ELLIOTT ET AL.: THE CAVE FAUNA OF CALIFORNIA 
75 
number of troglobites per cave varies between 2 and 11 within that range. Calaveras (SNN) has 
4 caves in the top 20, Shasta (KM) 3, Tuolumne (SNN) 2, and the following with 1 each: Inyo 
(SNS), Mariposa (SNS), Napa (BD), Santa Cruz (BD). Looking at new species of obligates, SNN 
has 9 while SNS has 18. The Sierra Nevada has 15 caves in the top 20, with 6 in SNN and 9 in 
SNS. 
Ecology 
This section focuses on ecological studies, especially of cave communities. Some ecological 
topics were covered in previous sections. Figures 96-113 and previously mentioned figures illus¬ 
trate cave community ecology over several regions. 
Richard E. Graham published many small studies of California cave ecology, generally on 
trogloxenes, troglophiles, and the twilight zone. He provided some of the earliest published data on 
temperature, humidity, zonation, and communities in California caves, as well as biogeography. As 
mentioned in Literature Review and History, he discovered seven new cave species, including the 
troglobitic scorpion, Uroctonus grahami. Most of his work was published in Caves and Karst (for¬ 
merly Cave Notes ) by Cave Research Associates, a journal that is now extinct, but digital copies 
can be downloaded from the Karst Information Portal at http://www.karstportal.org/. 
As mentioned in Herpetofauna, Graham (1962d, 1963a) established that Pseudacris sierra 
(formerly Hyla regilla ), the Sierran treefrog (Fig. 92), is a trogloxene that dwells in cave entrance 
areas during hot, dry weather, and returns to nearby streams during the winter rainy season. This 
behavior differs from the pickerel frog, Rana palustris, in the Ozarks region, which overwinters in 
caves and takes temporary refuge in wet caves from hot and cold weather (Elliott 2003). These 
strategies are adaptive with respect to the climates of these two very different regions. 
Graham studied the use of caves by the twilight moth, Triphosa haesitata (Fig. 89), a troglox¬ 
ene also known from caves in Nevada. The species occupies entrances to deep caves in a varied 
geographic area from nearly sea level to 3,000 m elevation, in deserts and humid forests, and in all 
seasons, but it seems to be absent from most Sierra Nevada caves so far (Graham 1962b). Two 
other Triphosa species are known to inhabit caves in Great Britain, Europe, southwest Asia, and 
Japan (Graham 1968b). Suggett (1982) observed a woodrat preying on this moth in one Marble 
Mountains cave and reported different roosting behavior as compared to that reported by Graham 
for the low elevation Santa Cruz caves. 
Graham studied the fauna of Subway Cave, a lava tube in the Hat Creek Lava Flow of Shasta 
County. Despite the cave being less than 2,000 years old, it has an extensive fauna list, including 
at least two troglobites, a host of troglophiles, and two species of bats. Graham reports, “Since 
cooling, it has developed many cracks and fissures, and the ceiling has collapsed in two places. At 
present, tree roots, wood rat nests, and bat guano provide food for an arthropod fauna which 
fluctuates according to the cyclic availability of moisture. Most of the species of Subway Cave are 
normally found in other northern California caves. . . the millipede [Plumatyla humerosa ] and the 
isopod Ligidium [kofoidi ] (Fig. 96) are restricted to damp, dark places and are two species which 
have dispersed into the Hat Creek flow since it cooled. . . .” Subway Cave runs close to the sur¬ 
face, has a wide temperature range, and contains a variety of habitats (Graham 1962d). 
Graham (1963b) studied rove beetles, Staphylinidae (Fig. 79), and their burrowing and feed¬ 
ing habits in caves related to wood rat nests, including Subway Cave. He thought that many 
staphylinids are troglophiles, such as Lobrathium subseriatum and Quedius ( Microsaurus ) 
spelaeus. According to his studies these two species are the only ones whose larvae are found in 
all cave zones. Q. spelaeus may feed on small arthropods associated with feces, nesting materials, 
or other organic matter. Most other rove beetles are in the deep entrance zone or just within the dark 
zone. 
