28 
PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES 
Series 4, Volume 64, Supplement 1 
from the Sierra Nevada North, and an undescribed species from Tulare County, Sierra Nevada 
South. A rhagidiid was photographed in a cave at Lava Beds National Monument (Fig. 18). 
Harvestmen (Opiliones) are common cavemicoles in California, with the exception of the 
most primitive members, the mite-like Cyphophthalmi, which have not yet been recorded from 
Californian caves, although ranging in proximity of karst areas. Nine representative harvestmen of 
six families are illustrated in Figures 19-27. The Eupnoi, by comparison, are the most conspicuous 
cave harvestmen, especially the long-legged Leiobunum species (Fig. 19), or “Daddy longlegs,” 
sometimes referred to as “cave hair” because of their dense aggregations. Members of this subor¬ 
der are trogloxenes and not cave restricted. The Dyspnoi are moderate-legged forms, represented 
by several probably troglophilic species in five genera, but with only a single troglobite in 
Taracus, T. fluvipileus, which is the only troglobitic arachnid from lava tubes and is characterized 
by much longer appendages than its troglophilic relatives, such as T. audisioae (Fig. 20). The Lani- 
atores family Phalangodidae, our shortest-legged harvestmen, is most diverse in the Californian 
region and is richly represented in caves. Of the 23 species (6 genera) so far recorded, 10 are troglo- 
bites, with eight in Banksula (Figs. 21-22), an endemic of Sierra Nevada North, and one each in 
Calicina (Fig. 23) from Tulare County, Sierra Nevada South, and Texella from Inyo County, 
Mojave Desert. Maps in Figures 7 and 8 illustrate the highly insular distribution of Banksula in the 
Sierra Nevada North region. Each of these genera also includes at least one troglophilic species 
known only from caves and likely cave restricted. The remaining eight troglophiles are of species 
with largely epigean distributions. With the exception of Calicina, which is the basal lineage of 
Phalangodidae, all other troglobites (including all Texan and eastern U.S.A. phalangodids) belong 
to the same, derived lineage, the bifurcate clade. The intermediate species, the sitalcinoid complex, 
are represented by several genera and many epigean species in California, but with only a few 
troglophiles in Sitalcina (Fig. 24) and Megacina (Ubick 2007). 
Pseudoscorpions have the highest cave diversity, with 23 troglobites distributed mostly among 
the Neobisiidae (10) and Chthoniidae (7), followed by Pseudogarypidae (2), Chemetidae (2), Lar- 
cidae (1), and a species not placed to family. Most of the troglobites are in the Sierran karst (17), 
and the two species in the Santa Cruz Karst are the sole representatives of troglobitic arachnids in 
the Bay Area/Delta Region. The highest diversity of U.S. troglobites is in the Interior Low Plateau, 
which has an astonishing 62 species (Culver et al. 2003), although the pseudogarypid troglobites 
are strictly Californian. Eight representative pseudoscorpions of four families are shown in Figures 
28-35. 
Spiders are very common cavemicoles in California (Tables 2 and 3) and about 145 species 
are recorded from caves. Most of these species are troglophiles (79) and troglobites (19), and the 
remainder largely accidentals (27). Cavemicoles are recorded in 35 families, or just over half of the 
61 families known for the region; 17 families include troglophiles and troglobites, 12 are repre¬ 
sented only by accidentals. Troglobites are found in six families which represent three major 
spider groups. Twenty representative photographs of 15 families are shown in Figures 36-55. 
The first spider group contains the haplogyne families, which are at the basal branches of tme 
spider phylogeny (Araneomorphae), and are extremely well represented in caves with troglobites 
recorded from most of the families. In California, haplogyne cavemicoles are known in 8 families 
(29 species), mostly troglophiles, but with four troglobites recorded for Telemidae and Leptoneti- 
dae, both small spiders that constmct delicate sheet webs. Telemids are represented by Usofila 
(Figs. 36-37), a wide-ranging genus in the western Nearctic of mostly epigean species, but with 
troglobites known only from California. The genus is in need of revision as the currently recorded, 
undescribed troglobite is widely-distributed and probably represents a species complex which has 
yet to be resolved (Ledford and Griswold 2010b). Leptonetids are represented in California by two 
