Raouliinae Števčić, 2005

Ng, Peter K. L. & Rahayu, Dwi Listyo, 2014, Revision of the family Acidopsidae Števčić, 2005, and the systematic position of Typhlocarcinodes Alcock, 1900, Caecopilumnus Borradaile, 1902, and Raoulia Ng, 1987, with descriptions of two new genera and five new species (Crustacea: Brachyura: Goneplacoidea), Zootaxa 3773 (1), pp. 1-63 : 18-19

Subfamily Raouliinae Števčić, 2005 , stat. nov.

Raouliidae Števčić, 2005: 71 .

Typhlocarcinodidae Števčić, 2005: 62 .

Caecopilumnidae Števčić, 2011: 127 .

Diagnosis. Eye fused to orbit or mobile, pear-shaped, tapering towards small or almost undiscernible cornea; cornea pigmented or unpigmented; basal antennal article transversely rectangular, distinctly wider than long ( Fig. 6C–F ); epistome prominently sunken ( Fig. 13D ); anterolateral angle of merus of third maxilliped rounded ( Fig. 5B–I ); lateral edge of a5 some distance from cx4, 5, male opening coxo-sternal, penis long to very long with at least part of it exposed in channel between st7, st8; male abdomen does not protect penis when closed; large part of st8 visible even when male abdomen closed; female sterno-abdominal cavity relatively shallower, transversely broader, vulvae positioned on median parts of st6 ( Fig. 4B, D, F, H ).

Remarks. Typhlocarcinus integrifrons was originally described by Miers (1881: 260, pl. 14 fig. 1) on the basis of one male and one female from Goree I. in Gambia (West Africa), which he provisionally referred to Typhlocarcinus Stimpson, 1858 . Alcock (1900: 326–327) commented on a badly damaged specimen in the Indian Museum (no provenance given) and suggested it may be Miers’ species although he did not formally identify it as such. In his discussion, he argued that since it differed significantly from Typhlocarcinus s. str., a new genus, Typhlocarcinodes , should be established but he did not formally name Typhlocarcinus integrifrons as the type species or even include it there. Alcock (1900) regarded Typhlocarcinodes as a rhizopine, and intermediate between Typhlocarcinus and Scalopidia Stimpson, 1858 . Borradaile (1902: 267, fig. 59) described a new genus and new species, Caecopilumnus hirsutus , on the basis of a single female specimen from the Maldives. He was not sure where it should be classified and treated it as an incerta sedis. He did realise it may be close to Typhlocarcinus integrifrons but he was apparently not aware of Alcock (1900) establishing a new genus for the species. In an amendment a year later, Borradaile (1903: 431) realised that Caecopilumnus was close to, if not identical, with Typhlocarcinodes , and he referred it to the Rhizopinae as well.

Tesch (1918) redefined Typhlocarcinodes and discussed its relationship with Caecopilumnus at length. He decided to synonymise the two genera, and recognised four species in Typhlocarcinodes : T. integrifrons ( Miers, 1881) , T. hirsutus ( Borradaile, 1902) , T. piroculatus ( Rathbun, 1911) and T. crassipes Tesch, 1918 . He also formally recognised Typhlocarcinus integrifrons Miers, 1881 , as the type species of Typhlocarcinodes Alcock, 1900 . He transferred Borradaile’s (1902) species to Typhlocarcinodes , recorded a male specimen from off Flores (Indonesia), and described a new species, T. crassipes , from Banda in Indonesia. Typhlocarcinops piroculata was originally described on the basis of one adult female and two juvenile females from Amirante Is. in the Indian Ocean ( Rathbun 1911: 239, pl. 20 figs. 1, 2). Although there was no male specimen, it was probably referred to the rhizopine genus Typhlocarcinops Rathbun, 1909 (characterised by a very wide male first abdominal somite) because of its superficial similarity with the type species, T. canaliculata Rathbun, 1909 . Tesch (1918) reported one male specimen from Talaud I. (= Talaut Archipelago, off northern Sulawesi, Indonesia), and referred Rathbun’s species to Typhlocarcinodes .

Balss (1922) appears to have accepted this classification although he inexplicably listed his record as " Typhlocarcinodes (Typhlocarcinus) integrifrons ". Balss (1957) later treated Typhlocarcinus as a separate genus. Balss (1938) also recorded Typhlocarcinodes piroculatus from the Gilbert Is. (= Kiribati) on the basis of a female specimen. Monod (1956: 356, figs. 466–468) reported another female specimen from West Africa (as “ Typhlocarcinodes integrifrons ”) and made a comment in the footnote that the Atlantic and Indo-Pacific taxa may not be congeneric ( Monod 1956: 356). Serène (1964: 237) followed Tesch’s (1918) classification and recorded T. piroculata from Jolo, Philippines.

Takeda (1973) and Takeda & Shimazaki (1974) suggested that it was not clear which is the type species of Typhlocarcinodes Alcock, 1900 , since Alcock (1900) did not name the constituent species in the genus when he established it. They suggested the name should perhaps be kept for the Indo-West Pacific taxa and a new name established for the Atlantic taxon instead. Manning & Holthuis (1981), however, affirmed that Tesch’s (1918) selection of a type species for Typhlocarcinodes Alcock, 1900 , was nomenclaturally valid, and that if the Atlantic and Indo-West Pacific taxa belong to separate genera, then the name Caecopilumnus Borradaile, 1902 , would have to be reinstated for the latter group of species.

Ng (1987: 92) reviewed the differences between the Atlantic and Indo-West Pacific species and decided to restrict Typhlocarcinodes Alcock, 1900 , for T. integrifrons ( Miers, 1881) , from the Atlantic. He transferred the three Indo-West Pacific species to Caecopilumnus Borradaile, 1902 : T. hirsutus ( Borradaile, 1902) , T. piroculatus ( Rathbun, 1911) and T. crassipes Tesch, 1918 . Accepting that Tesch’s (1918) material as true T. piroculatus, Ng (1987: 93) also argued that since the male of this species, as figured by Tesch (1918: pl. 15 2d) had all the male abdominal somites free in sharp contrast to the fused condition of somites 3–5 figured in Serène’s (1964: fig. 15) male, the latter taxon cannot be congeneric or conspecific with Typhlocarcinops piroculata Rathbun, 1909 . As such, he established a new genus, Raoulia Ng, 1987 , for Serène’s (1964) " Typhlocarcinodes piroculatus ", and treated it as a new species, Raoulia limosa Ng, 1987 . The latter is therefore the type species by monotypy (see also Ng et al. 2008: 76). Ng (1987: 94) also regarded the Typhlocarcinodes piroculatus of Barnard (1955: 35, fig. 16) from South Africa as conspecific with Raoulia limosa .

The subfamily Raouliinae Števčić, 2005 , as defined here contains four genera: Raoulia Ng, 1987 , Typhlocarcinodes Alcock, 1900 ; Caecopilumnus Borradaile, 1902 , and Thecaplax gen. nov., all from the Indo- West Pacific.

Figures

FIGURE 4. Female thoracic sterna, abdomens and vulvae. A, B, Raoulia piroculata (Rathbun, 1911), lectotype female (7.3 × 5.5 mm) (USNM 41359a), Amirante Is., Indian Ocean; C, D, Caecopilumnus crassipes (Tesch, 1918), female (13.0 × 11.0 mm) (MZB), Lombok, Indonesia; E, F, Thecaplax capillosa gen. et. sp. nov., paratype female (12.7 × 10.5 mm) (ZRC 2013.1714), Lombok, Indonesia; G, H, Typhlocarcinodes integrifrons (Miers, 1881), paralectotype female (7.0 × 5.4 mm) (NHM 81.24b), Goree I., Senegal.

FIGURE 5. Right third maxillipeds. A, Acidops cessacii (A. Milne-Edwards, 1878), syntype female (14.1 × 10.0 mm) (NHM 99.3.23.66) Cape Verde (laterally inverted); B, Parapilumnus cristimanus (A. Milne-Edwards, 1873), male (16.3 × 12.5 mm) (ZRC 2003.90), Nansha Is. (Spratly Is.), South China Sea (laterally inverted); C, Raoulia limosa Ng, 1987, holotype male (8.3 × 6.3 mm) (ZMUC M58), off Marongas, Jolo, Philippines; D, Raoulia piroculata (Rathbun, 1911), lectotype female (7.3 × 5.5 mm) (USNM 41359a), Amirante Is., Indian Ocean; E, Raoulia galea sp. nov., holotype male (9.4 × 7.4 mm) (ZRC 2000.0980), Phuket, Thailand; F, Raoulia fortis sp. nov., holotype male (12.0 × 9.2 mm) (MNHN), Santo, Vanuatu (laterally inverted); G, Raoulia calva sp. nov., holotype male (6.6 × 5.2 mm) (ZRC 2013.1721), Society Is., French Polynesia; H, Caecopilumnus hirsutus (Borradaile, 1902), male (7.1 × 5.6 mm) (ZMA 261027), Flores, Indonesia (laterally inverted); I, Caecopilumnus crassipes (Tesch, 1918), male (11.8 × 10.1 mm) (ZRC 2013.1710) Lombok, Indonesia; J, Caecopilumnus loculatus sp. nov., holotype male (8.0 × 6.7 mm) (ZRC 2013.1723), Moorea, French Polynesia (laterally inverted); K, Thecaplax capillosa gen. et. sp. nov., holotype male (11.6 × 9.5 mm) (MZB), Lombok, Indonesia; L, Typhlocarcinodes integrifrons (Miers, 1881), lectotype male (8.5 × 6.9 mm) (NHM 81.24a), Goree I., Senegal.

FIGURE 6. Left antennae and antennules. A, Acidops cessacii (A. Milne-Edwards, 1878), male (11.3 × 8.1 mm) (ZRC 1998.58), Ascension; B, Parapilumnus cristimanus (A. Milne-Edwards, 1873), male (16.3 × 12.5 mm) (ZRC 2003.90), Nansha Is. (Spratly Is.), South China Sea; C, Raoulia fortis sp. nov., holotype male (12.0 × 9.2 mm) (MNHN), Santo, Vanuatu; D, Caecopilumnus crassipes (Tesch, 1918), male (12.9 × 11.4 mm) (ZRC 2013.1711), Lombok, Indonesia; E, Thecaplax capillosa gen. et sp. nov., paratype male (13.2 × 10.7 mm) (MZB), Lombok, Indonesia; F, Typhlocarcinodes integrifrons (Miers, 1881), lectotype male (8.5 × 6.9 mm) (NHM 81.24a), Goree I., Senegal. Abbreviations: at3 and at4 = basal antennal articles 3 and 4, respectively; bs = basal antennal article; ba = basal antennular article. Scales = 0.5 mm.

FIGURE 13. Raoulia limosa Ng, 1987, holotype male (8.3 × 6.3 mm) (ZMUC M58), Jolo, Philippines. A, dorsal view; B, male abdomen; C, anterior thoracic sternum and sterno-abdominal cavity; D, frontal view of carapace; E, posterior thoracic sternum; F, outer view of right chela; G, outer view of left chela.