ELLIOTT ET AL.: THE CAVE FAUNA OF CALIFORNIA 
41 
Santa Cruz County. Calasellus longus occurs in one spring in Fresno County. Calasellus sp. occurs 
in 12 groundwater sites and Pinnacle Point Cave, Tuolumne County. Bowmanasellus is a genus 
endemic to California with its only species, B. sequoia known only from 8 caves and springs in 
Sequoia National Park. 
The crustacean Order Bathynellacea (Fig. 91) live interstitially in groundwater. Bathynel- 
laceans are minute, blind, worm-like animals with short, primitive legs, reaching a maximum 
length of 3.4 mm. Since 1997, a rich fauna of 10 species in two families has been discovered in the 
hyporheos, springs, and streams in the coastal zone of Los Angeles, San Diego, and Marin coun¬ 
ties. None have yet been found in caves. 
We examined high-biodiversity groundwater sites by tabulating the number of phreatobites per 
site and their single-site endemic scores (SSEs). Table 9 details 12 high-biodiversity groundwater 
sites, in which seven regions are compared. A site score was calculated by multiplying phreatobites 
by SSEs (single-site endemics). A regional score was summed from the site scores in each region. 
The top regional score of 24 is Sierra Nevada North because of Lake Tahoe’s 4 phreatobites and 
2 SSEs. The Coast Ranges ranked second with 7 sites and a regional score of 16; Bathynellaceans 
weighed heavily there. Bay Area/Delta ranked third with five sites and a regional score of 10 with 
a variety of species and sites. These scores are for general comparisons only, but it is apparent that 
the pattern of phreatobite biodiversity differs from that of caves in California (see below). 
Herpetofauna 
California supports a diverse array of amphibians and reptiles consistent with its latitude, com¬ 
plex geology, and diverse climatic regimes. Our numbers are approximate because of taxonomic 
flux: 14 genera and 64 species of native amphibians and 42 genera and 87 species of native, non¬ 
marine reptiles recorded from the state (Stebbins and McGinnis 2012). Despite this diversity, rela¬ 
tively few species have been reported from caves and related habitats. Records reported herein are 
limited to 13 species of salamanders in 6 genera, 3 species of anurans (frogs and toads) in 3 gen¬ 
era, one species of turtle, 5 species of lizards in 5 genera, and 8 species of snakes in 7 genera. 
All of these records likely represent individuals of the local epigean population that become 
trapped in caves or use caves for behavioral thermoregulation. With the notable exception of sala¬ 
manders, cave-restricted species of amphibians and reptiles are extremely rare or nonexistent glob¬ 
ally. Reptiles in particular, are precluded from occupying most cave environments due to low tem¬ 
peratures, which cause torpor. This is especially true in California where the maximum cave tem¬ 
perature is 18°C or less, typically much less. This is at the lower limits or below the preferred active 
temperature requirements of California reptiles. There is use of cave entrances as hibemacula by 
reptiles, a phenomenon widely observed in caves in many areas. Some of the records reported here¬ 
in likely represent hibernation sites. Most reptile records, however, represent transient animals near 
entrances or animals trapped in caves. 
Amphibians often tolerate lower temperatures and, therefore, have more potential to exist in 
temperate cave environments. There is no evidence of troglophilic or troglobitic adaptation to a 
cave environment among California species. Anurans (frogs and toads), due in part to their jump¬ 
ing locomotion and visual prey detection, are much less able to maneuver and thrive in deep cave 
habitats (Graham 1962d; Lannoo 2005). The primarily herbivorous diet of anuran larvae also poses 
a significant obstacle to the evolution of cave-adapted anurans. Consequently anurans are, like rep¬ 
tiles, generally lacking in cave environments except as transient animals from epigean populations. 
An exception would be populations seasonally restricted to a moist cave environment, typically 
near entrances, in a generally more arid surface environment, such as Pseudacris sierra (formerly 
Hyla regilla ), the Sierran treefrog (formerly Pacific treefrog, Fig. 92). Graham (1962d, 1963a) 
