Summary Species occurrence is influenced by environmental conditions and the presence of other species. Current approaches for multispecies occupancy modelling are practically limited to two interacting species and often require the assumption of asymmetric interactions. We propose a multispecies occupancy model that can accommodate two or more interacting species. We generalize the single‐species occupancy model to two or more interacting species by assuming the latent occupancy state is a multivariate Bernoulli random variable. We propose modelling the probability of each potential latent occupancy state with both a multinomial logit and a multinomial probit model and present details of a Gibbs sampler for the latter. As an example, we model co‐occurrence probabilities of bobcat (Lynx rufus), coyote (Canis latrans), grey fox (Urocyon cinereoargenteus) and red fox (Vulpes vulpes) as a function of human disturbance variables throughout 6 Mid‐Atlantic states in the eastern United States. We found evidence for pairwise interactions among most species, and the probability of some pairs of species occupying the same site varied along environmental gradients; for example, occupancy probabilities of coyote and grey fox were independent at sites with little human disturbance, but these two species were more likely to occur together at sites with high human disturbance. Ecological communities are composed of multiple interacting species. Our proposed method improves our ability to draw inference from such communities by permitting modelling of detection/non‐detection data from an arbitrary number of species, without assuming asymmetric interactions. Additionally, our proposed method permits modelling the probability two or more species occur together as a function of environmental variables. These advancements represent an important improvement in our ability to draw community‐level inference from multiple interacting species that are subject to imperfect detection.
Mule deer and black‐tailed deer Odocoileus hemionus have exhibited marked population fluctuations throughout their range over the past century. The relative contributions of predation, forage availability and weather to observed population changes remain unclear and controversial. We reviewed 48 studies on Odocoileus hemionus survival and predation from the past 30 years and quantified age‐specific vital rates, population growth rates (λ) and causes of mortality. We also evaluated the effect of environmental variables on variation in vital rates and the contribution of age‐specific survival to population growth. Age‐specific survival (ϕ) was the most frequently studied population parameter. Odocoileus hemionus have lower and more variable fawn survival than other ungulate species (ϕsummer = 0.44, CV = 0.42; ϕannual = 0.29, CV = 0.67). Adult female survival conversely appeared to be high and stable throughout the geographical range of the species (ϕannual = 0.84, CV = 0.06). Observed low fawn survival appears to be compensated for by high fecundity rates. Predation was the primary proximate cause of mortality for all age classes, and was an important source of summer fawn mortality and of mortality in multi‐prey, multi‐predator systems. However, predator removal studies suggest that predation is compensatory, particularly at high deer densities, and that nutrition and weather shape population dynamics. We propose three models to explain local population dynamics of Odocoileus hemionus: (i) populations are limited by forage availability and weather; (ii) adult females are limited by forage availability, fawns are limited by forage availability and predation, and population growth is constrained by fecundity and fawn predation; and (iii) large changes in the abundance of predators or alternative prey change predation risk and destabilize population dynamics. Future research should be focused on: the effects of age‐specific survival on population growth; possible interactions between predation, forage availability and weather; and the importance of multiple predator and prey species in shaping the population dynamics of Odocoileus hemionus.
Camera trapping has become an increasingly widespread tool for wildlife ecologists, with large numbers of studies relying on photo capture rates or presence/absence information. It is increasingly clear that camera placement can directly impact this kind of data, yet these biases are poorly understood. We used a paired camera design to investigate the effect of small-scale habitat features on species richness estimates, and capture rate and detection probability of several mammal species in the Shenandoah Valley of Virginia, USA. Cameras were deployed at either log features or on game trails with a paired camera at a nearby random location. Overall capture rates were significantly higher at trail and log cameras compared to their paired random cameras, and some species showed capture rates as much as 9.7 times greater at feature-based cameras. We recorded more species at both log (17) and trail features (15) than at their paired control cameras (13 and 12 species, respectively), yet richness estimates were indistinguishable after 659 and 385 camera nights of survey effort, respectively. We detected significant increases (ranging from 11–33%) in detection probability for five species resulting from the presence of game trails. For six species detection probability was also influenced by the presence of a log feature. This bias was most pronounced for the three rodents investigated, where in all cases detection probability was substantially higher (24.9–38.2%) at log cameras. Our results indicate that small-scale factors, including the presence of game trails and other features, can have significant impacts on species detection when camera traps are employed. Significant biases may result if the presence and quality of these features are not documented and either incorporated into analytical procedures, or controlled for in study design.
The recent development of fecal-genetic capture-mark-recapture (CMR) methods has increased the feasibility of estimating abundance of forest-dwelling ungulates that are difficult to survey using visual methods. Unless genetic markers differentiating sex are incorporated into such studies, however, genetic CMR approaches risk missing sex-specific differences in population trends. We developed a singlereaction genetic assay for sex and individual identification, including 10 microsatellites and an SRY marker, and applied it in the context of a post-fawning CMR study of Columbian black-tailed deer (Odocoileus hemionus columbianus) in forested habitat of coastal California during 2011 and 2012. We measured sexspecific abundance and sex ratios in high-quality summer habitats encompassing 4 distinct fawning areas. We detected a significant interaction between sex and year, indicating different trends in the abundance of males and females. We also detected a significant decline in abundance of females between years (P ¼ 0.045), which agreed with independent telemetry-based estimates, and significant differences in female abundance among fawning areas (P ¼ 0.020) but no significant differences in the abundance of males for either variable (F 1-3,20 < 0.710, P > 0.410). When sex was not considered in the analysis, we found no significant differences in abundance between the 2 years, suggesting that differing trends between the 2 sexes obscured the femalespecific patterns. We estimated average local (i.e., on the high-quality summer ranges) density (D) for females at 41.0 (AE 5.9) deer/km 2 in 2011 and 29.1 (AE 6.8) deer/km 2 in 2012, and local density of males at 15.7 (AE 3.0) deer/km 2 across the 2 study years. Accordingly, sex ratios differed between years (95% CI ¼ 3.0-4.2 F:M ratio in 2011, 2.0-2.3 F:M ratio in 2012). Incorporating sex and individual markers into a single assay provided a cost-effective means of applying CMR estimation based on fecal DNA to a high-density ungulate population in a forested ecosystem and emphasized the importance of explicitly modeling sex in abundance estimation. Ó
Summary Managed public wild areas have dual mandates to protect biodiversity and provide recreational opportunities for people. These goals could be at odds if recreation, ranging from hiking to legal hunting, disrupts wildlife enough to alter their space use or community structure. We evaluated the effect of managed hunting and recreation on 12 terrestrial wildlife species by employing a large citizen science camera trapping survey at 1947 sites stratified across different levels of human activities in 32 protected forests in the eastern USA. Habitat covariates, especially the amount of large continuous forest and local housing density, were more important than recreation for affecting the distribution of most species. The four most hunted species (white‐tailed deer, raccoons, eastern grey and fox squirrels) were commonly detected throughout the region, but relatively less so at hunted sites. Recreation was most important for affecting the distribution of coyotes, which used hunted areas more compared with unhunted control areas, and did not avoid areas used by hikers. Most species did not avoid human‐made trails, and many predators positively selected them. Bears and bobcats were more likely to avoid people in hunted areas than unhunted preserves, suggesting that they perceive the risk of humans differently depending on local hunting regulations. However, this effect was not found for the most heavily hunted species, suggesting that human hunters are not broadly creating ‘fear’ effects to the wildlife community as would be expected for apex predators. Synthesis and applications. Although we found that hiking and managed hunting have measureable effects on the distribution of some species, these were relatively minor in comparison with the importance of habitat covariates associated with land use and habitat fragmentation. These patterns of wildlife distribution suggest that the present practices for regulating recreation in the region are sustainable and in balance with the goal of protecting wildlife populations and may be facilitated by decades of animal habituation to humans. The citizen science monitoring approach we developed could offer a long‐term monitoring protocol for protected areas, which would help managers to detect where and when the balance between recreation and wildlife has tipped.
Human activity and land use change impact every landscape on Earth, driving declines in many animal species while benefiting others. Species ecological and life history traits may predict success in human‐dominated landscapes such that only species with “winning” combinations of traits will persist in disturbed environments. However, this link between species traits and successful coexistence with humans remains obscured by the complexity of anthropogenic disturbances and variability among study systems. We compiled detection data for 24 mammal species from 61 populations across North America to quantify the effects of (1) the direct presence of people and (2) the human footprint (landscape modification) on mammal occurrence and activity levels. Thirty‐three percent of mammal species exhibited a net negative response (i.e., reduced occurrence or activity) to increasing human presence and/or footprint across populations, whereas 58% of species were positively associated with increasing disturbance. However, apparent benefits of human presence and footprint tended to decrease or disappear at higher disturbance levels, indicative of thresholds in mammal species’ capacity to tolerate disturbance or exploit human‐dominated landscapes. Species ecological and life history traits were strong predictors of their responses to human footprint, with increasing footprint favoring smaller, less carnivorous, faster‐reproducing species. The positive and negative effects of human presence were distributed more randomly with respect to species trait values, with apparent winners and losers across a range of body sizes and dietary guilds. Differential responses by some species to human presence and human footprint highlight the importance of considering these two forms of human disturbance separately when estimating anthropogenic impacts on wildlife. Our approach provides insights into the complex mechanisms through which human activities shape mammal communities globally, revealing the drivers of the loss of larger predators in human‐modified landscapes.
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