DORIPPIDAE H. Milne Edwards, 1837

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 : 250-251

Family DORIPPIDAE H. Milne Edwards, 1837

Dorippiens H. Milne Edwards, 1837: 151, 153 pro parte.

Dorippidae – Dana 1852: 390, 398; 1853: 1427 pro parte. — Miers 1886: 326 pro parte. — Ortmann 1892: 553 pro parte. — Bouvier 1897a: 784 pro parte; 1897b: 54-70 pro parte; 1898: 103-105 pro parte; 1940: 195 pro parte. — A. Milne-Edwards & Bouvier 1902: 35-39 pro parte. — Gordon 1963: 55; 1966: 353. — Serène & Romimohtarto 1969: 1-35. — Kim 1973: 287, 290 pro parte. — Guinot 1978: 247; 1979a: 101, 130, 139, 171, 176, 195, 241, 260, tables 3, 5; 1979b: 45, table 1. — Chen 1993: 316. — Dai & Yang 1991: 50. — Guinot & Bouchard 1998: 649. — Ng et al. 2008: 59; 2017: 35. — Guinot et al. 2013: 10; 2019: 300. — Davie et al. 2015c: 1077. — Luque 2015: 253. — Emmerson 2016: 330. — Sasaki 2019: 7770.

Dorippinae – Alcock 1896: 136, 273, 275-286 pro parte. — Ihle 1916: 137 pro parte. — Rathbun 1937: 75 pro parte. — Balss 1957: 1609. — Manning & Holthuis 1981: 28. — Chen 1986a: 181; 1986b: 119. — Holthuis & Manning 1990: 1-151. — Chen & Xu 1991: 57. — Guinot & Bouchard 1998: 649. — Chen & Sun 2002: 208.

Dorippidae – Dorippinae – A. Milne-Edwards & Bouvier 1900: 21.

RECOGNITION OF SEVEN SUBFAMILIES

WITHIN THE DORIPPIDAE

The species of Dorippidae , before 1969 all assigned to the single genus Doripp e Weber, 1795, were subsequently distributed into three new genera and two new subgenera ( Serène & Romimohtarto 1969) and two further new genera ( Manning & Holthuis 1981, as Dorippinae ). Finally, a total of nine genera, including a new genus, were recognised by Holthuis & Manning (1990) in a comprehensive revision resolving a great number of previous misidentifications and confusions, which contributed considerably to a better understanding of the group. These authors were probably conscious of the differentiation within the group (at the time Dorippinae ) as they provided a separate key to the genera based on the G1s that highlighted their extreme diversity. Since then, two new Dorippe species from Australia have been added ( Manning 1993) and one species of Neodorippe described ( Ng & Rahayu 2002); no new dorippid taxa have been established recently. Currently, the family contains a total of only nine genera and 27 species ( Ng et al. 2008).

Although a small group with apparently similar general facies, the Dorippidae , on closer examination, displays great diversity in body form. The extent of variation and the occurrence of significant divergence patterns within the family have never led to the formal recognition of subfamilies, however. A tentative grouping of genera into a new taxonomic arrangement is proposed here, based on a re-evaluation of morphological characters, including the cephalic appendages, the configuration of the thoracic sternum, the male genital area, the gonopod structure, the vulvae, and the female reproductive system. Seven subfamilies are recognised: Dorippinae H. Milne Edwards, 1837 n. stat., Dorippoidinae n. subfam., Medorippinae n. subfam., Heikeopsinae n. subfam., Paradorippinae n. subfam., Philippidorippinae n. subfam., and Phyllodorippinae n. subfam.

Our results are congruent with known recovered molecular data. A first molecular analysis based on 16S rRNA gene sequence data from five species and four genera supported the recognition of the Dorippidae as a monophyletic family, consisting of two main clades, with the genus Dorippe appearing basal ( Fan et al. 2004: figs 1, 2). A more complete phylogenetic tree inferred from three mitochondrial genes (16S rRNA, 12S rRNA, and COI) ( Sin et al. 2009) showed several distinct clades, consistent with previous groupings based on overall carapace morphology and other traits, including G1 structure. The results were as follows: two main groups were supported, one with Dorippe , Dorippoides , Medorippe , and the other with Heikeopsis , Neodorippe and Nobilum ; Philippidorippe , somewhat ambiguous, could be associated with the preceding three genera, however with relatively low branch support; the species of Paradorippe also formed a distinct clade, with a high intra-generic divergence, and could be a sister to Heikeopsis / Neodorippe .

Dorippids are characterised by simultaneous carrying behaviour using reduced, mobile, dorsally carried and subchelate P4 and P5 to hold various materials), forward locomotion (mixed with movements in all directions, including sideways) and fast forward burying: they can be categorised as ‘forwardburiers’ by relying on their bodies to penetrate soft substrates rapidly (see below).