Carolinea 74

L amos : Tokukobelba gen. nov. (Acari: Oribatida: Damaeidae) 83 ley & H iggins , 1979, D . aphelesa W oolley & H ig - gins , 1979, and Tectodamaeus cordatus X ie & Y ang , 2010, have been described with a femoral setation formula of 5-5-3-3, based on the origi- nal descriptions. That the femoral setation of Tokukobelba is clearly more primitive than in all other Damaeidae was already noted many years ago by N orton (1977b). Indeed, it is reminiscent of numerous other basal brachypyline groups ( W oas 2002). Genual setation The genua I-III of Tokukobelba show a solenid- ion and a companion seta d originating from a shared alveolus with the solenidion being dis- tal to the seta. A solenidion is absent on genu IV but seta d is present. Tokukobelba shares a genual associated setal formula of 1-1-1-0 with all damaeids except three species of Damaeus K och , 1835, in which more than one seta d is absent on genu I-IV ( N orton 1977b, P érez - Í ñigo 1987, M iko 2006a). Among the Damaeidae with companion setae d on genua I-III the great ma- jority share a genual setation of 4-4-3-3. N orton (1977a) in his PhD considered a genual associ- ated setation of 1-1-1-0 to be plesiomorph for the Damaeidae. This is indeed very probable. The genual setation of 4-4-3-3, shown by both Belba and Tokukobelba is likely to represent the ances- tral character state. The larval seta d on genua I-III of Tokukobelba is extremely small, smooth and very difficult to perceive (Fig. 16). Tokukobelba shares this char- acter state, which is probably derived within the Eupheredermata ( N orton 1977a) with all mem- bers of its family except Quatrobelba N orton , 1980, where the seta is almost as long as the solenidion and barbed ( N orton 1980). Tibial setation Tokukobelba barbata is described as possess- ing an associated seta d on tibia I by F ujita & F ujikawa (1986), and the authors clearly illustrate this (p. 8, Fig. 2b). A oki (1984) mentions the pres- ence of 4 setae and 2 solenidia on the tibia of leg I of T japonica but his illustration of leg I (p. 113, Fig. 6c) shows the presence of 5 setae and the solenidia φ 1 and φ 2. In addition to setae v´, v´´, l´ and l´´ a short seta is depicted which in its proximal half appears to lie directly proximal to solenidion φ 1 in the antiaxial perspective shown. Although the insertion point of this seta is not vis- ible, its size and location match that of the as- sociated seta d of T . barbata . K ulczy ń ski (1902a, tab. IV, Fig. 65) similarly portrays a seta d asso­ ciated with the solenidion φ 1 on the tibia of leg I. In all specimens of T . compta , in which the tibial leg setation was examined in detail by me (n = 12), a short, very thin seta d was seen to be present on the tibia of leg I (Fig. 4). This was cou- pled to solenidion φ 1. The thickness and relative size of the seta d shows a very close correspond- ence to that drawn by K ulczy ń sk i (1902a), A oki (1984) and F ujita & F ujikawa (1986). An associ- ated seta d on tibia I has not been discovered for Russian or Chinese specimens of T. verru­ cosa and T sellnicki ( B ulanova - Z achvatkina 1962, 1967, 1975, W ang & N orton 1995), for T. mon­ golica ( B ayartogtokh 2000) or for T itsukiensis ( F ujikawa 2011). Possibly it is also present in these species as the very thin seta is extremely easy to overlook. A tibial associated setation of 1-1-1-1 is already found in basal Oribatida such as the Brachytho- nioidea T hor , 1934, ( M oritz 1976), the Pthirac- aroidea P erty , 1841, ( N iedbala 1992) and Cro- tonioidea T horell , 1876, such as Nothrus K och , 1836, ( O lszanowski 1996) and Camisia von H ey - den , 1826, ( C olloff 1993, O lszanowski 1996). Within the Brachypylina this pattern occurs in, for example, Tricheremaeus B erlese , 1908, ( W oas 2002), as well as in basal Circumdehiscentiae such as the Neoliodidae S ellnick , 1928, Plasmo- batidae G randjean , 1961, and the Hermanniel- lidae G randjean , 1934, if the tibial solenidia are present ( W oas 2002, N orton & B ehan - P elletier 2009, E rmilov et al. 2010b).Within the Ameroidea B ulanova -Z achvatkina , 1957, a tibial associated setal formula of 1-1-1-1 is so far apparently only documented for two species of Hungarobelba ( M iko & T ravé 1996) and for Costeremus yezoen­ sis ( F ujikawa & F ujita 1985). The presence of an associated seta on tibia I is exceptionally rare in Damaeidae besides Tokuko­ belba , having only once been noted by G randjean (1954) for some specimens of a single population of Porobelba spinosa ( S ellnick , 1920) from Italy. N orton (1979a) considered a tibial associated setal formula of 0-1-1-1 to be plesiomorph for the Damaeidae. The tibial associated setation of 1-1- 1-1 of Tokukobelba is evidently even more primi- tive than that, and the genus therefore lacks a major derived character state of the Damaeidae. Tokukobelba usually possesses setae d on tibiae II-IV which are quite short and thin; and at the same time displays a very elongate solenidion φ of tibia IV, about 5 times the length of its associ- ated seta d (Figs 6-7). This is unique in Damaei-

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