Seth P. D. Riley

A southern California freeway is a physical and social barrier to gene flow in carnivores.

Roads present formidable barriers to dispersal. We examine movements of two highly mobile carnivores across the Ventura Freeway near Los Angeles, one of the busiest highways in the United States. The two species, bobcats and coyotes, can disappear from habitats isolated and fragmented by roads, and their ability to disperse across the Ventura Freeway tests the limits of vertebrates to overcome anthropogenic obstacles. We combine radio‐telemetry data and genetically based assignments to identify individuals that have crossed the freeway. Although the freeway is a significant barrier to dispersal, we find that carnivores can cross the freeway and that 5–32% of sampled carnivores crossed over a 7‐year period. However, despite moderate levels of migration, populations on either side of the freeway are genetically differentiated, and coalescent modelling shows their genetic isolation is consistent with a migration fraction less than 0.5% per generation. These results imply that individuals that cross the freeway rarely reproduce. Highways and development impose artificial home range boundaries on territorial and reproductive individuals and hence decrease genetically effective migration. Further, territory pile‐up at freeway boundaries may decrease reproductive opportunities for dispersing individuals that do manage to cross. Consequently, freeways are filters favouring dispersing individuals that add to the migration rate but little to gene flow. Our results demonstrate that freeways can restrict gene flow even in wide‐ranging species and suggest that for territorial animals, migration levels across anthropogenic barriers need to be an order of magnitude larger than commonly assumed to counteract genetic differentiation.

FAST‐TRACK: A southern California freeway is a physical and social barrier to gene flow in carnivores

Roads present formidable barriers to dispersal. We examine movements of two highly mobile carnivores across the Ventura Freeway near Los Angeles, one of the busiest highways in the United States. The two species, bobcats and coyotes, can disappear from habitats isolated and fragmented by roads, and their ability to disperse across the Ventura Freeway tests the limits of vertebrates to overcome anthropogenic obstacles. We combine radio‐telemetry data and genetically based assignments to identify individuals that have crossed the freeway. Although the freeway is a significant barrier to dispersal, we find that carnivores can cross the freeway and that 5–32% of sampled carnivores crossed over a 7‐year period. However, despite moderate levels of migration, populations on either side of the freeway are genetically differentiated, and coalescent modelling shows their genetic isolation is consistent with a migration fraction less than 0.5% per generation. These results imply that individuals that cross the freeway rarely reproduce. Highways and development impose artificial home range boundaries on territorial and reproductive individuals and hence decrease genetically effective migration. Further, territory pile‐up at freeway boundaries may decrease reproductive opportunities for dispersing individuals that do manage to cross. Consequently, freeways are filters favouring dispersing individuals that add to the migration rate but little to gene flow. Our results demonstrate that freeways can restrict gene flow even in wide‐ranging species and suggest that for territorial animals, migration levels across anthropogenic barriers need to be an order of magnitude larger than commonly assumed to counteract genetic differentiation.

Use of highway undercrossings by wildlife in southern California

Abstract Roads, especially large highways, can have significant impacts on wildlife movement and survival. This is especially true for wide-ranging species, such as mammalian carnivores. Some of these impacts may be mitigated if wildlife can find and utilize passageways under highways. To determine if underpasses and drainage culverts beneath highways are used by wildlife as movement corridors, we monitored 15 such passages near Los Angeles, California using remotely triggered cameras and gypsum track stations. We found that passages were used by a variety of species, including carnivores, mule deer, small mammals, and reptiles. Many types of undercrossings were utilized, indicating that passages beneath highways, even when not originally designed for wildlife, can provide important safe avenues for animals to cross roads. For mammals of conservation concern, including native carnivores and deer, passage dimensions, surrounding habitat, and the extent of human activity were assessed to determine if these factors influenced passage use by these species. Our results show that while many native mammals used passages beneath highways, the presence of suitable habitat on either side of the passage was a particularly important factor predicting use. For deer and coyotes, passage dimensions were also important and should be considered with the presence of suitable habitat when wildlife passages are planned or evaluated. To increase the likelihood of utilization and to help prevent animals from crossing road surfaces, we suggest that simple improvements such as habitat restoration near crossing points and animal-proof fencing that serves to funnel wildlife to passages, can facilitate animal movement between fragmented habitats that are bisected by roads.

A Rapid, Strong, and Convergent Genetic Response to Urban Habitat Fragmentation in Four Divergent and Widespread Vertebrates

Background Urbanization is a major cause of habitat fragmentation worldwide. Ecological and conservation theory predicts many potential impacts of habitat fragmentation on natural populations, including genetic impacts. Habitat fragmentation by urbanization causes populations of animals and plants to be isolated in patches of suitable habitat that are surrounded by non-native vegetation or severely altered vegetation, asphalt, concrete, and human structures. This can lead to genetic divergence between patches and in turn to decreased genetic diversity within patches through genetic drift and inbreeding. Methodology/Principal Findings We examined population genetic patterns using microsatellites in four common vertebrate species, three lizards and one bird, in highly fragmented urban southern California. Despite significant phylogenetic, ecological, and mobility differences between these species, all four showed similar and significant reductions in gene flow over relatively short geographic and temporal scales. For all four species, the greatest genetic divergence was found where development was oldest and most intensive. All four animals also showed significant reduction in gene flow associated with intervening roads and freeways, the degree of patch isolation, and the time since isolation. Conclusions/Significance Despite wide acceptance of the idea in principle, evidence of significant population genetic changes associated with fragmentation at small spatial and temporal scales has been rare, even in smaller terrestrial vertebrates, and especially for birds. Given the striking pattern of similar and rapid effects across four common and widespread species, including a volant bird, intense urbanization may represent the most severe form of fragmentation, with minimal effective movement through the urban matrix.

EXPOSURE TO FELINE AND CANINE PATHOGENS IN BOBCATS AND GRAY FOXES IN URBAN AND RURAL ZONES OF A NATIONAL PARK IN CALIFORNIA

Exposure of bobcats (Lynx rufus) and gray foxes (Urocyon cinereoargenteus) to a range of common canine and feline pathogens was assessed in urban and rural zones of Golden Gate National Recreation Area, a National Park in the San Francisco Bay Area, (California, USA) from 1992 to 1995. Testing included serology for canine distemper virus, canine parvovirus (CPV), canine adenovirus, Leptospira interrogans, feline calicivirus (FCV), feline panleukopenia virus, feline herpesvirus, feline enteric coronavirus (FECV), feline immunodeficiency virus, feline leukemia virus, Toxoplasma gondii, and Bartonella henselae. Testing was also performed for Dirofilaria immitis. Significantly more gray foxes were seropositive for CPV in the urban zone than in the rural zone. In addition, radio-tracking of gray foxes in the rural zone indicated that all three of the rural CPV-seropositive foxes had traveled into adjoining small towns, whereas only one of the 11 seronegative animals had done so. Significantly more bobcats were seropositive for FCV in the rural zone than in the urban zone. Individual bobcats with positive FCV antibody titers had patterns of movement that intercepted park inholdings where domestic cats lived. Bobcat samples were seronegative for all five of the other viral feline pathogens, with the exception of a FECV-seropositive bobcat. High seroprevalence was detected for B. henselae and T. gondii in both zones. Variation in the seroprevalence for different pathogens might be related to differences in the exposure of bobcats and foxes to domestic animals: in the urban zone, gray foxes were located in residential areas outside the park, whereas bobcats were not. Although for most of the pathogens examined there was no relationship between urbanization and exposure, our results for CPV in foxes and FCV in bobcats indicated that proximity to urban areas or contact with humans can increase the risk of disease exposure for wild carnivore populations. Combining behavioral information from radio-tracking with data on pathogen exposure or disease incidence can provide valuable insights into the ecology of wildlife disease that might be missed with broadscale, population-level comparisons alone.

HYBRIDIZATION BETWEEN A RARE, NATIVE TIGER SALAMANDER (AMBYSTOMA CALIFORNIENSE) AND ITS INTRODUCED CONGENER

Exotic species threaten native biodiversity through predation, competition, and habitat alteration, but also by hybridizing with native species. A lack of reproductive isolation between exotic and native species can lead to genetic swamping, loss of native genetic diversity, and, in rare or endangered species, extirpation or extinction. We examined hybridization between a declining native salamander, the California tiger salamander, Am- bystoma californiense, and an introduced congener, A. tigrinum. Ambystoma californiense is restricted to central California where A. tigrinum has been deliberately introduced as fish bait. In the Salinas Valley, we sampled salamanders from four artificial ponds and two natural vernal pools. Based on mitochondrial DNA and two nuclear loci, we found that hybrids were present in all six ponds, and that these hybrids were viable and fertile. No potentially pure A. californiense were present in three of the six ponds, and only one pond had more than 8% possibly pure native animals. Despite a relatively ancient split and wide genetic divergence between these taxa, they are interbreeding and threatening the genetic purity of the native species. Our data also suggest that the extent of the genetic mixing depends on the breeding habitat. There is little evidence of barriers to gene exchange in the four artificial breeding ponds. However in the two vernal pools, we found significantly fewer larvae with hybrid genotypes and significantly more with pure parental genotypes than expected. Linkage disequilibria revealed positive associations between native alleles and genotypes, and neg- ative associations between native and introduced alleles and genotypes in these two ponds. Despite rampant hybridization, these data provide evidence of some constraints on hy- bridization in the native breeding habitats. Our results suggest that habitat characteristics of native species should be exploited in management strategies to limit hybridization with exotics.

Anticoagulant Exposure and Notoedric Mange in Bobcats and Mountain Lions in Urban Southern California

Abstract Humans introduce many toxicants into the environment, the long-term and indirect effects of which are generally unknown. We investigated exposure to anticoagulant rodenticides and evaluated the association between notoedric mange, an ectoparasitic disease, and anticoagulant exposure in bobcats (Lynx rufus) and mountain lions (Puma concolor) in a fragmented urban landscape in southern California, USA. Beginning in 2002, an epizootic of notoedric mange, a disease previously reported only as isolated cases in wild felids, in 2 years reduced the annual survival rate of bobcats from 0.77 (5-yr average) to 0.28. Anticoagulants were present in 35 of 39 (90%) bobcats we tested, multiple compounds were present in 27 of these 35 (77%), and total toxicant load was positively associated with the use of developed areas by radiocollared animals. Mange-associated mortality in bobcats showed a strong association with anticoagulant exposure, as 19 of 19 (100%) bobcats that died with severe mange were also exposed to the toxicants, and for bobcats with anticoagulant residues >0.05 ppm, the association with mange was highly significant (χ2 = 10.36, P = 0.001). We speculate that concomitant elevated levels of rodenticide exposure may have increased the susceptibility of bobcats to advanced mange disease. Bobcats were locally extirpated from some isolated habitat patches and have been slow to recover. In 2004, 2 adult mountain lions died directly from anticoagulant toxicity, and both animals also had infestations of notoedric mange, although not as advanced as in the emaciated bobcats that died with severe disease. Two other mountain lions that died in intraspecific fights also exhibited exposure to 2–4 different anticoagulants. These results show that the effects of secondary poisoning on predators can be widespread, reach even the highest-level carnivores, and have both direct and possibly indirect effects on mortality. Further research is needed to investigate the lethal and sub-lethal effects of anticoagulants and other toxicants on wildlife in terrestrial environments.

Three Pathogens in Sympatric Populations of Pumas, Bobcats, and Domestic Cats: Implications for Infectious Disease Transmission

Anthropogenic landscape change can lead to increased opportunities for pathogen transmission between domestic and non-domestic animals. Pumas, bobcats, and domestic cats are sympatric in many areas of North America and share many of the same pathogens, some of which are zoonotic. We analyzed bobcat, puma, and feral domestic cat samples collected from targeted geographic areas. We examined exposure to three pathogens that are taxonomically diverse (bacterial, protozoal, viral), that incorporate multiple transmission strategies (vector-borne, environmental exposure/ingestion, and direct contact), and that vary in species-specificity. Bartonella spp., Feline Immunodeficiency Virus (FIV), and Toxoplasma gondii IgG were detected in all three species with mean respective prevalence as follows: puma 16%, 41% and 75%; bobcat 31%, 22% and 43%; domestic cat 45%, 10% and 1%. Bartonella spp. were highly prevalent among domestic cats in Southern California compared to other cohort groups. Feline Immunodeficiency Virus exposure was primarily associated with species and age, and was not influenced by geographic location. Pumas were more likely to be infected with FIV than bobcats, with domestic cats having the lowest infection rate. Toxoplasma gondii seroprevalence was high in both pumas and bobcats across all sites; in contrast, few domestic cats were seropositive, despite the fact that feral, free ranging domestic cats were targeted in this study. Interestingly, a directly transmitted species-specific disease (FIV) was not associated with geographic location, while exposure to indirectly transmitted diseases – vector-borne for Bartonella spp. and ingestion of oocysts via infected prey or environmental exposure for T. gondii – varied significantly by site. Pathogens transmitted by direct contact may be more dependent upon individual behaviors and intra-specific encounters. Future studies will integrate host density, as well as landscape features, to better understand the mechanisms driving disease exposure and to predict zones of cross-species pathogen transmission among wild and domestic felids.

Spatial Ecology of Bobcats and Gray Foxes in Urban and Rural Zones of a National Park

Abstract Urbanization threatens the persistence of many wildlife populations, particularly those of wide-ranging and low-density species such as mammalian carnivores. Effective conservation of carnivore populations requires an understanding of the impacts of adjacent urbanization on carnivores in reserves. I compared the spatial ecology of bobcats (Lynx rufus) and gray foxes (Urocyon cinereoargenteus) between urban and rural zones of a national park in northern California, USA. In the urban zone, gray foxes used the entire landscape from interior natural areas across the park edges and into the neighboring developed areas, although fox core areas were always within the park. Bobcats never entered development, and radiocollared adult female bobcats maintained home ranges in the interior of the park, far from the urban edge. Bobcats appeared to avoid crossing paved roads, while foxes crossed roads regularly. For adult female bobcats, home ranges were smaller in the urban zone, and core areas were both smaller and overlapped more. Home range size and overlap did not differ between zones for gray foxes. Bobcats seem to be more affected by the proximity of urbanization than foxes, perhaps because of differences in diet and social structure. The more flexible use of the landscape by foxes may give them access to increased resources and habitat, but also may expose them to more human-associated risks. If female bobcats are more sensitive to urbanization, this sensitivity could affect the long-term viability of bobcat populations in urban areas. Knowledge of how bobcats and gray foxes use the landscape in urban areas will allow more effective conservation and improved coexistence with these widespread carnivores by helping to predict where and why conservation or management issues may occur.

ESTIMATING BOBCAT POPULATION SIZES AND DENSITIES IN A FRAGMENTED URBAN LANDSCAPE USING NONINVASIVE CAPTURE-RECAPTURE SAMPLING

Abstract Bobcats (Lynx rufus) are valuable indicators of connectivity in the highly fragmented landscape of coastal southern California, yet their population sizes and densities are largely unknown. Using noninvasive scat sampling in a capture–recapture framework, we estimated population sizes for 2 similar areas of natural habitat with differing levels of isolation by human development in Santa Monica Mountains National Recreation Area, California. We used scat transects with geographic information system land-use layers and home-range sizes of bobcats to estimate effective sampling area and population densities. Estimates of population size in the study area connected to a much larger habitat area (26–31 individuals) were similar to estimates for the area that was completely surrounded by development (25–28 individuals). Bobcat densities for the 2 study areas also were similar (ranging from 0.25 to 0.42 bobcat/km2) and likely represent recent population declines because of notoedric mange likely interacting with toxicants. These methods proved effective despite particularly low densities of bobcats and may be especially useful when study areas are geographically isolated, reducing the uncertainty in size of the sampling area.