Using global remote camera data of a solitary species complex to evaluate the drivers of group formation.

The social system of animals involves a complex interplay between physiology, natural history, and the environment. Long relied upon discrete categorizations of "social" and "solitary" inhibit our capacity to understand species and their interactions with the world around them. Here, we use a globally distributed camera trapping dataset to test the drivers of aggregating into groups in a species complex (martens and relatives, family Mustelidae, Order Carnivora) assumed to be obligately solitary. We use a simple quantification, the probability of being detected in a group, that was applied across our globally derived camera trap dataset. Using a series of binomial generalized mixed-effects models applied to a dataset of 16,483 independent detections across 17 countries on four continents we test explicit hypotheses about potential drivers of group formation. We observe a wide range of probabilities of being detected in groups within the solitary model system, with the probability of aggregating in groups varying by more than an order of magnitude. We demonstrate that a species' context-dependent proclivity toward aggregating in groups is underpinned by a range of resource-related factors, primarily the distribution of resources, with increasing patchiness of resources facilitating group formation, as well as interactions between environmental conditions (resource constancy/winter severity) and physiology (energy storage capabilities). The wide variation in propensities to aggregate with conspecifics observed here highlights how continued failure to recognize complexities in the social behaviors of apparently solitary species limits our understanding not only of the individual species but also the causes and consequences of group formation.

Dynamic occupancy modelling reveals a hierarchy of competition among fishers, grey foxes and ringtails.

Determining how species coexist is critical for understanding functional diversity, niche partitioning and interspecific interactions. Identifying the direct and indirect interactions among sympatric carnivores that enable their coexistence is particularly important to elucidate because they are integral for maintaining ecosystem function. We studied the effects of removing nine fishers (Pekania pennanti) on their population dynamics and used this perturbation to elucidate the interspecific interactions among fishers, grey foxes (Urocyon cinereoargenteus) and ringtails (Bassariscus astutus). Grey foxes (family: Canidae) are likely to compete with fishers due to their similar body sizes and dietary overlap, and ringtails (family: Procyonidae), like fishers, are semi-arboreal species of conservation concern. We used spatial capture-recapture to investigate fisher population numbers and dynamic occupancy models that incorporated interspecific interactions to investigate the effects members of these species had on the colonization and persistence of each other's site occupancy. The fisher population showed no change in density for up to 3 years following the removals of fishers for translocations. In contrast, fisher site occupancy decreased in the years immediately following the translocations. During this same time period, site occupancy by grey foxes increased and remained elevated through the end of the study. We found a complicated hierarchy among fishers, foxes and ringtails. Fishers affected grey fox site persistence negatively but had a positive effect on their colonization. Foxes had a positive effect on ringtail site colonization. Thus, fishers were the dominant small carnivore where present and negatively affected foxes directly and ringtails indirectly. Coexistence among the small carnivores we studied appears to reflect dynamic spatial partitioning. Conservation and management efforts should investigate how intraguild interactions may influence the recolonization of carnivores to previously occupied landscapes.

The effect of illumination and time of day on movements of bobcats (Lynx rufus).

Understanding behavioral changes of prey and predators based on lunar illumination provides insight into important life history, behavioral ecology, and survival information. The objectives of this research were to determine if bobcat movement rates differed by period of day (dark, moon, crepuscular, day), lunar illumination (<10%, 10 - <50%, 50 - <90%, >90%), and moon phase (new, full). Bobcats had high movement rates during crepuscular and day periods and low movement rates during dark periods with highest nighttime rates at 10-<50% lunar illumination. Bobcats had highest movement rates during daytime when nighttime illumination was low (new moon) and higher movement rates during nighttime when lunar illumination was high (full moon). The behaviors we observed are consistent with prey availability being affected by light level and by limited vision by bobcats during darkness.

Carnivore translocations and conservation: insights from population models and field data for fishers (Martes pennanti).

Translocations are frequently used to restore extirpated carnivore populations. Understanding the factors that influence translocation success is important because carnivore translocations can be time consuming, expensive, and controversial. Using population viability software, we modeled reintroductions of the fisher, a candidate for endangered or threatened status in the Pacific states of the US. Our model predicts that the most important factor influencing successful re-establishment of a fisher population is the number of adult females reintroduced (provided some males are also released). Data from 38 translocations of fishers in North America, including 30 reintroductions, 5 augmentations and 3 introductions, show that the number of females released was, indeed, a good predictor of success but that the number of males released, geographic region and proximity of the source population to the release site were also important predictors. The contradiction between model and data regarding males may relate to the assumption in the model that all males are equally good breeders. We hypothesize that many males may need to be released to insure a sufficient number of good breeders are included, probably large males. Seventy-seven percent of reintroductions with known outcomes (success or failure) succeeded; all 5 augmentations succeeded; but none of the 3 introductions succeeded. Reintroductions were instrumental in reestablishing fisher populations within their historical range and expanding the range from its most-contracted state (43% of the historical range) to its current state (68% of the historical range). To increase the likelihood of translocation success, we recommend that managers: 1) release as many fishers as possible, 2) release more females than males (55-60% females) when possible, 3) release as many adults as possible, especially large males, 4) release fishers from a nearby source population, 5) conduct a formal feasibility assessment, and 6) develop a comprehensive implementation plan that includes an active monitoring program.

A comparison of two field chemical immobilization techniques for bobcats (Lynx rufus).

Anesthetic protocols that allow quick induction, short processing time, and rapid reversal are necessary for researchers performing minimally invasive procedures (including morphometric measurements or attachment of radiocollars). The objective of this study was to evaluate the effectiveness of medetomidine and butorphanol as a substitute for xylazine in ketamine-based field immobilization protocols for bobcats (Lynx rufus) to reduce recovery and total field times. During 2008 and 2009, 11 bobcats were immobilized with an intramuscular combination of ketamine (10 mg/kg)-xylazine (0.75 mg/kg) (KX) or ketamine (4 mg/kg)-medetomidine (40 mcg/kg)-butorphanol (0.4 mg/kg) (KMB). Time to initial sedation, recumbency, and full anesthesia were recorded postinjection. Time to head up, sternal, standing, full recovery, and total processing times were recorded post-reversal. Throughout anesthesia, heart rate (HR), respiratory rate (RR), rectal temperature (RT), and noninvasive hemoglobin-oxygen saturation (SpO2) were recorded at 5-min intervals. The KX combination had a median time to full anesthesia of 10 min, a median recovery time of 46 min, and a median total processing time of 83 min. Alternatively, the KMB combination had a median time to full anesthesia of 21 min, a median recovery time of 18 min, and a median total processing time of 64 min. The KX protocol produced a median HR of 129 beats/min, RR of 25 breaths/min, RT of 38.3 degrees C, and SpO2 of 93%. The KMB protocol produced a median HR of 97 beats/min, RR of 33 breaths/min, RT of 38.4 degrees C, and SpO2 of 92%. Though both protocols provided safe and reliable sedation, the benefits of using medetomidine and butorphanol to lower ketamine doses and decrease processing time for brief nonsurgical sedation of bobcats in the field are presented.

Characterizing gonadal and adrenal activity by fecal steroid analyses in pygmy rabbits (Brachylagus idahoensis).

In 2001, the sudden collapse of the Columbia Basin population of pygmy rabbits prompted the initiation of a captive breeding program to facilitate reintroduction, but reproductive success in captivity has not met expectations. Therefore, the objective of this study was to characterize the reproductive and adrenal endocrinology of this endangered rabbit species so that appropriate management strategies could be developed to monitor animal welfare and increase reproduction. Fecal samples were collected from 27 female pygmy rabbits over three breeding and non-breeding seasons. HPLC analyses verified the presence of progesterone in the excreta of pygmy rabbits, but the majority of progestagen metabolites were unidentified polar compounds. By contrast, >70% of glucocorticoid immunoactivity was associated with cortisol. Longitudinal fecal hormone profiles during pregnancy were characterized by a large spike in progestagens shortly after mating, a gradual increase in progestagen and glucocorticoid concentrations throughout gestation and a return of hormones to baseline soon after birth (Day 24). The spike in progestagens 1 day after mating was a significant discovery for this species and appears to provide a reliable means of determining if a successful mating has occurred. Seasonal analyses of hormone excretion found that progestagen baselines did not vary between the breeding and non-breeding seasons, but, as expected, were highest during the breeding season in association with pregnancy. Across seasons, the lowest concentrations of glucocorticoids were associated with the highest rates of offspring production and survival, suggesting a possible link between heightened adrenal activity and lowered reproductive fitness in pygmy rabbits.

Animal ecology meets GPS-based radiotelemetry: a perfect storm of opportunities and challenges.

Global positioning system (GPS) telemetry technology allows us to monitor and to map the details of animal movement, securing vast quantities of such data even for highly cryptic organisms. We envision an exciting synergy between animal ecology and GPS-based radiotelemetry, as for other examples of new technologies stimulating rapid conceptual advances, where research opportunities have been paralleled by technical and analytical challenges. Animal positions provide the elemental unit of movement paths and show where individuals interact with the ecosystems around them. We discuss how knowing where animals go can help scientists in their search for a mechanistic understanding of key concepts of animal ecology, including resource use, home range and dispersal, and population dynamics. It is probable that in the not-so-distant future, intense sampling of movements coupled with detailed information on habitat features at a variety of scales will allow us to represent an animal's cognitive map of its environment, and the intimate relationship between behaviour and fitness. An extended use of these data over long periods of time and over large spatial scales can provide robust inferences for complex, multi-factorial phenomena, such as meta-analyses of the effects of climate change on animal behaviour and distribution.

The home-range concept: are traditional estimators still relevant with modern telemetry technology?

Recent advances in animal tracking and telemetry technology have allowed the collection of location data at an ever-increasing rate and accuracy, and these advances have been accompanied by the development of new methods of data analysis for portraying space use, home ranges and utilization distributions. New statistical approaches include data-intensive techniques such as kriging and nonlinear generalized regression models for habitat use. In addition, mechanistic home-range models, derived from models of animal movement behaviour, promise to offer new insights into how home ranges emerge as the result of specific patterns of movements by individuals in response to their environment. Traditional methods such as kernel density estimators are likely to remain popular because of their ease of use. Large datasets make it possible to apply these methods over relatively short periods of time such as weeks or months, and these estimates may be analysed using mixed effects models, offering another approach to studying temporal variation in space-use patterns. Although new technologies open new avenues in ecological research, our knowledge of why animals use space in the ways we observe will only advance by researchers using these new technologies and asking new and innovative questions about the empirical patterns they observe.

Building the bridge between animal movement and population dynamics.

While the mechanistic links between animal movement and population dynamics are ecologically obvious, it is much less clear when knowledge of animal movement is a prerequisite for understanding and predicting population dynamics. GPS and other technologies enable detailed tracking of animal location concurrently with acquisition of landscape data and information on individual physiology. These tools can be used to refine our understanding of the mechanistic links between behaviour and individual condition through 'spatially informed' movement models where time allocation to different behaviours affects individual survival and reproduction. For some species, socially informed models that address the movements and average fitness of differently sized groups and how they are affected by fission-fusion processes at relevant temporal scales are required. Furthermore, as most animals revisit some places and avoid others based on their previous experiences, we foresee the incorporation of long-term memory and intention in movement models. The way animals move has important consequences for the degree of mixing that we expect to find both within a population and between individuals of different species. The mixing rate dictates the level of detail required by models to capture the influence of heterogeneity and the dynamics of intra- and interspecific interaction.

Understanding contributions of cohort effects to growth rates of fluctuating populations.

1. Understanding contributions of cohort effects to variation in population growth of fluctuating populations is of great interest in evolutionary biology and may be critical in contributing towards wildlife and conservation management. Cohort-specific contributions to population growth can be evaluated using age-specific matrix models and associated elasticity analyses. 2. We developed age-specific matrix models for naturally fluctuating populations of stoats Mustela erminea in New Zealand beech forests. Dynamics and productivity of stoat populations in this environment are related to the 3-5 year masting cycle of beech trees and consequent effects on the abundance of rodents. 3. The finite rate of increase (lambda) of stoat populations in New Zealand beech forests varied substantially, from 1.98 during seedfall years to 0.58 during post-seedfall years. Predicted mean growth rates for stoat populations in continuous 3-, 4- or 5-year cycles are 0.85, 1.00 and 1.13. The variation in population growth was a consequence of high reproductive success of females during seedfall years combined with low survival and fertility of females of the post-seedfall cohort. 4. Variation in population growth was consistently more sensitive to changes in survival rates both when each matrix was evaluated in isolation and when matrices were linked into cycles. Relative contributions to variation in population growth from survival and fertility, especially in 0-1-year-old stoats, also depend on the year of the cycle and the number of transitional years before a new cycle is initiated. 5. Consequently, management strategies aimed at reducing stoat populations that may be best during one phase of the beech seedfall cycle may not be the most efficient during other phases of the cycle. We suggest that management strategies based on elasticities of vital rates need to consider how population growth rates vary so as to meet appropriate economic and conservation targets.

Trapping and marking terrestrial mammals for research: integrating ethics, performance criteria, techniques, and common sense.

We propose that researchers integrate ethics, performance criteria, techniques, and common sense when developing research trapping programs and in which members of institutional animal care and use committees address these topics when evaluating research protocols. To ask questions about ethics is in the best tradition of science, and researchers must be familiar with codes of ethics and guidelines for research published by professional societies. Researchers should always work to improve research methods and to decrease the effects on research animals, if for no other reason than to minimize the chances that the methods influence the animals' behavior in ways that affect research results. Traps used in research should meet performance criteria that address state-of-the-art trapping technology and that optimize animal welfare conditions within the context of the research. The proposal includes the following criteria for traps used in research: As Criterion I, killing-traps should render >/= 70% of animals caught irreversibly unconscious in /= 70% of animals with