ELLIOTT ET AL.: THE CAVE FAUNA OF CALIFORNIA 
23 
pede, Plumatyla humerosa, in caves of the Monument, and summarized the fauna in these and other 
volcanic caves in western North America (Peck 1973). In 1982, dipluran taxonomist Lynn Fergu¬ 
son (Longwood College) discovered a third species of Haplocampa (Ferguson 1992). 
Bat populations at Lava Beds have been monitored since at least 1988 by various researchers, 
including Dixie Pierson, Bruce Rogers, and David Cowan; that same year, the Cave Research 
Foundation established a long-term scientific study of the Monument’s caves. Under contract in 
1989, Rod Crawford (University of Washington), with colleagues Clyde Senger and David Cowan, 
bioinventoried some of the caves. In 1993, Christopher Richard, of the Oakland Museum of Cali¬ 
fornia, documented the diversity of ferns that exploited the microhabitats of the Monument’s caves. 
Under contract in 2005, Jean Krejca and Steve Taylor (Illinois Natural History Museum) invento¬ 
ried 29 caves in the Monument; these surveys revealed several new cave invertebrates that await 
taxonomic identification (Taylor and Krejca 2006; Shear et al. 2009). Diana Northup, National 
Cave and Karst Research Institute, and fellow microbiologists performed studies in the Monu¬ 
ment’s caves and other California caves to determine how microbes had adapted to these extreme 
environments (Northup and Boston 2006). 
Arachnologists at the California Academy of Sciences continue to make important contribu¬ 
tions to our understanding of cave faunas within California and beyond. Joel M. Ledford is docu¬ 
menting a huge diversity of species and adaptations in cave spiders in the genera Usofila and 
Calileptoneta, and with Charles Griswold, has described many new cave spiders in the family 
Leptonetidae (Ledford 2004; Ledford and Griswold 2010a-b). Griswold and colleagues also 
described a new family of cave spiders — Trogloraptoridae (Griswold et al. 2012); this new spider 
was first collected by G.O. Graening and fellow caver Ron Davis in a cave on the Califomia-Ore- 
gon border in 2011. 
Cave biologist G.O. Graening, Sacramento State University, began studying California caves 
in 2009. Teaming with cavers from California’s NSS (National Speleological Society) grottos and 
researchers from various museums and universities, Graening has performed bioinventories in 
more than 140 caves and springs in 18 counties. Discoveries to date include a new troglobitic 
silverfish, Speleonycta, from Crystal Stanislaus Cave, Calaveras Co.; a new cave amphipod, 
Stygobromus n. sp., from springs in Pinnacles National Park; a new genus and three new species 
of cave millipedes from the Marble Mountains, Opiona graeningi, etc.; and a new species of 
dipluran, Campodeidae (Shear 2011; Graening, Scherbanyik, and Arghandiwal 2012). Graening 
and colleagues published checklists of California’s isopods and amphipods and of North America’s 
ice crawlers that also summarized the distribution and diversity of these groups in subterranean 
habitats (Graening, Rogers, Holsinger, Barr, and Bottorff 2012; Graening and Rogers 2013; 
Schoville and Graening 2013). Graening created a statewide cave fauna occurrence database, 
Kryptos, and is collaborating with the other authors of this monograph and other scientists and 
conservationists. 
Recent taxonomic works include Gomez, Reddell, Will, and Moore (2016) on molecular 
systematics of the ground beetle Rhadine; Marek, Krejca, and Shear (2016) on the new millipede 
Illacme tobini; and Shear and Warfel (2016) on new species and revisions of Oskoron and Taracus 
harvestmen. 
Methods 
Besides the many literature records detailed above, collectively we have accumulated a large 
number of collections since the 1970s. Most collections were made by hand by the authors, their 
assistants and associates, cavers and entomologists working under permits or permissions from 
landowners. Most invertebrates were preserved in ethanol or other fluids appropriate for a particu- 
