Carolinea 74

60 Carolinea 74 (2016) to Norway ( G abrielsen et al. 1997). Tokukobelba compta has further been noted to occur in Po- land ( K ulczy ń ski 1902a, S ellnick 1920, W illmann 1931, 1939, 1956, S trenzke 1952, D ziuba & S ku - bala 1987, O lszanowski et al. 1996), Romania ( V asiliu et al. 1993), Russia ( K rivolutsky 1995, S idorchuk 2009, Z enkova et al. 2011), includ- ing the Kaliningrad Oblast ( S ellnick 1920), the Czech Republic ( S tarý 2013), Spain ( S ubías & S htanchaeva 2012), Sweden ( T rägårdh 1910, F orsslund 1943, W illmann 1943, D alenius 1960, L indberg & P ersson 2004, H onciuc & L undqvist 2009, R emén 2010) and Switzerland ( S trenzke 1952, S chweizer 1956). Tokukobelba verrucosa is known from Finland ( N iem i 1988, K oponen 1989, L aakso et al. 1995), Norway ( S eniczak et al. 2006, H ågvar et al. 2009), the Czech Republic ( S tarý 1994) and Pakistan ( H ammer 1977). The species shows a broad dis- tribution in Russia, ranging from Kandalaksha in the Murmansk Oblast and Arkhangelsk to Mos- cow, the Ural region, Eastern and Western Sibe- ria and the Russian Far East ( B ulanova -Z achvat - kina 1962, 1967, 1973, 1975, K rivolutsky 1995, K olesnikova et al. 2005). It has also been detect- ed in Dagestan ( G azaliev 2011). In China records for Tokukobelba verrucosa exist for the provinces Jilin, Heibei and Beijing ( W ang & N orton 1995, W ang et al. 2000, C hen et al. 2010). Tokukobelba farinosa has only been found in Lapland province in Northern Sweden ( T rägårdh 1902, 1910). Tokukobelba s ellnicki occurs in the Ukraine ( K arppinen et al. 1992), and is apparently rela- tively common in Russia with records from for example the Moscow region ( K rivolutsky & L eb - edeva 2004) and the Russian Far East ( B ulanova - Z achvatkina 1962, 1967, 1973, 1975, K rivolutsky 1995). C hoi & N amkoong (2002) describe the spe- cies from South Korea. In China T sellnicki has so far been only been discovered in the north- ern Liaoning and Beijing provinces bordering on Mongolia ( W ang & N orton 1995, A oki et al. 1997; W ang et al. 2000, C hen et al. 2010). Tokukobelba mongolica has only been recorded from cen- tral and northern Mongolia ( B ayartogtokh 2000, 2004a), while Tokukobelba japonica is known from Japan ( A oki 1984, K osuge 2005, I to et al. 2007, H arada et al. 2008, H asegawa et al. 2013) and Taiwan ( A oki 1995). Tokukobelba barbata has been collected in Nayoro on the Japanese island of Hokkaido ( F ujita & F ujikawa 1986) and on Hon- shu ( I to et al. 2007). Tokukobelba itsukiensis is only known from its type locality, the Itsuki Village on Kyushu Island, Japan ( F ujikawa 2011). Ecology Tokukobelba compta has frequently been found in mountainous regions such as the Alps ( S chwei ­ zer 1956, S chmölzer 1994, S chatz 1978, 2008), the Scandinavian mountains ( H eggen 2010), the Carpathians (K ulczy ń ski 1902a, S trenzke 1952), the Köszeg mountains in Hungary ( B a - logh 1943) and the Krkono š e mountain range in the Czech Republic ( S tarý 1994, 2005). Several authors such as F ranz ( 1943 ) and M iko (2006a) have assumed the species to predominantly oc- cur on high mountains. K ulczy ń ski ( 1902 a) him- self found the species not only at a subalpine site at an altitude of 2.000 m a.s.l. close to the summit Kozi Wierch, the highest mountain in Po- land, but also at a low altitude of about 450 m a.s.l. at Smreczyna in the Tatra region. T compta has subsequently often been encountered in low-lying areas such as the North European Plain ( S ellnick 1920, S trenzke 1952, W arncke et al. 1991), and the East European Lowlands ( K againis 2011). Tokukobelba verrucosa has been noted to occur on elevated areals in Pakistan at 2.600 m above sea level, but still well below the tree line ( H ammer 1977) and on glacier forelands in Fennoscandia ( S eniczak et al. 2006, H ågvar et al. 2009) but also is very common in regions of moderate to low elevation such as in the Russian White Sea area and the Moscow region ( B ulanova - Z achvatkina 1962, K rivolutsky 1995). B ayartogtokh (2000) initially collected T mongolica from litter of co- niferous larch and fir taiga forest at an altitude of 1.950 m a.s.l. but later found the species in a much lower situated grassland habitat ( B ayartog - tokh 2004a). The type localities of both T. japoni­ ca and T barbata lie at low elevations, relatively close to sea level ( A oki 1984, F ujita & F ujikawa 1986). The first of these species has subse- quently been encountered at altitudes ranging from 300 m to 960 m by H arada et al. (2008) but it has also been recorded from high altitude for- est at Nanhu Mountain, one of the highest peaks of Taiwan ( A oki 1995). Tokukobelba barbata was found at 1.200 m a.s.l. by I to et al. (2007). Tokukobelba therefore often occurs in montane to alpine habitats but is certainly not restricted to these. It is perhaps noteworthy in this con- text that in Alpine Fennoscandia T compta is found along the entire altitudinal gradient rang- ing from sub-alpine to high alpine zones without any apparent distributional preferences ( H eggen 2010 ). S chmölzer (1962) and S chatz (2008) view T. compta as most likely being a preglacial relict