Microtus californicus (Peale, 1848) were live trapped or retrapped 887 times, and fleas were collected over a 2.5-yr period in the San Francisco Watershed and Wildlife Refuge. Also, 179 M. californicus nests were collected monthly with observations to identify the environment of fleas. The ratio of the mean number of fleas per nest to the mean number collected on voles was 4.7:1 for Malaraeus telchinus (Rothschild, 1905), 13:1 for Hystrichopsylla occidentalis linsdalei Holland, 1957, 9:1 for Atyphloceras multidentatus multidentatus (C. Fox, 1909), and 76:1 for Catallagia wymani (C. Fox, 1909). The proportion of each flea species per nest to those per host was not closely associated seasonally or spatially. The average nest contained 37.5% moisture content in relation to its total weight and ranged between 3 and 92%. No relationship was observed between relative humidity of air within nests and flea number, but a significant relationship existed between the moisture of nesting materials and flea number. Malaraeus telchinus and A. m. multidentatus were collected in greatest numbers from nests having 30–39% moisture content by weight, and H. o. linsdalei and C. wymani were most numerous in nests that had 40–49% moisture content. Catallagia wymani had the greatest tolerance for high moisture and was severely affected by lack of moisture and nearly absent in drier nests. No ectoparasites were collected from nests that had <12% moisture content, and nests with >50% moisture content had few fleas. A static concept of nest fleas and host fleas as suggested by averages and often used in literature is questionable. In seasonal comparison of populations of fleas on hosts and in nests, M. telchinus and H. o. linsdalei reversed so that more of the flea population was on the host than in the nest for a short time during fall.