ELLIOTT ET AL.: THE CAVE FAUNA OF CALIFORNIA 
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genera and 7 species of cavemicoles, which includes three troglobites in Calileptoneta. Unlike 
telemids, which are restricted to the Californian region, leptonetids range into the eastern Nearctic, 
with many troglobitic species in Texas ( Tayshaneta ) and the Appalachians (Appaleptoneta). 
The second group with troglophilic spiders are the orbicularians, orb weavers and their rela¬ 
tives, and is also represented by eight fa mi lies of cavemicoles, but with more species (40). All of 
the families have troglophilic representatives but only two, Linyphiidae and Nesticidae, have 
troglobites. Linyphiidae are well-known as troglobites, especially the numerous species of Troglo- 
hyphantes in the Mediterranean region, as well as several genera with troglobitic species in 
Appalachia. Troglobitic linyphiids (Figs. 38-39) are very rare in California, mostly discovered in 
recent surveys in the Sierra Nevada, and known by four undescribed species belonging to different 
genera. By contrast, California’s Nesticidae are well known and the three species of Nesticus all 
occur in caves, but only N. potterius is troglobitic; the other two species (Figs. A0-A\) are 
troglophiles, with one mostly epigean in distribution (N. silvestrii ) and the other restricted to caves 
( N. sodanus ). The family is common in caves globally and troglobitic species occur in both the 
Appalachian region, represented by a different species group of Nesticus, and in Texas, by Eid- 
mannella. The families Pimoidae (Fig. 42) and Tetragnathidae include strongly troglophilic 
species, but are without troglobites in California. The tetragnathid genus, Meta, occurs in caves 
across the Holarctic and is known in California by Meta dolloff (Figs. 43-44), a species showing 
little troglomorphic modification but nonetheless appears to be cave restricted. 
The third group includes spiders related to funnel weavers, the Amaurobiodea and Dic- 
tynoidea, or amauro-dictynoids, and is represented in California by 38 species of cavemicoles in 
five families, with 9 species of troglobites in two, Cybaeidae and Dictynidae. Cybaeidae (Fig. 45) 
are widespread in the Nearctic but have their greatest diversity in the Pacific NW, especially at the 
Califomia-Oregon border where the three known troglobites occur (in Cybaeus and Cybaeozyga ). 
By contrast, cavemicolous Dictynidae (Figs. 46—47) are widespread in the Nearctic, in California 
by 17 species, mostly troglophilic, but with 6 troglobites in Blabomma and Yorima, both genera 
endemic to the Californian region (Ubick et al. 2017). Troglobitic dictynids also occur in the Texan 
and Appalachian karst, but in California the genus Cicurina is only troglophilic. In addition to these 
families, troglophiles also occur in Amaurobiidae and Hahniidae, especially the very common 
Calymmaria species (Fig. 48) with their conspicuous webs near cave entrances. 
Related to this group are the lycosoid spiders, which are typically large-eyed hunting spiders 
and only incidentally associated with caves. One exception is Zoropsidae, where the several 
species of Titiotus (Fig. 49) and Anachemmis are common troglophiles throughout the Californian 
karst. Although no troglobites are known from these genera, at least two species (of Anachemmis) 
show significant troglomorphy and are known only from caves. 
The two large groups of spiders not represented by troglobites are, interestingly, the most 
primitive and most derived. In the former group are mygalomorphs, sister to all remaining spiders 
(araneomorphs), with only one troglophilic genus, Calisoga (Fig. 50), and three species (3 fami¬ 
lies) of accidentals. Troglobitic mygalomorphs do occur in southern latitudes, such as Mexico. 
Even more poorly represented in caves are the Dionycha, 2-clawed hunting spiders that have aban¬ 
doned web-building and successfully radiated into a dozen families and thousands of species, but 
whose presence in Californian caves are 6 species (5 families) of accidentals. 
The cavemicolous arachnid fauna of California agrees well with that of the other major karst 
areas in the Nearctic, those of Texas and the greater Appalachians. Gertsch (1992) tabulated the 
spider troglobites of the three regions and recorded 20 from California, 18 from the Appalachians, 
and a whopping 61 from Texas, of which 46 are blind Cicurina species. Currently, by our count, 
there are 75 troglobitic spiders, including 53 Cicurina. The three regions also share troglobite taxa 
