Carolinea 74

L amos : Tokukobelba gen. nov. (Acari: Oribatida: Damaeidae) 85 expected, if the total setal count were 21. Since it is often very difficult to accurately determine the tarsal chaetotaxy with a light microscope, the tarsi here require re-examination. The illustra- tion of leg I given by B ayartogtokh (2000, p. 311, Fig. 28) does show 21 setae and two solenidia. However, the famulus is apparently not depicted. T mongolica therefore very likely possesses 22 setae on the tarsus of leg I. B ayartogtok h (2000) accordingly clearly pictures, but does not name, the ventral setae v1´, v1´´, v2´ and v2´´. The famulus on tarsus I is fully emergent in the larva and nymphs of Tokukobelba compta (Figs 16-18), ( M ourek & M iko 2010, S eniczak et al. 2013, pers. obs.) as in the ontogenetic stages of genera such as Belba ( G randjean 1954, N orton 1977b, 1979c, N orton & P alacios - V argas 1982, S eniczak & S eniczak 2013), Porobelba G randjean , 1936, ( G randjean 1954, E rmilov & L ochynska 2009), Quatrobelba ( N orton , 1980), Metabelba ( G randjean 1954, E rmilov 2010b, E rmilov et al. 2010a) and Metabelbella B ulanova - Z achvatkina , 1957, ( E rmilov & K haustov 2011) while a sunk- en famulus, reduced in size and submerged in a sclerotized cup, occurs in the larva and the nymphs of, for example, Damaeus ( G randjean 1935, 1954, 1960a, N orton 1977b, 1978a, E r - milov et al. 2010a), most Epidamaeus ( N orton 1977b, 1979c, E nami & F ujikawa 1989, E rmilov & L ochynska 2009) and Weigmannia M iko & N or - ton , 2010, ( N orton 1977b, p. 46). An emergent famulus appears to have been ancestral within the eupherederm mites and occurs in the large majority of Brachypylina ( G randjean , 1935) al- though some groups such as the Gymnodamaei- dae G randjean , 1954, ( P aschoal 1987) and Ze- torchestidae M ichael , 1898, ( G randjean 1951) display a sunken famulus in all stases. The emer- gent famulus of Tokukobelba appears to repre- sent a phylogenetically very ancient character. On the tarsus of leg I of adult Tokukobelba the primilateral pl´´ and primiventral seta pv´ do not insert on the tarsal bulb but in the middle region of the tarsus instead, whereas setae pl´ and pv´´ do insert on the bulb based on my own observa- tions, on W ang & N orton (1995) and on the illus- trations in A oki (1984), F ujita & F ujikawa (1986), B ayartogtokh (2000) and F ujikawa (2011), al- though in the last four papers these setae are not identified by name by the authors. In the larva of Tokukobelba compta all of these setae insert on the bulb, but seta pv´ and pl´´ are dis- tally displaced in later ontogenetic stages (Figs 10-13). This ontogenetic pattern differs strongly from genera such as Damaeus ( N orton 1977a, 1978a, B ernini & A rcidiano 1979, E nami & A oki 1988, X ie & Y ang 2009), almost all Epidamaeus ( M iko et al. 2011), Kunstidamaeus M iko , 2006, ( M iko 2006b, M iko & M ourek 2008) and Dyobelba ( N orton 1979b, N orton 1979c, N orton & R yabinin 1994, E nami & A oki 2001, B ayartogtokh & N or - ton , 2007) in which seta pv´´ additionally shifts anteriad in the adult stage. An ontogenetic setal shift of the primiventral setae corresponding to Tokukobelba occurs in Belba ( Protobelba ) ( N or - ton , 1979c), Weigmannia ( M iko & N orton , 2010) and also Lanibelba N orton , 1980, based on its original description. Setal ontogenies in Belba ( Belba ) ( G randjean 1936, N orton & P alacios - V argas 1982, E nami 1994, W ang & N orton 1995, T olstikov 1996, T olstikov & L yashchev 1996, B ayartogtok h 2000, 2004a), Metabelba ( M ourek et al. 2011a), Po­ robelba ( M iko 2008) and Acanthobelba E nami & A oki , 1993, based on its initial description and on C hoi 1997b) contrast with Tokukobelba in that on the tarsus I of leg I of these, with very rare exceptions, pl´´, and particularly pv´ and pv´´, experience only a minimal distal migration and all are inserted on the bulb in the adult. This pattern is also known from many other Brachy­ pylina, including species within the Autogneti- dae G randjean , 1960, ( G randjean 1960b, 1960c, B ayartogtokh 2012), Staurobatidae G randjean , 1966, ( G randjean 1966), Damaeolidae ( G rand - jean , 1965a) and Hungarobelba ( M iko & T ravé , 1996). Compared to this ancestral state the one in Tokukobelba is moderately derived and that of Damaeus highly derived. Spinae adnatae Spinae adnatae are found in approximately two thirds of the presently known species of Damaei- dae.Other than in Tokukobelba and Belba ( Belba ) ( B ayartogtokh 2000), the structures are absent in Dameobelba S ellnick , 1929, ( G randjean 1955, N orton 1977a, pers. obs.), Caenobelba ( N orton , 1979c), Subbelba B ulanova - Z achvatkina , 1967, ( S kubala 1992), Neobelba B ulanova - Z achvatkina , 1967, ( B ulanova - Z achvatkina 1975, p. 121, 131), Metabelba ( B ulanova - Z achvatkina 1965, M ourek et al. 2011a), Mirobelba P érez - Í ñigo & P eña , 1994, and Parabelbella ( B ulanova - Z achvatkina 1975, M iko et al. 2011). Belba ( Protobelba ) possesses spinae adnatae ( N orton 1979c). Spinae adnatae are variably present or absent within Dyobelba ( N orton 1979b, N orton & R yabinin 1994, B ayar - togtokh & N orton 2007), Epidamaeus ( N orton