ELLIOTT ET AL.: THE CAVE FAUNA OF CALIFORNIA 
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new species in three new genera: Bdellozonium cerviculatum, Buzonium crassipes, Endere disora 
and Plumatyla humerosa (Cook and Loomis 1928; Loomis 1938,1943; Shelley 1997b). 
Springs were studied sporadically, starting perhaps with the discovery of the pebblesnail, 
Fluminicola seminalis, in springs and lava tube streams in the upper Sacramento River in 1840; in 
1912, the Modoc pebblesnail, Fluminicola modoci, and Lost Creek pebblesnail, Fluminicola new 
species #20, were discovered in Fletchers Spring, Modoc Co. (Hershler and Frest 1996). This was 
just a hint of the large diversity of springsnails that exist in springs of arid regions of California and 
neighboring states. All of these groups ( Pyrgulopsis , Pristinicola, Tryonia, Vorticifex, Juga, etc.) 
are now in danger of extinction due primarily to ground water depletion 
The first systematic bioinventories of hot springs in California were performed by the Harvard 
entomologist Charles Braes in the late 1920s, who contributed to the knowledge of these heat-tol¬ 
erant, or thermophilous, animals (Braes 1928, 1932). In 1931 entomologist Edward Essig discov¬ 
ered the California cave isopod, Calasellus californicus, in a well in Lake Co. (M.A. Miller 1933). 
In 1937, E. Chace collected the Natural Bridge snail, Megomphix californicus, in Shasta Springs, 
Shasta Co. (Smith 1960). 
California’s vast diversity of spiders continued to be uncovered, above and below ground. In 
1934, the sheetweb spider, Bathyphantes diasosnemis, was discovered in the lava tube, Subway 
Cave (Ivie 1969; Gertsch 1984). In a single trip to Potter Creek Cave in 1937, Ralph Chamberlin 
discovered three new spiders: Cybaeus septatus, the dwarf sheet spider, Hahnia sanjuanensis, and 
the sheetweb spider, Arcuphantes potteri (R.V. Chamberlin and Ivie 1942, 1943). Another spider, 
Titiotus humboldt, was discovered in Potter Creek Cave in 1950 (Platnick and Ubick 2008). 
In the 1940s, several important speleological events occurred. In 1946, Erwin Bischoff (San 
Francisco State College) published the first list of the known caves of California. The Sequoia 
National Park naturalist, Frank Oberhansley, published a field book on Crystal Sequoia Cave, 
Sequoia National Park, which contained the first inventory of the fauna of a California cave (Ober¬ 
hansley 1946). Philip Kratzsch, a master’s student at San Diego State University, published an eco¬ 
logical study of the bats of San Diego County (Kratzsch 1948). California’s first three chapters of 
the National Speleological Society were started, including the Southern California Grotto (co¬ 
founded by speleologist Bill Halliday) in 1948. In this same year, the Stanford Grotto was started, 
mostly by students of the Stanford Alpine Club and the Geological Sciences Department of Stan¬ 
ford University. For the next ten years, this grotto led a statewide survey of caves and made many 
collections of cave fauna, discovering several new species, including the Dolloff Cave spider (Meta 
dolloff) in Empire Cave (Danehy 1951b; Levi 1980). Edward Danehy, a co-founder of the Grotto, 
published the first checklist of the animals collected in California caves (Danehy 1951b, 1952). The 
Stanford Grotto published various other important studies of cave animals (e.g., Lange 1951). Co¬ 
founders Edward Danehy and Arthur Lange discovered a new genus and species of sideband snail 
(Speleodiscoides spirellum ) in a cave in Amador Co. (Smith 1957). In 1957, Allyn Smith, a mala- 
cologist at California Academy of Sciences, published a summary of the snails from California 
caves, relying heavily on collections by the Stanford Grotto (Smith 1957). He concluded that there 
were no cave-adapted snails in California, although a few species are restricted to limestone caves 
and talus in California. In the early 1950s, the Stanford Grotto, along with the Southern California 
Grotto and Santa Barbara College, made the first expeditions to caves of the Channel Islands (Orr 
1951b). 
After the Stanford Grotto disbanded in 1954, some of the members and their associates formed 
new organizations that continued speleological studies in California, including the Cave Research 
Associates (CRA, 1958-1973) and the Western Speleological Institute (WSI, 1959-1973). These 
two organizations published the journals “Cave Studies”, “Cave Notes”, and “Caves and Karst” 
