PARADORIPPINAE, 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 : 279-284

Subfamily PARADORIPPINAE n. subfam.

TYPE GENUS. — Paradorippe Serène & Romimohtarto, 1969 (type species by original designation: Dorippe granulata De Haan, 1841 ). Other included species by Holthuis & Manning (1990: 108, 109): Dorippe australiensis Miers, 1884 ; Paradorippe cathayana Manning & Holthuis, 1986 ; Dorippe polita Alcock & Anderson, 1894 .

PRELIMINARY NOTE

We were able to examine only the type species Paradorippe granulata (De Haan, 1841) , so in this paper all our following descriptions, figures and mentions refer only to this one species. We believe that the genus Paradorippe sensu Holthuis & Manning (1990) and subsequent authors may not be monophyletic, with at least two groups of species (with possibly misidentified species): one, i.e., the genus Paradorippe sensu stricto, with P. granulata and P. australiensis , namely with a G1 bearing unequal processes, one being longer and hammer-shaped, and with a wide vulva on a slightly marked prominence; a second, i.e., a new genus, with P. polita (and probably also P. cathayana ), with a G1 bearing shorter processes of equal length and with a smaller, possibly recessed vulva. Genetic analyses of two of the four known species of Paradorippe , P. granulata and P. polita , support such a distinction as they indicate that they form a distinct clade “with high bootstrap and posterior probability support, with intrageneric divergence being 12.5% in 16S”, the latter being the highest recorded in the study of seven genera and 12 dorippid species by Sin et al. (2009: 229, tables 1; 2) (see our Figure 10 ). This problem still needs to be improved. However, the presumed distinctive generic characters do not affect the recognition of the subfamily Paradorippinae n. subfam.

DESCRIPTION (BASED ON PARADORIPPE GRANULATA ONLY ) Carapace ( Fig. 24A, B )

Carapace wider than long in adults, widening considerably in posterior part, flat. Dorsal surface sparsely sculptured but with distinct grooves and visible human facies, often naked, only with low, rare pubescence, lacking erect spines, coarsely granular or appearing smooth, at most slightly granular on magnification. Regions with well-delinated mesogastric region; metagastric region indistinct, urogastric region defined; generally, branchial lobes relatively large. Precervical and cervical grooves more or less distinct, maybe shallow across midline; precervical groove may be deeper than cervical, deepest at lateral margins; branchiocardiac groove quite distinct. A pair of oblique submedian gastric pits visible just at base of meso-metagastric region. Antero- and posterolateral margins not demarcated, epibranchial spine absent. Front consisting of two sharp or bluntly triangular teeth, not extending beyond outer orbital spines and separated by broadly U-shaped emargination revealing exhalant channels. Inner orbital teeth low and blunt or as rounded, lobes scarcely marked. Orbit small. Orbital fissure narrow, closed or open anteriorly. Inner suborbital tooth short, much smaller than outer orbital tooth, sometimes rudimentary. Carapace posterior rim not extending laterally at all along posterolateral margin, thus interrupted on each side, and lined posteriorly by well-defined strip, much more delineated in females ( Fig. 24B ) than in males where it is straight ( Fig. 24A ), often even appearing as separate sclerite.

Illustrations: Paradorippe australiensis: Miers 1884 : pl. 26, fig. D, as Dorippe australiensis (reproduced by Holthuis & Manning 1990: fig. 44; by Davie et al. 2015a: fig. 71-2.2D); Holthuis & Manning 1990: fig. 45a; Davie 2002: fig. p. 154; Poore 2004: fig. 95. P. cathayana: Shen 1932 : fig. 4, as Dorippe polita (reproduced by Holthuis & Manning 1990: fig. 46; by Sin et al. 2009: fig. 3G), pl. 1, fig. 11; Chen 1986b: fig. 7.33, as P. polita ; Holthuis & Manning 1990: fig. 47a; Chen & Sun 2002: fig. 98.1. P. granulata: De Haan 1839 : pl. 3, fig. 2, as Dorippe granulata (reproduced by Holthuis & Manning 1990: fig. 48); Shen 1931: pl. 6, fig. 3, as D. granulata ; 1932: fig. 8, as D. granulata (reproduced by Holthuis & Manning 1990: fig. 49), pl. 1, fig. 12; Takeda 1982b: pl. p. 94; Takeda 1983: 247, fig. p. 121; Miyake 1983: pl. 6, fig. 3; Chen 1986b: fig. 6.28; Holthuis & Manning 1990: fig. 50a; Chen & Sun 2002: fig. 97.1; Ng et al. 2008: fig. 44; Wong et al. 2021: fig. 14a, pl. 3B. P. polita: Alcock & Anderson 1894 : pl. 24, fig. 4, as D. polita (reproduced by Holthuis & Manning 1990: fig. 53); Holthuis & Manning 1990: figs 54a, 55a-c, 56.

Cephalic structures ( Figs 24A, B ; 25A ; 26A )

Eyes short, distally tapering; cornea ventrolateral; eye slightly extruded from orbit. Antennula retractable into fossa, directed anteriorly like antenna. Antenna well developed: articles 2+3 quadrangular; article 4 short but wide; article 5 much developed, foliaceous, fringed with dense setae, bent outwards nearly horizontally, extending along most of orbit and lying along eyestalk; flagellum bent inwards.

Illustrations: Paradorippe cathayana: Holthuis & Manning 1990 : fig. 46. P. granulata: Quintana 1987 : fig. 20C-c; Holthuis & Manning 1990: fig. 49. P. polita: Holthuis & Manning 1990 : figs 53, 54.

Oxystomatous disposition ( Fig. 26A )

Large development of endostomal roof, thus opening of exhalant channels clearly visible in dorsal view between rostral teeth.

Illustrations: Paradorippe cathayana: Shen 1932 : fig. 5a, b, as D. polita ; Holthuis & Manning 1990: fig. 46. P. granulata: Shen 1932 : fig. 9a, as D. granulata ; Holthuis & Manning 1990: figs 48, 49. P. polita: Holthuis & Manning 1990 : figs 53, 54.

Pereiopods ( Figs 24 ; 25A )

Chelipeds of females and non-adult males of same size and shape; marked heterochely in adult males, with globular palm having bulbous ventral protuberance, and short fingers. Carpus and merus smooth or granular, often with larger granules on upper and lower margins. Outer surface of palm smooth, punctate, or granular, with or without short hairs; dorsal margin of palm and proximal part of dactylus with fringe of long hairs; lower margin of palm, except in inflated chelae, variously fringed; lower margin of fixed finger without setae. In smaller specimens fingers very long, bent downwards, with teeth of equal size, more pointed and triangular at smaller size.

Illustrations: Paradorippe australiensis: Holthuis & Manning 1990 : figs 44, 45b, c. P. cathayana: Manning 1986 : fig. 1e; Chen 1986b: fig. 7.34, as P. polita ; Holthuis & Manning 1990: figs 46, 47; Chen & Sun 2002: fig. 98.2, 3. P.granulata: Chen 1986b : fig. 6.29; Yamaguchi et al. 1987: pl. 1, fig. 8; Holthuis & Manning 1990: figs 48, 49; Chen & Sun 2002: fig. 97.2, 3; Wong et al. 2021: fig. 14c. P. polita: Holthuis & Manning 1990 : fig. 53.

P2, P3 not very long and may be rather stout in both males and females, or longer and slender. P3 slightly longer, with flattened unarmed articles, smooth or granular; dactyli flattened, slightly twisted, with well-developed carinae on anterior and posterior surfaces and with at least a sparse layer of short setae in both sexes. Both P4, P5 reduced, with subeheliform apparatus.

Illustrations: Paradorippe australiensis: Holthuis & Manning 1990 : figs 44, 45d, e. P. cathayana: Holthuis & Manning 1990 : figs 46, 47c; Chen & Sun 2002: fig. 98.4. P. granulata: Holthuis & Manning 1990 : figs 48, 49, 50c; Wong et al. 2021: fig. 14b. P. polita: Holthuis & Manning 1990 : figs 53, 54, 55f, g, 56c.

Thoracic sternum ( Figs 25 ; 26 )

Thoracic sternum narrow, narrowing distinctly posteriorly, finely or coarsely granular. Sternite 1 with only a small portion visible dorsally; sternite 2 forming short, rectangular, sharply delineated shield with thick vertical margins, separated from sternite 3 by well-marked, straight depression. Lateral margin of sternite 2 straight, directed vertically, with slightly pointed anterior angles; lateral margin of sternite 4 slightly concave, directed obliquely; lateral margin of sternite 5 almost straight, directed vertically; lateral margin of sternite 6 clearly concave. Suture 3/4 short, curved into closed boutonniere in both sexes. Sternite 3 with distinct quadrate process bearing gynglyme for articulation of mxp3 sterno-coxal condyle close to edge of Milne Edwards opening; sternite 4 thickened on each side, with longitudinal median depression; sternites 4 and 5 with narrow extensions, i.e., laterally expanded between P1/P2 and P2/P3. Sutures 4/5-7/8 interrupted. Suture 4/5 horizontal, almost parallel to suture 5/6, only slightly oblique and weakly curved posteriorly; both sutures 4/5 and 5/6 with short interruption points; suture 6/7 interrupted in males but variable in females, being medially interrupted. Female thoracic sternum tilted backwards at level of raised ridge crossing whole sternite 6; sternite 8 lacking axial spine.

Illustrations: Paradorippe australiensis: Holthuis & Manning 1990 : fig. 45i. P. granulata: Shen 1931 : pl. 6, fig. 2; Holthuis & Manning 1990: fig. 50h; Vehof et al. 2018b: fig. 2A; 3. P. polita: Holthuis & Manning 1990 : fig. 56i.

Pleon and telson ( Figs 24A ; 25B )

Male pleon with all somites free; with articular membranes, especially at sutures 4/5 and 5/6; surface without strong tubercles or spines but often with distinctly granular ridges and bumps. Somites 1, 2 exposed dorsally in males; somites 1-3 and part of 4 exposed dorsally in adult females. Somite 1 trapezoidal, widening posteriorly, posterior margin excavated in middle; somite 2 widening posteriorly, with low transverse ridge, often appearing trilobed in dorsal view; somite 3 with swollen lateral parts; somites 4 and 5 short, each with low, transverse ridge; somite 6 with posterolateral angles produced; telson triangulary blunt, telson extending over much of sternite 5, with tip far exceeding level of suture 5/6.

Illustrations: Paradorippe australiensis: Serène & Rohmimohtarto 1969 : fig. 14. P. cathayana: Shen 1932 : fig. 5d, as D. polita ; Holthuis & Manning 1990: fig. 47d; Chen & Sun 2002: fig.98.5. P.granulata: Shen 1932 : fig. 9b, as D. granulata ; Holthuis & Manning 1990: figs 48, 50e. P.polita: Holthuis & Manning 1990 : figs 53, 54a, 56d.

Female pleon wide and rounded, smooth or variously granular. Somites 2-5 usually with blunt but distinct transverse carina; somites 4 and 5 widest, 5 and 6 longest; telson small, with semicircular or subtriangular posterior margin ( Figs 24B ; 26B ).

Illustrations: Paradorippe cathayana: Shen 1932 : fig. 5e, as D. polita ; Holthuis & Manning 1990: fig. 47e. P. granulata: Shen 1932 : fig. 9c, as D. granulata ; Holthuis & Manning 1990: figs 48, 50d. P. polita: Holthuis & Manning 1990 : figs 53, 54a, 55h.

Pleonal locking mechanism by press-button

( Figs 25 C-E; 26B-D)

Press-button quite large, located on suture 5/6 weakly curved backwards and situated far from G1 tip in males, very close (almost contiguous) to vulvae. Location of sockets clearly discernible dorsally on pleonal somite 6 by swollen lateral areas. No additional pleonal retention in females, unlike many other dorippids (see below). Sternite 8 only slightly enlarged posteriorly and, together with narrow sternite 7, only flanking the base of the pleon.

Additional female pleonal-retention mechanism

Unlike some genera ( Dorippe , Heikeopsis , Nobilum , Philippidorippe and Phyllodorippe ), in females the process emanating from the dorsally exposed sternite 8 not actually overhanging the pleonal somite 2. Here, sternite 8 only slightly enlarged posteriorly, with a square projection that just abuts the edge of the female pleon and, together with narrow sternite 7, merely flanking the base of the pleon ( Fig. 24B ).

Male gonopore and penis

Coxo-sternal condition: penis overhung over a rather long distance by sternites 7 and 8, close or not in contact; penis with inclined portion weakly sclerotised; vertical portion rather short, with long soft papilla.

Illustrations: Paradorippe granulata: Guinot et al. 2013 : fig. 17C; Vehof et al. 2018a: fig. 1B, G.

Gonopods ( Figs 25 C-E; 31G)

G1 stout throughout, short, filling most of sterno-pleonal cavity; shaft strongly swollen, abruptly constricted and angularly bent at obtuse angle towards about three-quarters of its length; bent portion wider, possibly flattened dorso-ventrally; terminal part swollen, elaborate with two distinct bulbs bearing several apical processes; these processes rather elongated, one of which being longer and hammer-shaped; no basal lobe.

Illustrations: Paradorippe australiensis: Serène & Romimohtarto 1969 : figs 26-28; Holthuis & Manning 1990: fig. 45f, g. P. cathayana: Chen 1986b : fig. 7.35, as P. polita ; Holthuis & Manning 1990: fig. 47f, g (reproduced by Sin et al. 2009: fig. 4G); Chen & Sun 2002: fig. 98.6. P. granulata Serène & Romimohtarto 1969 : figs 23-25; Chen 1986b: fig. 6.30, 31; Holthuis & Manning 1990: fig. 50f, g (reproduced by Davie et al. 2015a: fig. 71-2.22D); Dai & Yang 1991: fig. 23.1, as Dorippe (Paradorippe) granulata ; Chen & Sun 2002: fig. 97.4-6; Vehof et al. 2018a: fig. 1. P. polita: Holthuis & Manning 1990 : fig. 56e-h; Dai & Yang 1991: fig. 23.2, as Dorippe (Paradorippe) polita .

G2 straight, rather long

Illustrations: Paradorippe granulata: Chen 1986b : fig. 6.32; Serène & Romimohtarto 1969: fig. 29; Chen & Sun 2002: fig. 97.7; Vehof et al. 2018a: fig. 1.

Vulvae ( Figs 26 B-D; 32C)

Vulvae not juxtaposed, close to suture 5/6, and near distal part of raised ridge; opening large to very large, rounded, surrounded by numerous setae, clusters of setae projecting to some extent above vulva opening; vulva located on more or less prominent part of sternite 6; wider and on slightly marked prominence. Histologically, lateral margin bulging in median direction, forming a muscle-operated vulvar cover that partially roofs vulva opening, so that the opening actually visible externally is crescent-shaped.

Illustrations: Paradorippe australiensis: Holthuis & Manning 1990 : fig. 45h, i. P. granulata: Holthuis & Manning 1990 : fig. 50h, i; Vehof et al. 2018a: fig. 2. P. polita: Holthuis & Manning 1990 : figs 55i, 56i-k.

Female reproductive system

Studied in Paradorippe granulata by Vehof et al. (2018a: 68; 2018b: 82, figs 1-5; Vehof 2020: 78, 86, figs 17, 20, 22) ( Figs 35D ; 37 ). See below, The female reproductive system in Brachyura , its evolution and unique disposition in Dorippidae .

DISTRIBUTION AND HABITAT

(see Preliminary note, p. 279)

Paradorippe australiensis , known from Australia (Western Australia, Queensland), is a shallow water species, caught under stones on reef flat at 22 m, on mud at 18 m, on sand and occasional sponges and corals at 15 m (Holthuis & Manning 1990; McEnnulty et al. 2011).

Paradorippe cathayana is known mainly from many localities in China ( Shen 1932, 1937a, 1940b, as Dorippe polita ; Yang 1986, as D. polita ; Chen 1986b, as P. polita ; Chen & Sun 2002); found in tidepools on sandy beaches ( Shen 1932, as D. polita ) and shallow pools of clear water, along sand beach ( Shen 1937a, as D. polita ), Gulf of Tonkin ( Dai & Song 1986, as D. polita ) and from Vietnam ( Serène 1937, as D. polita ); also from India, Gulf of Mannar ( Rajan et al. 2017: 3, 11; Zoological Survey of India 2018 fig. p. 73).

Paradorippe granulata lives much further north than any of the Indo-West Pacific dorippids, occurring as far east in Russia as Vladivostock and Peter the Great Gulf, as well as in the Sakhalin Island ( Balss 1922; Urita 1942; Kobjakova 1955, 1966; Vinogradov 1950; Levin 1976, all as Dorippe granulata ). It is abundant in many localities of Japan (De Haan 1841; Herklots 1861; Bouvier 1899; Parisi 1914; Sakai c. 1930, 1936, 1940, 1956; Nishimura & Suzuki 1971; Yang 1986, all as Dorippe granulata ; Sakai 1976, 1985; Takeda 1975, 1982 a, b, 1983; Yamaguchi et al. 1976; Muraoka 1982; Miyake 1983; Sakai et al. 1983; K. Sakai & Nakano 1983; Minemizu 2000); of Korea ( Kim 1970, as D. granulata ; Kim 1973; Koh & Lee 2013; Lee et al. 2021); and of China, Hong Kong ( Shen 1932, 1940a, b, as D. granulata ; Takeda & Miyake 1970; Huang 1994; Chen 1986b; Dai et al. 1986; Chen & Sun 2002; Wang 2005, 2009; Wong et al. 2021) and Taiwan ( Ng et al. 2017). For complete and recent distribution of P. granulata , see Holthuis & Manning 1990; Wang et al. 2013, 2017; Ng et al. 2001, 2017.

Paradorippe polita is known from a few localities: southeast India ( Alcock & Anderson 1894; Alcock 1896; Sankarankutty 1966; all as Dorippe polita ; Dev Roy 2008; Venkataraman et al. 2004, as D. polita ; Krishnamoorthy 2007: 90, as Paradorippe granulata ; Trivedi et al. 2018: table 1) and Malaysia (Holthuis & Manning 1990). The figure by Jeyabaskaran et al. (2000: pl. 32a) of a P. granulata from the Gulf of Mannar is too small to be sure that it is P. polita instead of P. granulata . The Paradorippe polita from the Gulf of Tonkin ( Zarenkov 1972: 250) should be confirmed.

CARRYING BEHAVIOUR

Paradorippine crabs are known to exclusively select valves of bivalve molluscs, often intimately associated with sea anemones in cases of symbioses ( Castro 2015). This longrecognised behaviour has been widely documented (for full literature, see Holthuis & Manning 1990). Nevertheless, the question of how and to what extent the P4 and P5 subcheliform device is specialised to grasp the valve of lamellibranchs rather than a sponge or leaf like in other dorippids has apparently not been studied. A small male of Paradorippe australiensis was found carrying on its back a 16 mm long valve of the venerid genus Antigona Schumacher, 1819 ( Rathbun 1924: 27, as Dorippe australiensis ). Paradorippe cathayana is always found carrying a bivalve shell, which is thrown away when it buries itself ( André 1937, as Dorippe polita ; Shen 1937a, as D. polita ). For P. granulata (see Döderlein 1883: 109, as D. sima ; Patton 1967, as Dorippe ; Levin 1976: fig. 106a, as D. granulata ), the carried shells seem to belong mainly to the tellinid lamellibranch genus Macoma Leach, 1819 , on which one and sometimes two or three sea anemones are fastened; a dead shell of Macoma with three longitudinally striped anemones of the actinarian Carcinactis ichikawai Uchida, 1960 has been found carried ( Uchida 1960: 595, pl. 1, fig. 1, as Dorippe granulata ); the crab disappears completely under the shell ( Kobjakova 1966: pl. 49: fig. 1a; Minemizu 2000: fig. p. 189; Koh & Lee 2013: pl. 15). According to Quintana (1987: figs 15C-H, 17E, F, 19F, 24B), in laboratory the megalopa and first crab stages of P. granulata did not swim, had benthic habits like adults, and carried dorsally over the carapace fragments of shells or small rocks provided in rearing vessels: the dorippid early-carrying behaviour is the unique case known within the Brachyura (the homolid megalopa has a P5 that may be held in a dorsal position and bears a recurved ending with long setae corresponding to a ‘feeler’, not functional for carrying; see Rice 1964: figs 4, 8f; Williamson 1965: fig. 2).

Illustrations and data: Paradorippe australiensis: Rathbun 1924: 27 , as Dorippe australiensis ; Davie 2002: 156; Thoma 2007: 301. Paradorippe cathayana: Shen 1932: 11 ; 1937a: 171, as D.polita ; André 1937: 79, as D. polita . P.granulata: Döderlein 1883: 109 , as D. sima ; Sakai 1956: 26, fig. 9, as D. granulata ; Uchida 1960: 595, 600, pl. 1, fig. 1, as D. granulata ; Kobjakova 1955: 155, pl. 49, fig. 1a, as D. granulata (reproduced by Holthuis & Manning 1990: fig. 51); 1966: pl. 49, fig. 1a, as D. granulata ; Patton 1967: 1232, as Dorippe ; Burton 1969: fig. p. 64, as Japanese crab (redrawn by Holthuis & Manning 1990: fig. 62); Levin 1976: fig. 106a, as D. granulata ; Sakai et al. 1983: fig. p. 29; Tan & Ng 1988; 1992: 149; Guinot et al. 1995: fig. 5B; Ng et al. 2008: fig. 44; Guinot & Wicksten 2015: 599, fig. 77-11.9.J.

REMARKS

Von Siebold had collected a total of 38 specimens of Dorippe granulata , described a little later in the Fauna Japonica by De Haan (1839: pl. 31, fig. 2; 1841: 122) and which became the type species of Paradorippe Serène & Romimohtarto, 1969 . The holotype and many paralectotypes are deposited at RMNH ( Fransen et al. 1997), and a dry paralectotype is deposited at MNHN (MNHN-IU-2000-11383, ex MNHNB11383) ( Yamaguchi & Baba 1993: 304, fig. 90A, B; Yamaguchi 1993: 586).

Paradorippinae n. subfam. is monotypic, with the genus Paradorippe known from four species (see Preliminary note p. 279). We agree with Alcock & Anderson (1894) that the flattish, smooth and naked carapace of P. polita resembles more that of an ethusid than that of a dorippid.

Figures

FIG. 10. — Dorippinae H. Milne Edwards, 1837 n. stat.: habitus: A, B, Dorippe quadridens (Fabricius,1793):A, ♂ 36.3 × 38.1 mm, China Sea near Singapore, Hee Huat, ZRC 1984. 6308; B, ♀ 29.5 × 30 mm, NW Madagascar, Ambaro Bay, MNHN-IU-2018-5193 (= MNHN-B18279). C, D, Dorippe sinica Chen, 1980, China, Guangdong, Nanao Island, ZRC 1999.0470: C, ♂ 36.2 × 39.5 mm; D, ♀ 34.8 × 38.9 mm, specimen brushed.

FIG. 24. — Paradorippinae n. subfam. Paradorippe granulata (De Haan, 1841): habitus: A, ♂ 23.6 × 27.3 mm, NE Taiwan,I-Lan county, ZRC 2001.0014; B, ovigerous ♀ 23. 2 × 25.7 mm, Japan, off Hota, ZRC 1999.0082.

FIG. 25. — Paradorippinae n. subfam. Paradorippe granulata (De Haan, 1841): A, C, E, ♂ 24.0 × 26.0 mm, China, Tuandao, off Quingdao, MNHN-IU-2016-10753. B, D, ♂ 23.6 × 27.3 mm, NE Taiwan, I-Lan county, ZRC 2001.0014; A, chelae; B, thoracic sternum and pleon; C, D, thoracic sternum without pleon, with G1, G2 and press-buttons; E, G1 and G2 in situ.

FIG. 26. — Paradorippinae n. subfam. Paradorippe granulata (De Haan, 1841): A-C, ovigerous ♀ 23.2 × 25.7 mm, Japan, off Hota, ZRC 1999.0082. D, ♀ 22.7 × 24, 7 mm, China, Tuandao, off Quingdao, MNHN-IU-2016-10753. A, anterior ventral view; B, C, thoracic sternum and pleon; D, vulvae.

FIG. 35. — Schematic illustrations of female reproductive systems in representatives of four dorippid subfamilies:A, B, Dorippinae n. stat.: A, Dorippe sinica Chen, 1980, as interpreted by Hayer et al. (2016a: fig. 2); B, Dorippe quadridens (Fabricius, 1793) and D. sinica, as interpreted by Vehof et al. (2017: fig. 2A); C, Medorippinae n. subfam.: Medorippe lanata (Linnaeus, 1767), as interpreted by Vehof et al. (2017: fig. 2B); D, Paradorippinae n. subfam.: Paradorippe granulata (De Haan, 1841), as interpreted by Vehof et al. (2018b: fig. 3). E, Heikeopsinae n. subfam.: Heikeopsis japonica (von Siebold, 1824), with the same pattern shared by Neodorippe callida (Fabricius, 1798) and Nobilum histrio (Nobili, 1903), as interpreted by Vehof (2020: fig. 4). Abbreviations: a, apodeme; amb, anteromedian bursa; bu, bursa;cu, cuticle;cv, cuticular valves;bu, bursa; ev, extension of vulva;ge, glandular epithelium; mu, musculature;oc, oocyte; od, oviduct; ov, ovary; plb, posterolateral bursa; sr, seminal receptacle; vg, vagina; v, vulva.

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).