 mm, South China Sea, MNHN-IU-2018-5195 (= MNHN-B18802): vulvae.

FIG. 18. — Dorippoidinae n. subfam.: A-C, Dorippoides facchino (Herbst, 1785), ovigerous ♀ 20.3 × 26.2 mm, South China Sea, Hee Huat, ZRC 1984.5347: A, thoracic sternum with pleon; B, without pleon, vulvae. C, ovigerous ♀ 21 × 25 mm, South China Sea, trawl, 37 m, Chen leg. 1987, MNHN-IU-2018-5195 (= MNHN-B18802).D-F, Dorippoides nudipes Manning & Holthuis 1986: D, ♂ 16.2 × 19.4 mm, Iran, ZRC 2017.1227; thoracic sternum with pleon; E, F, ♀ 21.6 × 29.5 mm, South Africa, vicinity of Tugela River mouth, ZRC 2009.0885: E, thoracic sternum with pleon; F, thoracic sternum without pleon; vulvae. See the spurlike process on P3 ischium of females on A, B.

FIG. 9. — Particular structures of some dorippids: A, spur-like process on ischium of P3 and sometimes P2: Dorippoides facchino (Herbst, 1785), postpubertal ♀ 19.0 × 22.0 mm,Pondicherry Bay,Maindron coll.1882,MNHN-IU-2018-5199 (= MNHN-B19817).B, erect spine on sternite 8: Neodorippe callida (Fabricius,1795), ovigerous ♀ 13.5 × 14.3 mm, Amoy, MNHN-IU-2021-8738 (= MNHN-B11170). C, D, callosity at base of P3: C, Dorippe sinica Chen, 1980, ♀ 34.8 × 38.9 mm, China, Guangdong, Nanao Island, ZRC 1999.0470; D, Dorippe tenuipes Chen, 1980, ♂ 13.2 × 14.2 mm, South China Sea, ZRC 1999.0009 (the granules on P2 and P3 are not visible in the photograph). Abbreviations: c, callosity; cx2-cx5, coxae of P2-P5; e, erect spine; m, articulating membrane; p, process of sternite 8; pl6, exposed pleurite 6; p2-p5, pereiopods 2-5; r, rim; s, strip; sp, spur-like process; t, special texture area.

FIG. 31. — Diversity of G1s in the seven dorippid subfamilies (schematic drawings): A, Dorippinae n. stat. (Dorippe quadridens); B, Dorippoidinae n. subfam. (Dorippoides facchino); C, Medorippinae n. subfam.(Medorippe lanata); D-F, Heikeopsinae n.subfam.:D, Heikeopsis?japonica; E, Neodorippe callida; F, Nobilum histrio; G, Paradorippinae n. subfam. (Paradorippe cathayana); H, Philippidorippinae n. subfam. (Philippidorippe philippinensis); I, Phyllodorippinae n. subfam. (Phyllodorippe armata). (A-H, after Holthuis & Manning 1990; I, after Manning & Holthuis 1981). From Sin et al. (2009: fig. 4). Setae not always figured.

FIG. 32. — Diversity of vulvae in the seven dorippid subfamilies (schematic drawings): A, Dorippinae n. stat. (Dorippe quadridens); B, Dorippoidinae n. subfam. (Dorippoides facchino); C, Medorippinae n.subfam.(Medorippe lanata); D-F, Heikeopsinae n. subfam.;D, Heikeopsis?japonica; E, Neodorippe callida; F, Nobilum histrio; G, Paradorippinae n. subfam. (Paradorippe cathayana); H, Philippidorippinae n. subfam. (Philippidorippe philippinensis); I, Phyllodorippinae n. subfam. (Phyllodorippe armata, ♀ 13.0 × 19.0 mm, Guinean Trawling Survey, MNHN-IU-2021-8731 [= MNHN-B24199]). Setae represented only on B. A-H, modified from Holthuis & Manning (1990).

FIG. 37. — Schematic representation of character mapping of sperm storage organs in the different dorippid subfamilies recognised here (except Phyllodorippinae n.subfam.not studied) and in a typical eubrachyuran:ovary with oocytes; oviduct; cuticular bursae (thin black line), of equal or unequal size; seminal receptacle either bilateral twin- or bilateral single,and either completely cuticular (thin black line) or partially glandular (thick black line). The twin system occurs in Dorippinae n. stat., Dorippoidinae n. subfam., Paradorippinae n. subfam. and Philippidorippinae n. subfam.; the single system occurs in Heikeopsinae n. subfam.,Medorippinae n. subfam.and in the typical Eubrachyura (most, see e.g. Ethusidae). Modified from Vehof 2020: figs 20-22).

FIG. 33. — Callosities in Dorippoidinae n. subfam. (Dorippoides) and Dorippinae n. stat. (Dorippe): A, callosity reduced to a thin calcified band: Dorippoides facchino (Herbst, 1785), ♂ 25.3 × 32.0 mm, Malaysia, Johore, Pontian, ZRC 1991.6672. B, no apparent callosity in male Dorippoides nudipes Manning & Holthuis, 1986, ♂ 16.2 × 19.4 mm, Iran, ZRC 2017.1227. C, callosity as a thick, elongated bulge: Dorippe quadridens (Fabricius, 1793), ♂ 39.0 × 41.4 mm, Thailand, Pattani Province, ZRC 2003.0126. D-F, callosity as a hemicircular structure: close-up views from different angles: D, E. Dorippe sinica Chen, 1980. D, ♀ 36.2 × 39.5 mm, China, Guangdong, Nanao Island, ZRC 1999.0470; E, ♂ 36.6 × 38.6 mm, Japan, Kochi, Shikoku, SMF 57855. F, D. frascone (Herbst, 1785), ♂ 29.7 × 31.0 mm, Philippines, Exp. Panglao 2004, ZRC 2008.0076. G, H, callosity as an arched double cup: D. tenuipes Chen, 1980, South China Sea, ZRC 1999.0009: G, ovigerous ♀ 17.3 × 19.6 mm; H, ♂ 13.2 × 14.2 mm. Abbreviations: b, branchiostegite; c, callosity; cx2-cx5, P2-P5 coxae; f, sulcated part of coxa; l, onepiece lateroventral part of coxa; m, membrane; p, pleon; pr, process of retention of female pleonal somite 2; pl6, exposed pleurite 6; P2-P5, pereiopods 2-5; r, rim of carapace posterior margin; s, strip along posterior rim; 1, 2, pleonal somites 1, 2; t, bottom of callosity with special texture; 7, 8, thoracic sternites 7, 8.