Carolinea 74

82 Carolinea 74 (2016) with Tokukobelba , including the presence of pro- dorsal apophyses Aa and Ap, absence of spinae adnatae, femora III and IV with 3 and 2 setae, respectively, and an epimeral setation of 3-1-3-3. Costeremus barbatus is mentioned as possess- ing 10 notogastral setae ( C hoi 1997a). Based on the text of the original description ( F ujikawa & F ujita 1985), C . yezoensis similarly seems to differ from Tokukobelba in the presence of only 10 notogastral setae. The illustrations in C hoi (1997a) and F ujikawa & F ujita (1985), however, distinctly show the presence of 11 notogastral se- tae in both of these species. The leg chaetotaxy of C barbatus is not known, while C yezoensis is characterized by a tibial associated setal formula of 1-1-1-1. Tokukobelba differs from Costeremus ornatus A oki , 1970, the type of its genus and from C cor­ nutus W ang & C ui , 1996, in: 1) ratio of distance between insertion points of prodorsal setal pairs ro-ro to la-la ranges from 1:1 to 1,3:1 instead of 4:1; 2) notogastral setae setae c2, la, lm, lp insert more or less in a line along the notogastral mar- gin instead of presence of a conspicuous cen- trodorsal setal pair, inserted very close together; the ratio of distance between the insertion points of the setal pairs la-la or lm-lm to that between those of the centrodorsal setal pair, termed dp by W ang & C ui (1996), is between 3:1 to 4:1; 3) anterior prodorsal ridges do not extend to inser- tion points of rostral setae; 4) notogastral and adanal setae not short, with long barbs and a stellate appearance; 5) anteriormost genital seta not hypertrophied and not 3x length of setae g3, g4; 6) substantially longer legs; 7) notogaster posteriorly without a distinct protuberance; 8) bothridium without spinelike processus; 9) hu- meral processes or spinae adnatae absent; 10) absence of two pairs of ridges on the anterior of the notogaster, posterior to the structures called first notogastral ridges by both A oki (1970) and W ang & C ui (1996), but which I interpret as being humeral processes or spinae adnatae instead. Neither C yezoensis nor C . barbatus or the re- cently discovered C . stebaevae B ayartogtokh & E rmilov , 2015, display any of these ten attributes in which the type species of Costeremus A oki , 1970, differs from Tokukobelba , and I consider their inclusion in Costeremus most uncertain. I agree with B ayartogtokh & E rmilov (2015) in lo- cating these 3 species in the Hungarobelbidae, however, based on our current knowledge about them. Like Hungarobelba B alogh , 1943 they may be considered proto-damaeids, and their phylo- genetic relationship to Tokukobelba requires in- vestigation. 6 Comparative morphology and evolutionary systematics of Tokukobelba In this section I will comment in depth on sev- eral of the extraordinary traits characterizing the new genus and also on those characters which it shares with Belba . This is followed by an at- tempt to shed some light on the position of Toku­ kobelba within the Damaeoidea. Trochanteral setation Tokukobelba shares a leg trochanteral seta- tion formula of 1-1-2-1 with the great majority of Damaeidae, including all representatives of Belba except B . californica ( B anks , 1904), B . ros­ sica B ulanova - Z achvatkina , 1962, and B . crassi­ setosa B ayartogtokh , 2000, in which the seta- tion is 1-1-2-2 based on the original descriptions and the redescription of B . californica by N orton (1979 c ). A trochanteral setation of 1-1-2-1 char- acterizes numerous basal Brachypylina ( W oas 2002) and appears to represent the ancestral state for the Damaeidae and also for the Brachy­ pylina as a whole. Higher setal counts appear to have evolved independently within many families of higher Oribatida. Femoral setation Tokukobelba is characterized by a strongly re- duced femoral setation with 3 setae usually found on femur III and 2 on femur IV (Figs 6-7). Here setae d, ev´ and l´ are developed on femur III, while on femur IV only setae d and ev´ occur. Compared to Belba corynopus , ventral setae v´ and v´´ are both absent on femora III and IV, with seta l´ additionally missing on the femur of leg IV. In the Damaeidae setae d and ev´ are found on femur III in the larval stage, while l´ is deutonym- phal in appearance on this segment, whereas in sharp contrast setae d and ev´ are deutonym- phal in appearance on femur IV ( N orton 1977b). Ventral setae in the Damaeidae are added in the tritonymph or more often in the adult, as in B . corynopus and B . sculpta (pers. obs.). In the ontogeny of Tokukobelba no regression or loss of femoral setae occurs. Instead the ventral setae on femora III and IV, and seta l´ on femur IV are simply never formed. Such a low femoral setal count is unique in Damaeidae where reli- ably known, although Dasybelba perona W ool -

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