Carolinea 74

88 Carolinea 74 (2016) lar ridges to the latter and a complete prodorsal enantiophysis also appears to exist ( W ang & C ui 1996). In Costeremus yezoensis and C . barbatus the apophyses Aa and Ap are both present ( F u - jikawa & F ujita 1985, C hoi 1997a). The phylogenetic distribution of the prodorsal enantiophysis suggests that in possessing this, Tokukobelba retains a highly primitive character state, linking it to Hungarobelba , Costeremus and also several basal Eupheredermata, that has become lost in virtually all other Damaeidae. The combined presence of Aa and Ap may however also be interpreted as an autapomorphy of Toku­ kobelba . Interbothridial protuberance An interbothridial protuberance forming a circu- lar structure when viewed from above (Figs 1, 3) ‑appears to be characteristic of Tokukobelba , based on the contributions of B ulanov a- Z achvat - kina (1962, 1967, 1975), B ayartogtokh (2000), F ujikawa (2011), C ho i & N amkoong (2012) and my own observations, although the character is not mentioned or clearly depicted in the brief de- scriptions of T japonica or T barbata ( A oki 1984, F ujita & F ujikawa 1986). Such a prodorsal protu- berance is apparently absent in all Damaeidae besides Tokukobelba . Clearly homologous pro- dorsal elevations exist in Hungarobelba ( M iko & T ravé 1996) and in at least three species of Costeremus ( A oki 1970, W ang & C ui 1996; B ayar - togtokh & E rmilov 2015) but are otherwise very seldomly encountered in brachypyline oribatid mites. Epimeral setation A setal formula of 3-1-3-3 for epimeres I-IV (Figs 2, 3) is shown by all members of Tokukobelba ( B ulanova - Z achvatkina 1962, A oki 1984, F ujita & F ujikawa 1986, B ayartogtokh 2000, F ujikawa 2011) where reliably known. Such a pattern is so far unknown for Belba . In Belba ( Protobelba ), 3 setae are found on epimere II ( N orton 1979c). Similarly 3 or 4 setae are present on epimere II in the Belba corynopus group ( G randjean 1936, M ärkel & M eyer 1960, E nami 1994). In Belba ( Protobelba ) ( N orton , 1979c) and within the Belba corynopus species group, 4 or more setae are inserted on epimere IV, where dependably known ( G randjean 1936, E nami 1994, P érez - Í ñigo 1997) . All members of Belba without epimere II neotrichy and for which the epimeral setation is known, such as B . cor­ nuta W ang & N orton , 1995, B . flammeisetosa T olstikov , 1995, B . heterosetosa B ayartogtokh , 2000, and B . prasadi B ayartogtokh , 2000, pos- sess an epimeral setation of 3-1-3-4 as evi- denced by the original descriptions. Within the Damaeidae, an epimeral setation of 3-1-3-3 is quite rare and has so far been noted for Metabelba platynotus G randjean , 1954, Me­ tabelba glabriseta M ahunka , 1981, ( E rmilov et al. 2010a), Porobelba weigmanni M iko , 2008, Nodo­ damaeus monticola H ammer , 1977, as well as for some Epidamaeus such as E . tenuissimus H am - mer , 1967, ( B ehan - P elletier & N orton 1985) and E . berlesei M ichael , 1898, ( B ernini 1970). Like nu- merous other features of significance in damaeid systematics, the epimeral setation formula may be subject to variability within some species. In Damaeus flagellifer Michael, 1890, for example, epimeral setations of 3-1-3-3 coexist with those of 3-1-3-4, with the latter formula being the most common ( B ernini & A rcidiano 1979). Tokukobelba shares an epimeral setation of 3-1-3-3 with all five species of Costeremus ( A oki 1970, F ujikawa & F ujita 1985, W ang & C ui 1996, C ho i 1997a, B ayartogtokh & E rmilov 2015) and Veloppia ( N orton 1978b, C hen & W ang 2002). This characteristic is also found in virtually all Ameroidea ( G randjean 1959b, 1961, 1965a, 1965b, 1966, B eck 1962, W oas 1986, 2002, A oki & Y amamoto 2000, C hen et al. 2004, M a - hunka 2009, 2010, E rmilov et al. 2011, 2012, E rmilov & C oetzee 2012) except apparently only the Rhynchoribatidae with a setation formula of 3-1-2-5 or 3-1-3-5 ( M ahunka 1985, 1986, W oas 1986) and Hungarobelba which features 3 or 4 setae on epimere II ( N orton 1977a , M iko & T ravé 1996). The possession of 2 or 3 setae on epimere IV is typical of the great majority of basal Brachy­ pylina ( G randjean 1934, W oas 2002) such as most Neoliodidae ( W oas 2000), Gymnodamaei- dae ( P aschoal 1987, W oas 1992, B ayartogtokh 2001, H ugo 2010) and Plateremaeidae ( P ascho - al 1987, W oas 1992, H unt 1996), as well as the Licnodamaeidae G randjean , 1954, ( P érez -Í ñigo 1997), Pheroliodidae ( P érez -Í ñigo 1997), Licno- belbidae ( G randjean 1965, P érez - Í ñig o 1993), Cymbaeremaeidae S ellnick , 1928, ( W oas 2000) and Eremaeidae O udemans , 1910, ( B ehan - P elle - tier 1993). N orton (1979a) suggests that an epi- meral setation of 3-1-3-4 is plesiomorph for the Damaeidae. I agree with this conclusion insofar that I regard the rich epimeral setations of the Belba corynopus group as being derived. I be- lieve, however, that the epimeral setation of 3-1-