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86 Carolinea 74 (2016) 1979b, L yashchew & T olstikov 1993, M iko et al. 2011), Porobelba ( S ubías 1977, M iko 2008), Damaeus ( Tectodamaeus) ( A oki 1984, W ang & C ui 1994, L u & W ang 1995, X ie & Y ang 2009), and Metabelbella ( B ulanova - Z achvatkina 1967, 1975, A rillo & S ubías 2006, E rmilov & K haustov 2011, E rmilov , S htanchaeva & S ubía s 2012). Only exceptionally rarely are spinae adnatae present in Brachypylina other than the Damaei- dae. They are seen, however, in a few isolated members of the Ameroidea such as Hunga­ robelba ( N orton 1977a, M iko & T ravé 1996), Costeremus ornatus ( A oki 1970) and C . cornu­ tus ( W ang & C ui 1996) within the Damaeolidae, and also Roynortonia E rmilov , 2011, in the Am- erobelbidae G randjean , 1961, ( E rmilov 2011). In the overwhelming majority of Ameroidea spinae adnatae are absent, but in many, including the Ameridae B ulanova - Z achvatkina , 1957, ( C hen et al. 2004), Heterobelbidae B alogh , 1961, ( B eck 1962), Eremulidae G randjean , 1965, ( P érez - Í ñigo 1997), Amerobelbidae ( G randjean 1961), Staurobatidae ( G randjean 1966), Ctenobelbidae G randjean , 1965, ( M ahunka 1974, 1977, W oas 1986, B aran 2012) and Rhynchoribatidae B a - logh , 1961, ( B alogh 1970, M ahunka 1985, W oas 1986), humeral condyles are often visible in an analogous position. N orton (1977a) sees the spinelike spinae ad- natae as derived from such humeral tubercles and (1977a, 1979a) assumes their presence to be plesiomorph for the Damaeidae. However, it is quite possible that spinae adnatae arose one or more times within the Damaeidae and that the character state of Tokukobelba may represent the ancestral one. Spinae adnatae appear to have been gained or lost independently or in par- allel in several lineages or clades of Damaeidae. Nevertheless their presence is a very strong indi- cator that a mite does belong to the Damaeidae. Tokukobelba misses this major family character. Propodolateral apophyses An apophysis P is generally absent in Tokuko­ belba (Fig. 1), although C hoi & N amkoong (2002, p. 27, Fig. 4) depict what appears to be a minute tip-like apophysis P for T sellnicki , and is also not detected in any species of Belba except B . cor­ nuta W ang & N orton , 1995, B . sarvari T olstikov , 1996, and B . flammeisetosa T olstikov , 1996, based on their original descriptions. The apophy- sis P is missing in Quatrobelba ( N orton 1980), Dameobelba ( N orton 1977a), Porobelba ( M iko 2008), Caenobelba ( N orton 1979c), Subbelba ( S kubala 1992), Acanthobelba ( E nami & A oki 1993, C hoi 1997b), Mirobelba ( P érez -Í ñigo & P eña 1994), Dasybelba ( W oolley & H iggins 1979) and Lanibelba ( N orton 1980). The structure is variably present or absent within Epidamaeus ( N orton 1979c, 1979d, L yashchew & T olstikov 1993), Metabelba ( M ourek et al. 2011a), Belbo­ damaeus ( B ayartogtokh 2004a) and Dyobelba ( N orton 1979c, 1979d, W ang & N orton 1993, E nami & A oki 2001). Within the latter genus it is not even a constant feature of the otherwise morphologically very ho- mogeneous and clearly monophyletic Dyobelba tectopediosa species group ( B ayartogtokh & N or - ton 2007). An apophysis P is seldomly encoun- tered outside of the Damaeidae, but is known, for example from Pheroliodidae P aschoal , 1987, ( G randjean 1964) and the genera Veloppia H am - mer , 1955, ( N orton 1978b, C hen & W ang 2002), Hungarobelba ( N orton 1977a; M iko & T ravé 1996) and Costeremus ( A oki 1970, F ujikawa & F ujita 1985, W ang & C ui 1996, C hoi 1997a). N orton (1979a), following G randjean (1960a), ar- gues that the apophysis P is derived from the pe- dotectum I. The latter is commonly found in pyc- nonotic Oribatida ( W oas 2002). Pedotecta I are auriculate or scale-like structures projecting from the body wall immediately posterior to acetabu- lum I, whereas the apophyses P are toothlike or ceratiform projections between legs I and II which are not directly adjacent to an acetabulum ( G randjean 1960a, N orton & B ehan - P elletier 2009). N orton (1977a, 1979a) assumes the presence of an apophysis P to be the ancestral state for the Damaeidae. If so, the character state of absence of apophysis P appears to have been subject to considerable convergent or more likely parallel evolution within the Damaeidae. Anterolateral apophyses Both anterior Ala and posterior anterolateral apo- physes Alp are present in Tokukobelba compta (Figs 1, 3). The simultaneous occurrence of both of these structures, which may be attributes of other species in the genus, as they are very easy to overlook, has amongst Damaeidae so far only been noted for Metabelba denscanis by M ourek et al. (2011a). Due to the strongly laterally dis- placed bothridia and apophyses Ba of this spe- cies, the homology of its anterolateral apophyses with those in Tokukobelba is somewhat unclear, but very likely. A clear homologue of the apophy- sis Ala of T compta is found in four species of the Dyobelba tectopediosa species group, based