38 
PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES 
Series 4, Volume 64, Supplement 1 
compared to S. laticeps, and it was collected only in caves whereas S. laticeps was collected at 
night outside the caves. Furthermore, S. nunenmacheri has perhaps one of the largest eggs of any 
tenebrionid in the world. 
3. Troglophiles and troglobites include Anobiidae (Ptininae, spider beetles), Carabidae, Leio- 
didae, Staphylinidae, and Tenebrionidae. Common species include the leiodid, Ptomaphagus 
nevadicus (Fig. 77) and rove beetles (Staphylinidae, Fig. 78), which are found near dung. Table 7 
lists the known troglobites. From just a few concentrated cave studies in California, remarkable 
troglophilic and troglobitic cave beetles are already known or are in the process of being described. 
Notable examples include the Anobiidae (Ptininae), Niptus arcanus, a troglobite endemic to 
Mitchell Caverns, San Bernardino County, Mojave Desert (Fig. 79). Despite having connected two 
adjacent caves in 1969 by a tunnel to facilitate tours, this species is still restricted to only one side 
of Mitchell Caverns: El Pakiva. Among the Carabidae are three noteworthy species, two of which 
are from the Anillina subtribe. Anillines are often completely blind beetles found in deep soil 
deposits often associated with groundwater. The completely blind, undescribed anilline troglobite 
from Clay Cave, Napa County, Bay Area/Delta, is a rather large member of this group and repre¬ 
sents a new genus and species. The completely blind undescribed anilline troglobite (Fig. 80) from 
the Sierra Nevada North, in Moaning Cave, Calaveras County, and Crystal Palace, Tuolumne 
County, also represents a new genus and species. This giant new species is the largest known 
species in this tribe in the world! (D. Kavanaugh, in lit.). A third carabid, Tribe Pterostichini, from 
Paul Gibson Cave (limestone), Trinity County, Pterostichus (Leptoferonia) enyo Will, 2007, has 
very small eyes, and represents an endogean (soil interstitial species) that occasionally occurs 
under deep rocks/logs or in caves. 
Among the Tenebrionidae, the genus Eleodes with about 200 species ranging from the West¬ 
ern U.S.A. to northern Guatemala, and the subgenus Caverneleodes contain a number of cave 
endemics from Mexico to California. Many are cave-adapted with long appendages and reduced 
eyes. The tenebrionid species from Clay Cave represents a new tribe of Tenebrionidae, along with 
another found under a rock at a spring mouth in Marin County. Both species are troglobites that 
prefer wet soil. The California species E. ( Caverneleodes ) microps (Fig. 81), a troglobite, has the 
Table 7. Ten troglobitic beetles of California; only four are described species 
Family/ Subfamily 
Species 
Cave 
Ecol. Class. 
Anobiidae, Ptininae 
Niptus arcanus 
San Bernardino Co.: Mitchell Cav¬ 
erns (El Pakiva Cave) 
troglobite 
Carabidae, Trechinae 
Anillaspis explanata 
El Dorado Co.: Alabaster Cave 
troglobite 
Carabidae, Trechinae 
Anillina n. gen., n. sp. 1 
Tuolumne Co.: McLean's Cave, 
Transplant Mine 
troglobite 
Carabidae, Trechinae 
Anillina n. gen., n. sp. 2 
Tulare Co.: Lost Soldier's Cave. 
troglobite 
Carabidae, Trechinae 
Anillina n. gen., n. sp. 3 
Napa Co.: Clay Cave 
troglobite 
Carabidae, Trechinae 
Anillina n. gen., n. sp. 4 
Calaveras Co.: Moaning Cave. 
Tuolumne Co.: Crystal Palace 
troglobite 
Curculionidae, 
Raymondionyminae 
Gilbertiola sp. 
Tulare Co.: Clough Cave 
troglobite 
Leiodidae, Leiodinae 
Ptomaphagus inyoensis 
Inyo Co.: Poleta Cave 
troglobite 
Tenebrionidae, 
Lagriinae 
Eschatoporis n. sp. 
Napa Co.: Clay Cave 
troglobite 
Tenebrionidae, Tene- 
brioninae 
Eleodes ( Caverneleodes ) microps 
Inyo Co.: Microps Cave, Defense 
Cave, Poleta Cave 
troglobite 
