ELLIOTT ET AL.: THE CAVE FAUNA OF CALIFORNIA 
15 
Literature Review And History 
The first scientific studies and collections of animals in California’s caves were sporadic and 
opportunistic, and began with the discovery of California’s rich diversity of bats in various cave 
and noncave habitats. Circa 1840, the famous naturalist John Audubon discovered the California 
Bat {Myotis californicus (Audubon and Bachman 1842)). In the next two decades, several new 
species of bats were discovered: California Leaf-nosed Bat ( Macrotus californicus ) and the West¬ 
ern Pipistrelle ( Parastrellus hesperus) in Fort Yuma (Imperial Co.); the Long-Eared Bat {Myotis 
evotis ) in Monterey; the Silver-Haired Bat {Lasionycteris noctivigans ) in Fort Reading (Shasta 
Co.); and the Pallid Bat {Antrozous pallidus) in the Mojave Desert (LeConte 1831; Baird 1858, 
1859; H. Allen 1864, 1893). Joel Allen, a mammalogist at the American Museum of Natural His¬ 
tory, described many of these western bat species and in the 1890s added two new species from 
California: the Fringed Bat {Myotis thysanodes ) from Dulzura (San Diego Co.) and Fort Tejon 
(Kern Co.); and the Spotted Bat {Euderma maculatum ) from Pirn (Ventura Co.) (J.A. Allen 1891). 
In this same decade, Clinton Merriam, known as the father of mammalogy and former head of the 
U.S. Bureau of Biological Survey, described two more new species of bats from California: the 
Western Bonneted Bat {Eumops perotis californicus ) and the Pocketed Free-Tailed Bat {Nyctin- 
omops femorosaccus ) (Merriam 1889, 1890). 
Under the tutelage of Joseph Grinnell (Director of the University of California Berkeley’s 
Museum of Vertebrate Zoology), Hilda Grinnell published her doctoral studies on the bats of Cal¬ 
ifornia in 1918. This comprehensive monograph described the distribution and habits of 63 bat 
species and subspecies, many of which are still valid today (H.W. Grinnell 1918). 
In 1879, Franz Steindachner described the blind goby fish {Typhlogobius californiensis ) from 
dark holes in rocky tidal pools in southern California. Because of its troglomorphic characters 
(degenerate eyes, lack of pigment, slow metabolism), this goby was studied by the ichthyologists 
Carl and Rosa Eigenmann alongside the typical cavefish that they were studying (R. Smith 1881; 
Eigenmann 1890). 
In 1866, national attention was brought to the marble caves of Calaveras County when a gold 
miner discovered a human skull thought to be very old (Anonymous 1881). Josiah Whitney, then 
the State Geologist of California as well as a geology professor at Harvard University, published 
that this “Calaveras Skull” was proof of humans’ presence in North America a million years ago 
(during the Pliocene Epoch he thought). It was later determined to be a hoax, with the skull, only 
a few hundred years old, transplanted from a nearby looted Indian burial. The date of the arrival of 
humans onto the North American continent is still debated to this day, with cave deposits being the 
primary evidence. 
Around this same time, California’s large diversity of cave invertebrates started to be discov¬ 
ered by naturalists and taxonomists who visited some of the well-known caves in the state. In 1868, 
L.G. Yates discovered a new genus and species of snail inside Cave City Cave (Calaveras Co.) — 
the Tight Coin Snail {Ammonitella yatesii [J.G. Cooper 1868]). James Cooper, then curator of gen¬ 
eral zoology at the California Academy of Sciences, described this new species, and in the 1870s, 
made various collections of his own, including the discovery of the Sierra Sideband Snail {Mon- 
adenia mormonum) in Pioneer Cave (El Dorado Co.) (Pilsbry 1939). In the 1890s, arachnologist 
George Marx discovered the California cave harvestman, Banksula californica, and the troglobitic 
spider, Usofila gracilis, in Alabaster Cave (El Dorado Co.); about this same time the troglophilic 
ground beetle, Anillaspis explanata, was also discovered in the now-destroyed Alabaster Cave 
(Packard 1888; Marx 1891; Banks 1900). In 1888, the entomologist and paleontologist Alpheus 
