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PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES 
Series 4, Volume 64, Supplement 1 
998 caves: (693 karst caves and features, 181 lava tubes, and 124 sea caves), 143 
groundwater sites, and 160 mines and tunnels. 
The richest regions for obligate subterranean species are the Sierra Nevada, Kla¬ 
math Mountains, and lava flows in the northern portion of the State. The high num¬ 
ber (72) of single-site endemic species is indicative of the insular distribution of karst, 
large differences in elevation, and the many river systems cutting across the state’s 
mountain ranges. 
In our database 1,366 taxa are recorded; 134 were determined to Family or high¬ 
er taxa only. There are 102 troglobites (terrestrial cave obligates), 12 stygobites 
(aquatic cave obligates), and 32 phreatobites (obligate groundwater forms). Of those 
146 obligate subterranean taxa, 11 are still undetermined beyond Order or Family, 
and represent an uncertain number of distinct species, although some may be more 
than single new species. Our species list includes 109 new (currently undescribed) 
species of all types, including 72 obligate subterranean species: 61 troglobites, 3 sty¬ 
gobites, and 8 phreatobites, significantly adding to the knowledge of California’s bio¬ 
diversity. 
California has a long, complex geologic history. Karst with highly endemic, troglobitic (cave- 
obligate or troglobiotic) species occurs in small outcrops of marble or limestone that originated 
from island terranes that were accreted onto the North American plate. Extensive lava and ash 
flows, mostly in northeastern California, also have rich cave fauna. More than 39,000 inactive 
mines occur all across the state, many of them containing bats and interesting invertebrates. One 
troglobitic species occurs in two littoral (sea) caves among the hundreds of sea caves on the coast 
and adjacent Channel and Farallon Islands, but many marine species are known from sea caves and 
there are some interesting terrestrial relicts. Extensive groundwater systems also provide habitat for 
subterranean life, some narrowly restricted to specific aquifers or river substrates. Boreal, montane, 
and tropical relicts are represented in the terrestrial cave fauna, but the true geographic origins of 
much of the fauna remain to be elucidated. 
Records of groundwater fauna in California date back to 1840, and cave fauna to 1863. Early 
20th century archaeologists and paleontologists conducted surveys of caves containing Pleistocene 
deposits, and they discovered many new extinct and extant species. Since 1975 the rate of discov¬ 
ery of new species has accelerated in most of the important cave areas of the state. 
In 1979, five of the nine current authors began compiling an exhaustive faunal list and bibli¬ 
ography. Many new species and records were added during intensive field work sponsored by the 
U.S. Army Corps of Engineers in 1977-1978. Most of this work was concentrated in the Stanislaus 
River canyon of the Northern Sierra Nevada because of a new dam for the New Melones Lake 
(reservoir) and the scheduled inundation of numerous caves. In 1979 a study funded by the U.S. 
Fish and Wildlife Service expanded the study of caves across northern California. 
By 1985 about 650 species and subspecies had been recorded from 281 caves, 23 sea caves, 
26 mines and 10 groundwater localities. In the 2000s projects were funded by the National Park 
Service at Sequoia and Kings Canyon National Parks, Yosemite National Park, and Lava Beds 
National Monument. The number of known subterranean species has doubled since 1985. The nine 
authors collectively have done extensive field work across the state, and this paper and our associ¬ 
ated Kryptos database (see Methods) document an effort to compile all biological records for sub¬ 
terranean sites in California. 
We summarize data from 1,296 biological sites, which are detailed below in maps, tables, and 
discussions. Figure 1 is a map depicting California cave regions with density of caves per county 
and 500 representative sites (localities) of all types. 
