ELLIOTT ET AL.: THE CAVE FAUNA OF CALIFORNIA 
31 
to varying degrees and four spider families (Leptonetidae, Linyphiidae, Nesticidae, and 
Dictynidae) are represented in all regions, but with different genera. One exception is Nesticus, 
which also has troglobites in the greater Appalachians. Pseudoscorpion troglobites are much more 
diverse in greater Appalachia, which has about 100 species in 9 genera, or about 5 times that in 
California. The families Chthoniidae, Neobisiidae and Chemetidae occur in all regions, with some 
shared genera. In addition to California, troglobitic Aphrastochthonius and Apochthonius also 
occur in Texas and Appalachia, respectively. In Opiliones, troglobitic Taracidae are represented by 
Taracus (Fig. 26) in California and Hesperonemastoma in the Interior Low Plateaus. The family 
Phalangodidae is well represented in the three regions with endemic genera, except for Texella, 
which occurs in both California and Texas. 
Myriapods 
Myriapods (“many feet”) are an ancient Subphylum of arthropods grouped in four Classes. 
The first terrestrial arthropod was a millipede from the middle Silurian Period, about 428 Ma, its 
fossil discovered in Scotland (Wilson and Anderson 2004). But all myriapods may be descended 
from a basic body plan established in the late Cambrian period about 500 Ma. The evolutionary 
relationships of these classes are still being researched. 
Myriapods are common in moist forests and caves. In caves millipedes are usually found near 
moist organic matter. Most are detritivores, but centipedes are generally nocturnal predators. These 
characteristics have led many species to adapt to caves (Table 4). 
Myriapods have a single pair of antennae and, in most cases, simple eyes. Some groups, such 
as polydesmoid millipedes and geophilomorph centipedes, lack eyes even if not troglobitic. The 
mouthparts lie on the underside of the head, with an “epistome” and labrum forming the upper lip, 
and a pair of maxillae forming the lower lip. A pair of mandibles lie inside the mouth. 
Class Chilopoda, centipedes, predators, one pair of legs per body segment, first pair of legs 
modified as venomous fangs under the head. The Orders Geophilomorpha and Lithobiomorpha are 
illustrated in Figures 56-58. One Californian troglobite, the geophilomorph Stenophilus californi- 
cus, is known from two caves in two regions: Brown Deer Cave, Calaveras County; and Potter 
Creek Cave (type locality), Shasta County. It is described from one male specimen, about 90 mm 
long and pale yellow. The legs were described as “short.” This may be the “large myriapod” 
referred to by Sinclair (1904). An undescribed lithobiomorph is known from Porcupine Cave 
Tuolumne County. This species is depigmented, eyeless, and has elongated appendages. Cen¬ 
tipedes prey on small cave invertebrates, as shown in Figure 57. 
Class Symphyla, garden centipedes, one pair of legs per segment, white and eyeless endo- 
geans (soil dwellers). Undetermined species of tiny endogeans have been collected in 29 caves and 
9 counties across the state, except in the deserts. Symphylella oviceps was found in gravelly Trans¬ 
plant Mine tailings, Tuolumne County. At least four species of Scutigerellidae have been found in 
caves of the Mother Lode Region (Sierra Nevada North). Scutigerella causeyae is known from 
three caves, and it was introduced from McLean’s Cave to the Transplant Mine. It is also found in 
the mine tailings and on the surface in the vicinity. It ranges widely throughout the United States. 
Figure 59 is a symphylan from a cave in Sierra Nevada South, in Sequoia-Kings Canyon National 
Parks. 
Class Pauropoda, dwarf millipedes, tiny endogeans rarely found in caves, possibly the most 
primitive myriapods, with semi-fused body rings and a short body. An undetermined species of 
endogean was found in Overhang Cave, Tulare County, the first pauropod record from a Califor¬ 
nia cave. 
