American marten and fisher do not segregate in space and time during winter in a mixed-forest system.

Understanding the mechanisms of coexistence between ecologically similar species is an important issue in ecology. Carnivore coexistence may be facilitated by spatial segregation, temporal avoidance, and differential habitat selection. American martens Martes americana and fishers Pekania pennanti are medium-sized mustelids that occur sympatrically across portions of North America, yet mechanisms of coexistence between the two species are not fully understood. We assessed spatial and temporal partitioning in martens and fishers in the Upper Peninsula of Michigan, USA, using camera trap data collected during winter 2013-2015. To investigate spatial segregation, we used a dynamic occupancy model to estimate species' occupancy probabilities and probabilities of persistence and colonization as a function of covariates and yearly occupancy probability for the other species. Temporal segregation was assessed by estimating diel activity overlap between species. We found weak evidence of spatial or temporal niche partitioning of martens and fishers. There was high overlap in forest cover selection, and both marten and fisher occupancy were positively correlated with deciduous forests (excluding aspen [Populus tremuloides]). There was strong temporal overlap ( Δ ^ 4 = 0.81 ; CI = 0.79-0.82) with both species exhibiting largely crepuscular activity patterns. Co-occurrence of martens and fishers appears to be facilitated by mechanisms not investigated in this study, such as partitioning of snow features or diet. Our results add additional insights into resource partitioning of mesocarnivores, but further research is required to enhance our understanding of mechanisms that facilitate marten and fisher coexistence.

Using multiple data types and integrated population models to improve our knowledge of apex predator population dynamics.

Current management of large carnivores is informed using a variety of parameters, methods, and metrics; however, these data are typically considered independently. Sharing information among data types based on the underlying ecological, and recognizing observation biases, can improve estimation of individual and global parameters. We present a general integrated population model (IPM), specifically designed for brown bears (Ursus arctos), using three common data types for bear (U. spp.) populations: repeated counts, capture-mark-recapture, and litter size. We considered factors affecting ecological and observation processes for these data. We assessed the practicality of this approach on a simulated population and compared estimates from our model to values used for simulation and results from count data only. We then present a practical application of this general approach adapted to the constraints of a case study using historical data available for brown bears on Kodiak Island, Alaska, USA. The IPM provided more accurate and precise estimates than models accounting for repeated count data only, with credible intervals including the true population 94% and 5% of the time, respectively. For the Kodiak population, we estimated annual average litter size (within one year after birth) to vary between 0.45 [95% credible interval: 0.43; 0.55] and 1.59 [1.55; 1.82]. We detected a positive relationship between salmon availability and adult survival, with survival probabilities greater for females than males. Survival probabilities increased from cubs to yearlings to dependent young ≥2 years old and decreased with litter size. Linking multiple information sources based on ecological and observation mechanisms can provide more accurate and precise estimates, to better inform management. IPMs can also reduce data collection efforts by sharing information among agencies and management units. Our approach responds to an increasing need in bear populations' management and can be readily adapted to other large carnivores.

Estimating Lion Abundance using N-mixture Models for Social Species.

Declining populations of large carnivores worldwide, and the complexities of managing human-carnivore conflicts, require accurate population estimates of large carnivores to promote their long-term persistence through well-informed management We used N-mixture models to estimate lion (Panthera leo) abundance from call-in and track surveys in southeastern Serengeti National Park, Tanzania. Because of potential habituation to broadcasted calls and social behavior, we developed a hierarchical observation process within the N-mixture model conditioning lion detectability on their group response to call-ins and individual detection probabilities. We estimated 270 lions (95% credible interval = 170-551) using call-ins but were unable to estimate lion abundance from track data. We found a weak negative relationship between predicted track density and predicted lion abundance from the call-in surveys. Luminosity was negatively correlated with individual detection probability during call-in surveys. Lion abundance and track density were influenced by landcover, but direction of the corresponding effects were undetermined. N-mixture models allowed us to incorporate multiple parameters (e.g., landcover, luminosity, observer effect) influencing lion abundance and probability of detection directly into abundance estimates. We suggest that N-mixture models employing a hierarchical observation process can be used to estimate abundance of other social, herding, and grouping species.

Effects of Prey Presence and Scale on Bobcat Resource Selection during Winter.

Factors relevant to resource selection in carnivores may vary across spatial and temporal scales, both in magnitude and rank. Understanding relationships among carnivore occupancy, prey presence, and habitat characteristics, as well as their interactions across multiple scales, is necessary to improve our understanding of resource selection and predict population changes. We used a multi-scale dynamic hierarchical co-occurrence model with camera data to study bobcat and snowshoe hare occupancy in the Upper Peninsula of Michigan during winter 2012-2013. Bobcat presence was influenced at the local scale by snowshoe hare presence, and by road density at the local and larger scale when hare were absent. Hare distribution was related primarily to vegetation cover types, and detectability varied in space and time. Bobcat occupancy dynamics were influenced by different factors depending on the spatial scale considered and the resource availability context. Moreover, considering observed co-occurrence, we suggest that bobcat presence had a greater effect on hare occupancy than hare presence on bobcat occupancy. Our results highlight the importance of studying carnivore distributions in the context of predator-prey relationships and its interactions with environmental covariates at multiple spatial scales. Our approach can be applied to other carnivore species to provide insights beneficial for management and conservation.

Estimating upper bounds for occupancy and number of manatees in areas potentially affected by oil from the Deepwater Horizon oil spill.

The explosion of the Deepwater Horizon drilling platform created the largest marine oil spill in U.S. history. As part of the Natural Resource Damage Assessment process, we applied an innovative modeling approach to obtain upper estimates for occupancy and for number of manatees in areas potentially affected by the oil spill. Our data consisted of aerial survey counts in waters of the Florida Panhandle, Alabama and Mississippi. Our method, which uses a Bayesian approach, allows for the propagation of uncertainty associated with estimates from empirical data and from the published literature. We illustrate that it is possible to derive estimates of occupancy rate and upper estimates of the number of manatees present at the time of sampling, even when no manatees were observed in our sampled plots during surveys. We estimated that fewer than 2.4% of potentially affected manatee habitat in our Florida study area may have been occupied by manatees. The upper estimate for the number of manatees present in potentially impacted areas (within our study area) was estimated with our model to be 74 (95%CI 46 to 107). This upper estimate for the number of manatees was conditioned on the upper 95%CI value of the occupancy rate. In other words, based on our estimates, it is highly probable that there were 107 or fewer manatees in our study area during the time of our surveys. Because our analyses apply to habitats considered likely manatee habitats, our inference is restricted to these sites and to the time frame of our surveys. Given that manatees may be hard to see during aerial surveys, it was important to account for imperfect detection. The approach that we described can be useful for determining the best allocation of resources for monitoring and conservation.

Modeling trends from North American breeding bird survey data: a spatially explicit approach.

Population trends, defined as interval-specific proportional changes in population size, are often used to help identify species of conservation interest. Efficient modeling of such trends depends on the consideration of the correlation of population changes with key spatial and environmental covariates. This can provide insights into causal mechanisms and allow spatially explicit summaries at scales that are of interest to management agencies. We expand the hierarchical modeling framework used in the North American Breeding Bird Survey (BBS) by developing a spatially explicit model of temporal trend using a conditional autoregressive (CAR) model. By adopting a formal spatial model for abundance, we produce spatially explicit abundance and trend estimates. Analyses based on large-scale geographic strata such as Bird Conservation Regions (BCR) can suffer from basic imbalances in spatial sampling. Our approach addresses this issue by providing an explicit weighting based on the fundamental sample allocation unit of the BBS. We applied the spatial model to three species from the BBS. Species have been chosen based upon their well-known population change patterns, which allows us to evaluate the quality of our model and the biological meaning of our estimates. We also compare our results with the ones obtained for BCRs using a nonspatial hierarchical model (Sauer and Link 2011). Globally, estimates for mean trends are consistent between the two approaches but spatial estimates provide much more precise trend estimates in regions on the edges of species ranges that were poorly estimated in non-spatial analyses. Incorporating a spatial component in the analysis not only allows us to obtain relevant and biologically meaningful estimates for population trends, but also enables us to provide a flexible framework in order to obtain trend estimates for any area.

Dynamic occupancy models for analyzing species' range dynamics across large geographic scales.

Large-scale biodiversity data are needed to predict species' responses to global change and to address basic questions in macroecology. While such data are increasingly becoming available, their analysis is challenging because of the typically large heterogeneity in spatial sampling intensity and the need to account for observation processes. Two further challenges are accounting for spatial effects that are not explained by covariates, and drawing inference on dynamics at these large spatial scales. We developed dynamic occupancy models to analyze large-scale atlas data. In addition to occupancy, these models estimate local colonization and persistence probabilities. We accounted for spatial autocorrelation using conditional autoregressive models and autologistic models. We fitted the models to detection/nondetection data collected on a quarter-degree grid across southern Africa during two atlas projects, using the hadeda ibis (Bostrychia hagedash) as an example. The model accurately reproduced the range expansion between the first (SABAP1: 1987-1992) and second (SABAP2: 2007-2012) Southern African Bird Atlas Project into the drier parts of interior South Africa. Grid cells occupied during SABAP1 generally remained occupied, but colonization of unoccupied grid cells was strongly dependent on the number of occupied grid cells in the neighborhood. The detection probability strongly varied across space due to variation in effort, observer identity, seasonality, and unexplained spatial effects. We present a flexible hierarchical approach for analyzing grid-based atlas data using dynamical occupancy models. Our model is similar to a species' distribution model obtained using generalized additive models but has a number of advantages. Our model accounts for the heterogeneous sampling process, spatial correlation, and perhaps most importantly, allows us to examine dynamic aspects of species ranges.

Assessing hypotheses about nesting site occupancy dynamics.

Hypotheses about habitat selection developed in the evolutionary ecology framework assume that individuals, under some conditions, select breeding habitat based on expected fitness in different habitat. The relationship between habitat quality and fitness may be reflected by breeding success of individuals, which may in turn be used to assess habitat quality. Habitat quality may also be assessed via local density: if high-quality sites are preferentially used, high density may reflect high-quality habitat. Here we assessed whether site occupancy dynamics vary with site surrogates for habitat quality. We modeled nest site use probability in a seabird subcolony (the Black-legged Kittiwake, Rissa tridactyla) over a 20-year period. We estimated site persistence (an occupied site remains occupied from time t to t+1) and colonization through two subprocesses: first colonization (site creation at the timescale of the study) and recolonization (a site is colonized again after being deserted). Our model explicitly incorporated site-specific and neighboring breeding success and conspecific density in the neighborhood. Our results provided evidence that reproductively "successful" sites have a higher persistence probability than "unsuccessful" ones. Analyses of site fidelity in marked birds and of survival probability showed that high site persistence predominantly reflects site fidelity, not immediate colonization by new owners after emigration or death of previous owners. There is a negative quadratic relationship between local density and persistence probability. First colonization probability decreases with density, whereas recolonization probability is constant. This highlights the importance of distinguishing initial colonization and recolonization to understand site occupancy. All dynamics varied positively with neighboring breeding success. We found evidence of a positive interaction between site-specific and neighboring breeding success. We addressed local population dynamics using a site occupancy approach integrating hypotheses developed in behavioral ecology to account for individual decisions. This allows development of models of population and metapopulation dynamics that explicitly incorporate ecological and evolutionary processes.

Hierarchical modeling of an invasive spread: the Eurasian Collared-Dove Streptopelia decaocto in the United States.

Invasive species are regularly claimed as the second threat to biodiversity. To apply a relevant response to the potential consequences associated with invasions (e.g., emphasize management efforts to prevent new colonization or to eradicate the species in places where it has already settled), it is essential to understand invasion mechanisms and dynamics. Quantifying and understanding what influences rates of spatial spread is a key research area for invasion theory. In this paper, we develop a model to account for occupancy dynamics of an invasive species. Our model extends existing models to accommodate several elements of invasive processes; we chose the framework of hierarchical modeling to assess site occupancy status during an invasion. First, we explicitly accounted for spatial structure and how distance among sites and position relative to one another affect the invasion spread. In particular, we accounted for the possibility of directional propagation and provided a way of estimating the direction of this possible spread. Second, we considered the influence of local density on site occupancy. Third, we decided to split the colonization process into two subprocesses, initial colonization and recolonization, which may be ground-breaking because these subprocesses may exhibit different relationships with environmental variations (such as density variation) or colonization history (e.g., initial colonization might facilitate further colonization events). Finally, our model incorporates imperfection in detection, which might be a source of substantial bias in estimating population parameters. We focused on the case of the Eurasian Collared-Dove (Streptopelia decaocto) and its invasion of the United States since its introduction in the early 1980s, using data from the North American BBS (Breeding Bird Survey). The Eurasian Collared-Dove is one of the most successful invasive species, at least among terrestrial vertebrates. Our model provided estimation of the spread direction consistent with empirical observations. Site persistence probability exhibits a quadratic response to density. We also succeeded at detecting differences in the relationship between density and initial colonization vs. recolonization probabilities. We provide a map of sites that may be colonized in the future as an example of possible practical application of our work.

Estimating the occupancy rate of spatially rare or hard to detect species: a conditional approach.

We consider the problem of estimating the occupancy rate of a target species in a region divided in spatial units (called quadrats); this quantity being defined as the proportion of quadrats occupied by this species. We mainly focus on spatially rare or hard to detect species that are typically detected in very few quadrats, and for which estimating the occupancy rate (with an acceptable precision) is problematic. We develop a conditional approach for estimating the quantity of interest; we condition on the presence of the target species in the region of study. We show that conditioning makes identifiable the occurrence and detectability parameters, regardless of the number of visits made in the sampled quadrats. Compared with an unconditional approach, it proves to be complementary, in that this allows us to deal with biological questions that cannot be addressed by the former. Two Bayesian analyses of the data are performed: one is noninformative, and the other takes advantage of the fact that some prior information on detectability is available. It emerges that taking such a prior into account significantly improves the precision of the estimate when the target species has been detected in few quadrats and is known to be easily detectable.