Habitat selection when killing primary versus alternative prey species supports prey specialization in an apex predator

© 2019 The Zoological Society of London Many predators specialize on one or several prey species that they select from the range of potential prey. Predator specialization on primary versus alternative prey is driven in part by encounter rates with prey and a predator’s habitat selection. Although habitat selection changes with behavioural state, this has not been well-recognized in the resource selection function (RSF) literature to date, often because auxiliary data on the predator’s behavioural states (e.g. hunting) are absent. We monitored habitat selection of pumas Puma concolor in a multi-prey system in northern California, where pumas specialized on black-tailed deer Odocoileus hemionus columbianus. We employed multiple RSF analyses on different datasets to test the following three hypotheses: (1) Pumas utilize habitats in proportion to their availability; (2) Pumas select specific habitat features when killing black-tailed deer, their primary prey; (3) Pumas do not select distinct habitats from those identified under hypothesis 1 when killing alternative prey. We found that pumas in our study selected for specific habitats and habitat features in general, but that their selection was more pronounced when killing black-tailed deer. In summer, kill sites of deer were associated with rugged terrain, but gentle slopes and northerly aspects. In winter, pumas killed deer at low elevations, on gentle slopes and on northerly and westerly aspects. Overall, evidence suggested that pumas tracked their primary prey across seasonal migrations, which were short in distance but resulted in pronounced changes in elevation. When killing alternative prey, pumas showed little evidence of habitat selection, suggesting they may kill alternative prey opportunistically. Our results hold implications for how data should be partitioned when modelling baseline habitat selection of predators, hunting habitat selection and predation risk for prey species, as well as for how we model ecological processes such as apparent competition.

Habitat selection when killing primary versus alternative prey species supports prey specialization in an apex predator

© 2019 The Zoological Society of London Many predators specialize on one or several prey species that they select from the range of potential prey. Predator specialization on primary versus alternative prey is driven in part by encounter rates with prey and a predator’s habitat selection. Although habitat selection changes with behavioural state, this has not been well-recognized in the resource selection function (RSF) literature to date, often because auxiliary data on the predator’s behavioural states (e.g. hunting) are absent. We monitored habitat selection of pumas Puma concolor in a multi-prey system in northern California, where pumas specialized on black-tailed deer Odocoileus hemionus columbianus. We employed multiple RSF analyses on different datasets to test the following three hypotheses: (1) Pumas utilize habitats in proportion to their availability; (2) Pumas select specific habitat features when killing black-tailed deer, their primary prey; (3) Pumas do not select distinct habitats from those identified under hypothesis 1 when killing alternative prey. We found that pumas in our study selected for specific habitats and habitat features in general, but that their selection was more pronounced when killing black-tailed deer. In summer, kill sites of deer were associated with rugged terrain, but gentle slopes and northerly aspects. In winter, pumas killed deer at low elevations, on gentle slopes and on northerly and westerly aspects. Overall, evidence suggested that pumas tracked their primary prey across seasonal migrations, which were short in distance but resulted in pronounced changes in elevation. When killing alternative prey, pumas showed little evidence of habitat selection, suggesting they may kill alternative prey opportunistically. Our results hold implications for how data should be partitioned when modelling baseline habitat selection of predators, hunting habitat selection and predation risk for prey species, as well as for how we model ecological processes such as apparent competition.

Adenoviral hemorrhagic disease in California mule deer, 1990–2014

We reviewed case records from the California Animal Health and Food Safety (CAHFS) laboratory and the California Department of Fish and Wildlife (CDFW) spanning 25 years (1990–2014) for all deer accessions submitted to CAHFS for pathology and/or histopathology, with and without a diagnosis of adenoviral hemorrhagic disease (AHD), in order to determine the prevalence of AHD in California. We also examined spatial and temporal distribution, age, and mule deer subspecies in deer that died from AHD. Of 483 deer submitted to CAHFS for diagnostic testing in 1990–2014, 17.2% were diagnosed with confirmed AHD, and 26.5% were confirmed plus suspected cases of AHD. Columbian black-tailed deer ( Odocoileus hemionus columbianus), particularly fawns and juveniles, were most frequently affected. Deer adenovirus ( Odocoileus adenovirus 1; OdAdV-1) was detected by immunohistochemistry in archived CDFW formalin-fixed, paraffin-embedded tissues from deer that died in mortality events in 1981, 1983, and 1986–1987. OdAdV-1 is a common cause of hemorrhagic disease mortality events in California deer, and mortality as a result of AHD is documented as early as 1981.

Antler and Body Size in Black-Tailed Deer: An Analysis of Cohort Effects

For long-lived species, environmental factors experienced early in life can have lasting effects persisting into adulthood. Large herbivores can be susceptible to cohort-wide declines in fitness as a result of decreases in forage availability, because of extrinsic factors, including extreme climate or high population densities. To examine effects of cohort-specific extrinsic factors on size of adults, we performed a retrospective analysis on harvest data of 450 male black-tailed deer (Odocoileus hemionus columbianus) over 19 years in central California, USA. We determined that population density of females had a more dominant effect than did precipitation on body size of males. Harvest of female deer resulted in increases in the overall size of males, even though a 6-year drought occurred during that treatment period. Body size was most influenced by female population density early in life, while antler size was highly affected by both weather early in life and the year directly before harvest. This study provides insights that improve our understanding of the role of cohort effects in body and antler size by cervids; and, in particular, that reduction in female population density can have a profound effect on the body and antler size of male deer.

Seasonal variation in the feeding ecology of pumas (Puma concolor) in northern California

Direct effects of predators depend upon factors that can vary across seasons, including variations in the abundance and vulnerability of migrating prey. Past studies show conflicting results of whether puma (Puma concolor (L., 1771)) feeding ecology varies among seasons. We employed GPS collars to study puma feeding ecology in a single-prey system with migratory black-tailed deer (Odocoileus hemionus columbianus (Richardson, 1829)). We hypothesized that puma feeding ecology would vary based on changes in prey abundance and spatial distribution, as well as competition with scavengers and decomposers. Our results supported these hypotheses. Kill rates in number of ungulates/week were significantly higher in summer and autumn than in winter, likely owing to the increased availability and density of black-tailed deer fawns. The handling times of black-tailed deer ≥1 year old were significantly higher in winter than in spring, summer, or autumn. We speculated that reduced handling time in summer may have been influenced by black bear (Ursus americanus Pallas, 1780) kleptoparasitism and the decomposition of kills. Pumas killed black-tailed deer at higher elevations in summer than in winter, spring, or autumn, and the elevations correlated significantly with seasonal elevations used by black-tailed deer, suggesting that pumas exhibited seasonal foraging behaviours and tracked prey availability in a system with migrating prey.

Conditional daily and seasonal movement strategies of male Columbia black-tailed deer (Odocoileus hemionus columbianus)

Animals move to maximize fitness via resource acquisition, predator avoidance, thermoregulation, or mate access. Variations in movement strategies among and within populations often reflect habitat- or demographic-specific variations in fitness trade-offs. To examine these conditional movement strategies, we modeled seasonal and diel movement patterns of radio-collared adult male Columbia black-tailed deer (Odocoileus hemionus columbianus (Richardson, 1829)) on a temperate, predator-free island. Linear 10 h displacement and home-range areas reached annual maxima during autumn and minima during late winter, corresponding with known dates for breeding season and lowest quality forage, respectively. For all males in all years, initiation of increased movements began during spring and again, abruptly, in late September, immediately prior to peak breeding season. Larger antlered males continued increased movements longer into December, suggesting increased breeding effort relative to smaller antlered males. Time of day predicted movements during all seasons; however, we observed no strong evidence of the crepuscular or nocturnal movement bias typically noted in deer, likely relating to the lack of predators in our study area. In this way, male black-tailed deer adopted conditional, seasonally specific movement strategies to balance fitness trade-offs in resource acquisition, thermoregulation, and mate access.

Encounter Competition between a Cougar, Puma concolor, and a Western Spotted Skunk, Spilogale gracilis

Encounter competition occurs frequently over food resources and may include kleptoparasitism, where scavengers usurp prey killed by carnivores. Scavenging may have important adverse effects on carnivores and may result in higher than expected kill rates by predators. Using camera traps placed on a Black-Tailed Deer (Odocoileus hemionus columbianus) carcass killed by a Cougar (Puma concolor) in California, we observed a series of encounters in which a Western Spotted Skunk (Spilogale gracilis) temporally usurped the carcass from the Cougar. The Western Spotted Skunk also successfully defended the carcass when the Cougar returned and attempted to feed. The Spotted Skunk was about 1% of the mass of the Cougar. Our observation is the largest reported size differential of a mammalian species engaging in successful encounter competition.