PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES 
Series 4, Volume 64, Supplement 1, pp. 1-311, 118 figs., 25 tables 
July 28, 2017 
The Cave Fauna of California 
William R. Elliott James R. Reddell 2 , D. Craig Rudolph 3 , G.O. Graening 4 , 
Thomas S. Briggs 5 , Darrell Ubick 5 , Rolf L. Aalbu 5 , Jean Krejca 6 , Steven J. Taylor 7 
1 Cave biologist {retired), Missouri Department of Conservation, 914 Bannister Drive, Jefferson City, 
Missouri, 65109 {Email:speodesmus@gmail.com); 2 Curator Emeritus of Cave Invertebrates, Texas Natural 
History Collections, The University of Texas at Austin, Texas 78712; 3 USD A Forest Service, Southern 
Research Station, 506 Hayter St., Nacogdoches, Texas 75965; 4 Department of Biological Sciences, 
California State University, Sacramento, 6000 J Street, Sacramento, California 95819-6077; 5 Department 
of Entomology, California Academy of Sciences, San Francisco, California 94118; 6 Zara Environmental 
LLC, 1707 W FM1626, Manchaca, Texas 78652; 1 Illinois Natural History Survey, Prairie Research 
Institute, University of Illinois at Urbana-Champaign, Champaign, Illinois 61820-6953 
Contents 
Abstract.1 
Introduction.2 
Geology and Cave Regions .3 
Literature Review and History.15 
Methods .23 
Overviews of Fauna.25 
Biogeography and Biodiversity.66 
Ecology.75 
Conservation.94 
Discussion.99 
Acknowledgments .101 
Literature Cited.103 
Appendix 1. Annotated Species List .149 
Appendix 2. Site Index.286 
Appendix 3. Glossary.296 
Index to taxonomic and major geographic features discussed in Section 1 (pp. 1-103) . 301 
Hidden biodiversity is revealed in this study of California’s subterranean fauna, 
which contains distinctive elements that differentiate it from other North American 
regions. Since 1975, the rate of discovery of new species has accelerated with funded 
projects in most of the important cave areas of the state, including our own studies. 
Here we compile all available biological records for subterranean sites in California 
dating back to 1840. 
California’s karst is primarily comprised of small outcrops of marble or lime¬ 
stone with thousands of caves. Additionally, lava and ash flows, tens of thousands of 
mines, hundreds of sea caves on the mainland coast and islands, and extensive 
groundwater systems provide habitat for subterranean life. 
At least 4,600 caves of all types are known in California, of which 22% have been 
biologically sampled. We summarize 1,301 biological sites, and analyze data from 
1 
