Belliidae, , Grapsoidea and Ocypodoidea, 1852

GUINOT, DANIÈLE, TAVARES, MARCOS & CASTRO, PETER, 2013, Significance of the sexual openings and supplementary structures on the phylogeny of brachyuran crabs (Crustacea, Decapoda, Brachyura), with new nomina for higher-ranked podotreme taxa, Zootaxa 3665 (1), pp. 1-414 : 97

Family Belliidae

The male gonopore is coxal and the penis emerges from the coxo-sternal condyle in the four genera of the family. Acanthocyclus shows a penis emerging practically from the extremity of the well-developed condyle, which articulates on sternum under the penis and is wedged between episternite 7 and an extension of sternite 8. The penis of Heterozius and Corystoides emerges from the anterior margin of the condyle, the posterior margin of which articulates with sternite 8; the large gonopore is protected by an extension of sternite 7.

The condylar protection of the penis (see Modalities of penis protection: Condylar protection; Table 4) is assumed to be a synapomorphy of Belliidae , a feature that was previously illustrated, although not explicitly stated ( Guinot 1976: fig. 11E). The status, familial ( Belliidae ) or suprafamilial (Bellioidea Dana, 1852; see Guinot 1976: 57; Ng, Guinot & Davie 2008: 46), consisting of four genera seemingly as different as Acanthocyclus , Bellia , Corystoides , and Heterozius , is therefore confirmed here. Salva & Feldmann (2001: 23) questioned whether Heterozius should be in Belliidae . Heterozius rotundifrons , which is a true Belliidae in terms of the evolution of its reproductive characters, occupies an intermediate position between the ancestral and the most derived conditions in families such as the Corystidae ( Thompson & McLay 2005: 1114; see also Hazlett et al. 2008). In any case, a separate family Heteroziidae Števčić, 2005 , is an inappropriate rank for the seemingly apomorphic Heterozius (not plesiomorphic as commented in Ng, Guinot & Davie 2008: 46), a genus that only deserves a subfamilial status.