ELLIOTT ET AL.: THE CAVE FAUNA OF CALIFORNIA 
37 
where they feed on detritus. At lower elevations they can live in lava tubes that produce perennial 
ice, and almost all California ice caves that have been studied contain Grylloblatta. In California 
Grylloblatta occurs in caves from the Lava Flows North to the Sierra Nevada South, Tulare Coun¬ 
ty, where Mays Cave is the second-most southern record for any grylloblattid to date (Lee 1967). 
Grylloblatta marmoreus is restricted to caves and snow fields in the Marble Mountains. 
G. gurneyi is restricted to lava tube caves in or near Lava Beds National Monument, with the pos¬ 
sible addition of a few records on the ice fields of Mount Shasta. One new species of Grylloblatta 
(Fig. 74) is found only in the caves and snow fields of Sequoia National Park. Another species, 
G. chandleri, is widespread in the Modoc Plateau and Basin Ranges of Eastern California (Kamp 
1963, 1973). All are glacial relicts from times when montane glaciers descended to lower eleva¬ 
tions than now (Schoville and Graening 2013). 
Beetles (Order Coleoptera) 
Worldwide, one of every five species is a beetle (Figs. 75-81, 84). They are easily the largest 
group of animals in California. Cave Coleoptera comprise a number of families, most of which 
happen to be found in caves incidentally. Others are both habitual and occasional trogloxenes often 
being found in caves. A number of troglophiles exist with an even smaller number of troglobites. 
Many take advantage of the sometimes rich organic matter that is brought in either from under¬ 
ground streams or by bats, birds, or mammals often associated with caves. California cave beetles 
are poorly known as only a few publications have concentrated on caves or have included cave bee¬ 
tles as part of a larger study. 
Most troglobitic cave beetles in California only recently have been formally described. In our 
experience the general diversity of beetles in the western United States is higher than the eastern 
U.S.A., and so far western cave beetles are about as diverse as in the east. According to the latest 
taxonomic studies, some of which are not yet published, the diversity of California’s troglobitic 
beetles may eventually prove to be higher than in the east. 
In California, cave Coleoptera can be divided into three groups: 
1. Accidentals or incidentals, which contain families such as Bostrichidae, Cantharidae, 
Carabidae, Coccinelidae, Cerambycidae, Chrysomelidae, Dytiscidae, Elateridae (adults), Elmidae, 
Gyrinidae, Hydrophilidae, Lampyridae, Meloidae, Melyridae, Monotomidae, Psephenidae, Scir- 
tidae, Scydmaenidae, Trogidae and Zopheridae. Scaphinotus sp., a snail-eating carabid beetle, is an 
example of an accidental or incidental species. One unusual discovery in this group was finding the 
fungus-feeding Usechimorpha montanus, a rare species found at much higher elevations, deep in 
Sutter Creek Cave, Amador County, at a much lower elevation of about 707 m. 
2. Trogloxenes, which include the families habitually found in caves, such as Agyrtidae, some 
Carabidae, Cleridae, Cryptophagidae, Curculionidae, Elateridae (larvae), Dermestidae, Histeridae, 
Lathridiidae, Phengodidae, Ptiliidae, Scarabaeidae, Silphidae, and Staphylinidae. Trechus sp. 
(Fig. 75) is a trogloxenic or troglophilic carabid, member of the Tribe Trechini, with important cav- 
emicoles in the eastern U.S.A. The discovery of the carabid (ground beetle), Amblycheila 
schwartzi, prompted a study of the entire ecosystem of Mitchell Caverns. Someone found a “rare 
large tiger beetle” in the cave (tiger beetles are now classified in the Carabidae as the Subfamily 
Cicindelinae). This species is neither rare nor limited to caves, and it is often encountered at night 
in the Mojave Desert’s Providence Mountains, San Bernardino County, and other eastern Califor¬ 
nia ranges, or in caves. The staphylinid subfamily Pselaphinae, the ant-like litter or mold beetles 
(Fig. 76), is a group with important troglobites in Texas and the eastern U.S.A. Two species of tene- 
brionid (darkling) beetles in the genus Schizillus are found in the Providence Mountains: 
S. laticeps and. S. nunenmacheri. The latter has more elongate appendages and reduced eyes 
