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Matthew J. Kauffman

Publications and source records attributed to Matthew J. Kauffman.

At least 55 records · Page 3Linked to original sources

Ungulate migrations of the western United States, volume 3

Ungulates (hooved mammals) have a broad distribution across the western United States and play an important role in maintaining predator-prey dynamics, affecting vegetation communities, and providing economic benefits to regional communities through tourism and hunting. Throughout the diverse landscapes they occupy, many ungulate populations undertake seasonal migrations to exploit spatially and temporally variable resources and to avoid predation or other threats. As the human footprint continues to expand across the western United States, ungulates increasingly face more obstacles on their migratory journeys. These obstacles threaten the long-term persistence of existing migrations. As a result, wildlife management agencies across the western United States have worked to identify and protect (or enhance) ungulate migration corridors and seasonal ranges identified from global positioning system (GPS) collar data. These efforts garnered additional support through the U.S. Department of the Interior Secretarial Order (SO) 3362, which was initiated in 2018 and provided Federal support for enhancing habitat quality of big-game winter ranges and migration corridors across the western states. Further, SO 3362 prompted the U.S. Geological Survey (USGS) to establish the Corridor Mapping Team (CMT): a collaboration between USGS and participating State and Federal wildlife management agencies, as well as numerous Tribal Nations. The CMT works collaboratively to map ungulate migrations and seasonal ranges throughout the western United States within the Ungulate Migrations of the Western United States report series. Volume 1 of the series was published in 2020 and contained migrations and winter ranges from 42 herds across 5 states. Volume 2 was published in 2022 and contained migrations and seasonal ranges from an additional 65 herds. This report, Volume 3 in the series, details migrations and seasonal ranges from an additional 45 herds throughout most western states. In aggregate, the report series has detailed and mapped the migrations and seasonal ranges of 152 ungulate herds and serves as a map-based inventory of the documented ungulate migrations across the western United States. The data layers for most of the herds included in the report series are also available to the public by the USGS. In addition to the included herd maps, this volume provides an overview of the many ways the mapping efforts associated with the CMT are being integrated into local conservation, management, and policy throughout the western United States.

Arizona, California, Idaho, Nevada, New Mexico, Or↗

Evaluating risks associated with capture and handling of mule deer for individual-based, long-term research

Capture and handling techniques for individual-based, long-term research that tracks the life history of animals by recapturing the same individuals for several years has vastly improved study inferences and our understanding of animal ecology. Yet there are corresponding risks to study animals associated with physical trauma or capture myopathy that can occur during or following capture events. Rarely has empirical evidence existed to guide decisions associated with understanding the magnitude of capture-related risks, how to reduce these risks when possible, and implications for mortality censoring and survival estimates. We used data collected from 2,399 capture events of mule deer ( Odocoileus hemionus ) via helicopter net-gunning to compare daily survival probabilities within a 10-week period centered on a capture event and evaluated how animal age, nutritional condition (body fat), and various handling methods influenced survival before, during, and following a capture event. Direct mortality resulting from capture efforts was 1.59%. Mean daily survival was 0.9993 ± 0.0001 (SE) during the 5-week pre-capture window, was depressed the day of capture at 0.9841 ± 0.0004, and rebounded to 0.9990 ± 0.0008 during the 5-week post-capture window. Neither capture nor handling had a detectable effect on post-capture survival, including handling time ( x̄ = 13.30 ± 1.87 min), capture time of year (i.e., Dec or Mar), tooth extraction, and the number of times an animal had been recaptured (2–17 times). Although mortality rate was slightly elevated during capture (resulting from physical trauma associated with capture), age and nutritional condition did not influence the probability of mortality during a capture event. Following a capture event, nutritional condition influenced survival; however, that relationship was consistent with expected effects of nutritional condition on winter survival and independent of capture and handling. Overall survival rates 5 weeks before capture and 5 weeks after capture were not different. A specified window of time with depressed survival following capture and handling was not evident, which contradicts the implementation of a predetermined window often used by researchers and managers for censoring mortalities that occur after capture. Previous notions that censorship of all mortality data in the 2 weeks following capture is unwarranted and risks removal of meaningful data. With previous evidence guiding our protocols for capture (e.g., reduced chase time) and handling (e.g., temperature mitigation), low direct mortality and almost undetectable indirect mortality post capture reinforces the efficacy of helicopter net-gunning for capture and recapture of mule deer in long-term, individual-based studies.

Wyoming↗

Industrial energy development decouples ungulate migration from the green wave

The ability to freely move across the landscape to track the emergence of nutritious spring green-up (termed ‘green-wave surfing’) is key to the foraging strategy of migratory ungulates. Across the vast landscapes traversed by many migratory herds, habitats are being altered by development with unknown consequences for surfing. Using a unique long-term tracking dataset, we found that when energy development occurs within mule deer ( Odocoileus hemionus ) migration corridors, migrating animals become decoupled from the green wave. During the early phases of a coalbed natural gas development, deer synchronized their movements with peak green-up. But faced with increasing disturbance as development expanded, deer altered their movements by holding up at the edge of the gas field and letting the green wave pass them by. Development often modified only a small portion of the migration corridor but had far-reaching effects on behaviour before and after migrating deer encountered it, thus reducing surfing along the entire route by 38.65% over the 14-year study period. Our study suggests that industrial development within migratory corridors can change the behaviour of migrating ungulates and diminish the benefits of migration. Such disruptions to migratory behaviour present a common mechanism whereby corridors become unprofitable and could ultimately be lost on highly developed landscapes.

Wyoming↗

A statistical framework for modelling migration corridors

1. Management of animal populations requires spatially explicit knowledge of movement corridors, such as those used during seasonal migrations. GPS tracking data allows for mapping of corridors from directly observed movements, providing important insights, but tracking data is absent for many populations. 2. We developed a novel statistical corridor modeling approach that predicts movement corridors from cost-distance movement models fit directly to migration tracking data. Unlike existing predictive approaches, this does not require the ad-hoc transformation of habitat suitability surfaces into resistance surfaces. We tested the ability of the approach to recover parameters used to generate simulated movements. We then used GPS data from three migrating mule deer (Odocoileus hemionus) herds in Idaho and Wyoming to model corridors as a function of elevation, slope, aspect, percent shrub, date of peak green-up, snow-off date, and human footprint. We assessed the predictive ability of the fitted models using validation tracks from the same herd as well as from the other herds. 3. The approach reproduced parameters used to generate the simulated movements, predicted the corridors used by migratory populations, and described the direction, magnitude, and confidence levels of the effects of environmental variables on corridors. The effects environmental variables had on corridors differed depending on the herd. Within-herd validation indicated that fitted corridor models are more accurate at predicting migration corridors than null models, and cross-herd validation indicated that fitted models for some herds accurately predicted the observed migrations of other herds. 4. In addition to the practical benefit of mapping corridors for management, our statistical corridor modeling framework sets the stage for evaluating fundamental questions about the fitness tradeoffs, navigation, learning, fidelity, and movement constraints that influence migratory and other corridor-generating behavior. Models of predictive corridors can inform management and planning for the conservation of migrations across taxa, including the potential restoration of corridors. Our corridor modeling approach is also readily applied to non-migratory animal movements.

Idaho, Wyoming↗

Migration Mapper: Identifying movement corridors and seasonal ranges for large mammal conservation

Modern tracking technology has facilitated a novel understanding of terrestrial mammal movement while revealing that movements are being truncated and lost. The first step towards conserving mobile animals is identifying movement corridors and key seasonal ranges. Yet, the identification and subsequent mapping of these important areas has remained a challenge due to the analytical skills necessary to conduct such analyses. Migration Mapper (MM) is a user-friendly software that provides tools to analyse global positioning system (GPS) collar data to create season-specific, population-level polygons representing areas where most of a population moves (i.e. movement corridors) and areas where most of a population spends time (e.g. high-use areas, seasonal ranges). MM consists of six standalone modules including data cleaning and review, seasonal movement delineation, movement model application, calculation of population-level outputs and visualization of results. Analysis of GPS data using MM can provide the spatial polygons necessary to facilitate conservation and policy planning. New initiatives at the local and global levels are already beginning to use MM to facilitate conservation of large, terrestrial mammals.

Methods in Ecology and Evolution↗

Trade-offs between utility-scale solar development and ungulates on western rangelands

Utility-scale solar energy (USSE) has become an efficient and cost-effective form of renewable energy, with an expanding footprint into rangelands that provide important habitat for many wild ungulate populations. Using global positioning system data collected before and after construction, we documented the potential impacts of USSE on pronghorn ( Antilocapra americana ), including direct habitat loss, indirect habitat loss, and barrier effects to both resident and migratory population segments. Our case study highlights the challenges that USSE poses to ungulate conservation, including (1) impermeable security fencing that blocks access to and reduces connectivity between formerly available habitats, and (2) the lack of guidelines for minimizing USSE impacts on ungulates. Improved siting and ungulate-specific best management practices would help to minimize habitat loss and retain landscape connectivity. Ungulate biodiversity and ecosystem services (for example, services provided by long-distance migratory species) in arid rangelands are important considerations when balancing the global benefits of renewable energy with local wildlife impacts.

Wyoming↗

Ungulate migrations of the western United States, volume 2

Migration is widespread across taxonomic groups and increasingly recognized as fundamental to maintaining abundant wildlife populations and communities. Many ungulate herds migrate across the western United States to access food and avoid harsh environmental conditions. With the advent of global positioning system (GPS) collars, researchers can describe and map the year-round movements of ungulates at both large and small spatial scales. The migrations can traverse landscapes that are a mix of different jurisdictional ownership and management. Today, the landscapes migrating herds traverse are increasingly threatened by fencing, high-traffic roads, oil and gas development, and other types of permanent development. Through the use of GPS collars, a model of science-based conservation emerged in which migration corridors, stopovers, and winter ranges can be mapped in detail, thereby allowing threats and conservation opportunities to be identified and remedied. In 2018, the U.S. Geological Survey (USGS) assembled a Corridor Mapping Team (CMT) to work collaboratively with western states to map migrations of Odocoileus hemionus (mule deer), Cervus canadensis (elk), and Antilocapra americana (pronghorn). Led by the USGS Wyoming Cooperative Fish and Wildlife Research Unit, the team consists of Federal scientists, university researchers, and biologists and analysts from participating State and Tribal agencies. The first set of maps described a total of 42 migrations across 5 western states and was published in 2020 as the first volume of this report series. This second volume describes an additional 65 migrations mapped within 9 western states and select Tribal lands. As the western United States continues to grow, this report series and the associated map files released by the USGS will allow for migration maps to be used for conservation planning by a wide array of State and Federal stakeholders to reduce barriers to migration caused by fences, roads, and other development.

western United States↗

Wildlife migrations highlight importance of both private lands and protected areas in the Greater Yellowstone Ecosystem

Formally protected areas are an important component of wildlife conservation, but face limitations in their effectiveness for migratory species. Improved stewardship of working lands around protected areas is one solution for conservation planning, but private working lands are vulnerable to development. In the Greater Yellowstone Ecosystem (GYE), ungulates such as elk ( Cervus canadensis ) use both protected areas and private lands throughout their annual migrations. We studied patterns of landownership, protection, and conservation challenges within the ranges of migratory elk in the GYE. We used GPS data from 1088 elk in 26 herds to define herd-level seasonal ranges, and extracted covariates related to landownership and protection, land use, and human infrastructure and development. All elk herds used land encompassing >1 ownership type. Most elk herds (92.3 % of herds, n = 24) used the highest proportion of private land in the winter (mean = 36.2 % private land). Most elk herds' winter ranges contained the highest building densities (mean = 1.24 buildings/km 2 ), fence densities (mean = 1.02 km fence/km 2 ), and cattle grazing (mean = 1.9 cows/km 2 ), compared to migratory and summer ranges. Out of all ranges, 36.5 % of ranges did not have any zoning regulations, indicating the potential for future development. Our results show that elk in the GYE rely on habitat outside of protected areas, and face landscape-scale conservation challenges such as habitat fragmentation from human development, particularly in winter ranges. Future conservation strategies for wildlife in this system need to encompass coordination across both public and private land to ensure migratory connectivity.

Idaho, Montana, Wyoming↗

Extending body condition scoring beyond measurable rump fat to estimate full range of nutritional condition for moose

Moose ( Alces alces ) populations along the southern extent of their range are largely declining, and there is growing evidence that nutritional condition — which influences several vital rates – is a contributing factor. Moose body condition can presently be estimated only when there is measurable subcutaneous rump fat, which equates to animals with >6% ingesta-free body fat (IFBFat). There is need for a technique to allow body fat estimation of animals in poorer body condition (i.e., <6% body fat). We advance current methods for moose, following those used and validated with other ungulate species, by establishing a moose-specific body condition score (BCS) that can be used to estimate IFBFat in the lower range of condition. Our modified BCS was related strongly ( r 2 = 0.89) to IFBFat estimates based on measurable rump fat. By extending the predicted relationship to individuals without measurable fat, the BCS equated severe emaciation with 0.67% IFBFat, supporting the accuracy of the method. The lower end of nutritional condition is important for identifying relationships involving life-history characteristics because most state-dependent changes occur at lower levels of condition. Therefore, until the BCS can be validated with moose carcasses, we believe our method to estimate body fat across the full range of condition should yield better understanding of the drivers underlying declining moose populations.

Alces↗

Variable effects of wind-energy development on seasonal habitat selection of pronghorn

In the face of climate change, wind energy represents an important alternative to oil and gas extraction to meet increasing energy demands, but it has the potential to disrupt wildlife populations. Because behavioral adjustments, such as altered habitat selection, are a primary way that long-lived species respond to novel disturbances, we evaluated effects of wind energy development on pronghorn ( Antilocapra americana ) space use and habitat selection. Using data from GPS-collared female pronghorn in the Shirley Basin of south-central Wyoming, USA, we tested four potential effects of wind turbines on pronghorn space use during the summer and winter: (1) displacement away from wind turbines, (2) increase in size of home ranges, (3) short-term avoidance behavior within home ranges, and (4) changes in avoidance behavior within home ranges over time. We monitored 166 individuals over five summers (2010, 2011, 2018, 2019, and 2020) and 142 individuals over five winters (2009/2010, 2010/2011, 2011/2012, 2018/2019, and 2019/2020) and used resource selection functions to evaluate selection relative to turbines after controlling for other habitat factors, such as snow depth. Although a lack of consistent negative effects of wind turbines on pronghorn across years suggested that wind energy development may have less severe and more intermittent effects on pronghorn than oil and gas development has had on other ungulates, there was a trend toward increased displacement during the study and behavioral avoidance was apparent for individuals in close proximity to turbines. However, pronghorn were highly variable in their fine-scale habitat selection, across both individuals and years, which could make effects of wind energy development difficult to detect. Nevertheless, some individuals, particularly those close to wind-energy facilities, did avoid turbines, which could translate to population-level behavioral or demographic changes over time and affect the resilience and stability of the population. Over time, the accumulation of development, including wind turbines, roads, and fences, can both limit movement and fragment habitat, potentially reaching a critical threshold beyond which populations are negatively impacted.

Wyoming↗

Body size and digestive system shape resource selection by ungulates: A cross-taxa test of the forage maturation hypothesis

The forage maturation hypothesis (FMH) states that energy intake for ungulates is maximised when forage biomass is at intermediate levels. Nevertheless, metabolic allometry and different digestive systems suggest that resource selection should vary across ungulate species. By combining GPS relocations with remotely sensed data on forage characteristics and surface water, we quantified the effect of body size and digestive system in determining movements of 30 populations of hindgut fermenters (equids) and ruminants across biomes. Selection for intermediate forage biomass was negatively related to body size, regardless of digestive system. Selection for proximity to surface water was stronger for equids relative to ruminants, regardless of body size. To be more generalisable, we suggest that the FMH explicitly incorporate contingencies in body size and digestive system, with small-bodied ruminants selecting more strongly for potential energy intake, and hindgut fermenters selecting more strongly for surface water.

Ecology Letters↗

U.S. Geological Survey science for the Wyoming Landscape Conservation Initiative—2018 annual report

The Wyoming Landscape Conservation Initiative (WLCI) was established in 2007 as a collaborative interagency partnership to develop and implement science-based conservation actions. During the past 11 years, partners from U.S. Geological Survey (USGS), State and Federal land management agencies, universities, and the public have collaborated to implement a long-term (more than 10 years) science-based program that assesses and enhances the quality and quantity of wildlife habitats in the southwest Wyoming region while facilitating responsible development. The USGS WLCI Science Team completes scientific research and develops tools that inform and support WLCI partner planning, decision making, and on-the-ground management actions. In fiscal year 2018, the USGS initiated 3 new projects and continued efforts on 21 ongoing science and web-development projects. The first new project was initiated to support Secretarial Order 3362 which calls on the USGS to assist Western States in mapping big-game migration corridors and developing new mapping tools. During 2018, the USGS hosted a workshop in Laramie, Wyoming, which included more than 70 State and Federal wildlife experts from Colorado, New Mexico, Texas, and Wyoming. Most of the mapping and migration tool curricula used in the workshop were derived from prior WLCI studies and mapping efforts of big-game migration movement in habitats undergoing large-scale energy development. The second new project was in response for WLCI partners to better understand sedimentation and hydrogeomorphic processes in a cold-desert headwater and the third new project was designed to improve our approach for people to access, manage, and analyze WLCI data and WLCI resource information. The USGS published 18 products (including peer-reviewed journal articles, USGS series publications, and data releases) and provided more than a dozen professional oral and poster presentations at scientific meetings and numerous informal presentations to WLCI partners at meetings and workshops. This report summarizes the objectives and status of each project and highlights the USGS 2018 accomplishments and products.

Wyoming↗

Short-term responses to a human-altered landscape do not affect fat dynamics of a migratory ungulate

According to risk-sensitive foraging theory, animals should make foraging decisions that balance nutritional costs and gains to promote fitness. Human disturbance is a form of perceived risk that can prompt avoidance of risky habitat over acquisition of food. Consequently, behavioural responses to perceived risk could induce nutritional costs. Population declines often coincide with increases in human disturbance, which likely is associated with direct and indirect habitat loss. Nevertheless, behavioural and physiological responses to perceived risks associated with human disturbance could be an added nutritional deficit with population-level repercussions. Using GPS-collar data from three populations of migratory mule deer Odocoileus hemionus exposed to a gradient of established industrial energy development on winter ranges where direct and indirect habitat loss were well documented, we evaluated whether exposure and behavioural responses to human disturbance alter changes in nutritional condition (i.e. fat reserves) over winter. Although animals exhibited behaviours indicative of perceived risk of human disturbance, such as increased movement rates and avoidance of infrastructure, exposure and behavioural responses to human disturbance had little to no measurable effect on fat loss over a winter. Instead, catabolism of fat reserves occurred primarily as a function of the amount of fat animals had entering winter, suggesting that, in the short term, animals were able to mitigate the energetic costs of perceived risk of human disturbance over winter. Animals, however, did not appear to overcome persistent food limitations, and animals with less food availability lost more fat over winter. Our findings heed caution in using short-term behavioural and physiological responses to inform long-term nutritional consequences of human disturbance. Although animals appear to mitigate the energetic costs of perceived risk, food limitations exacerbated by broader-scale avoidance of food near human disturbance may be the primary pathway causing the frequently observed population declines following human disturbance to pristine landscapes.

Wyoming↗

Functional connectivity in a continuously distributed, migratory species as revealed by landscape genomics

Maintaining functional connectivity is critical for the long-term conservation of wildlife populations. Landscape genomics provides an opportunity to assess long-term functional connectivity by relating environmental variables to spatial patterns of genomic variation resulting from generations of movement, dispersal and mating behaviors. Identifying landscape features associated with gene flow at large geographic scales for highly mobile species is becoming increasingly possible due to more accessible genomic approaches, improved analytical methods and enhanced computational power. We characterized the genetic structure and diversity of migratory mule deer Odocoileus hemionus using 4051 single nucleotide polymorphisms in 406 individuals sampled across multiple habitats throughout Wyoming, USA. We then identified environmental variables associated with genomic variation within genetic groups and statewide using a stepwise approach to first evaluate nonlinear relationships of landscape resistance with genetic distances and then use mixed-effects modeling to choose top landscape genomic models. We identified three admixed genetic groups of mule deer and found that environmental variables associated with gene flow varied among genetic groups, revealing scale-dependent and regional variation in functional connectivity. At the statewide scale, more gene flow occurred in areas with low elevation and mixed habitat. In the southern genetic group, more gene flow occurred in areas with low elevation. In the northern genetic group, more gene flow occurred in grassland and forest habitats, while highways and energy infrastructure reduced gene flow. In the western genetic group, the null model of isolation by distance best represented genetic patterns. Overall, our findings highlight the role of different seasonal ranges on mule deer genetic connectivity, and show that anthropogenic features hinder connectivity. This study demonstrates the value of combining a large, genome-wide marker set with recent advances in landscape genomics to evaluate functional connectivity in a wide-ranging migratory species.

Wyoming↗

Migration distance and maternal resource allocation determine timing of birth in a large herbivore

Birth timing is a key life-history characteristic that influences fitness and population performance. For migratory animals, however, appropriately timing birth on one seasonal range may be constrained by events occurring during other parts of the migratory cycle. We investigated how the use of capital and income resources may facilitate flexibility in reproductive phenology of migratory mule deer in western Wyoming, USA, over a 5-yr period (2015–2019). Specifically, we examined how seasonal interactions affected three interrelated life-history characteristics: fetal development, birth mass, and birth timing. Females in good nutritional condition at the onset of winter and those that migrated short distances had more developed fetuses (measured as fetal eye diameter in March). Variation in parturition date was explained largely by fetal development; however, there were up to 16 d of plasticity in expected birth date. Plasticity in expected birth date was shaped by income resources in the form of exposure to spring green-up. Although individuals that experienced greater exposure to spring green-up were able to advance expected birth date, being born early or late with respect to fetal development had no effect on birth mass of offspring. Furthermore, we investigated the trade-offs migrating mule deer face by evaluating support for existing theory that predicts that births should be matched to local peaks in resource availability at the birth site. In contrast to this prediction, only long-distance migrants that paced migration with the flush of spring green-up, giving birth shortly after ending migration, were able to match birth with spring green-up. Shorter-distance migrants completed migration sooner and gave birth earlier, seemingly trading off more time for offspring to grow and develop over greater access to resources. Thus, movement tactic had profound downstream effects on birth timing. These findings highlight a need to reconsider classical theory on optimal birth timing, which has focused solely on conditions at the birth site.

Wyoming↗

Sex-specific migratory behaviors in a temperate ungulate

Sexual segregation has been intensely studied across diverse ecosystems and taxa, but studies are often limited to periods when animals occupy distinct seasonal ranges. Some avian and marine studies have revealed that habitat segregation, when sexes differ spatially or temporally in use of the physical landscape, is common during the migratory period and characterized by sex-specific differences in migratory behaviors. Recent research highlights the importance of understanding movement patterns in the context of the full annual life cycle and highlights the need to extend relevant theories of sexual segregation to the migratory period. We tested predictions from two leading hypotheses of sexual segregation, the forage-selection hypothesis (FSH) and the reproductive strategy hypothesis (RSH) as applied to the migratory period. We collected global positioning system (GPS) location data for male and female mule deer ( Odocoileus hemionus ) in south-central Wyoming and northwest Colorado and tested the main predictions of the FSH and RSH. Both sexes showed high fidelity to their migratory routes, but route fidelity was more variable in males. Males also started spring migrations earlier, ended spring and autumn migrations later, and spent 22% more time on stopover sites during spring migrations. Consequently, males took twice as long in spring and 44% longer in autumn to complete migration. Our results revealed clear sex-specific migratory behaviors and supported predictions of the RSH that male foraging behaviors optimize body condition for the autumn rut, and females prioritize foraging while balancing reproductive constraints. Specifically, males timed their movements with spring green-up as optimally as females, and the timing of male migrations and use of stopovers suggested that males prioritized time in areas of high-quality forage. This refutes predictions of the FSH during the migratory period that males should consistently choose habitats with abundant, low-quality forage. Our findings provide an important contribution to sexual segregation theory by extending relevant theories to understand male and female movements during the migratory period.

Ecosphere↗

Life-history theory provides a framework for detecting resource limitation: A test of the Nutritional Buffer Hypothesis

For ungulates and other long-lived species, life-history theory predicts that nutritional reserves are allocated to reproduction in a state-dependent manner because survival is highly conserved. Further, as per capita food abundance and nutritional reserves decline (i.e., density dependence intensifies), reproduction and recruitment become increasingly sensitive to weather. Thus, the degree to which weather influences vital rates should be associated with proximity to nutritional carrying capacity—a notion that we refer to as the Nutritional Buffer Hypothesis. We tested the Nutritional Buffer Hypothesis using six moose ( Alces alces ) populations that varied in calf recruitment (33–69 calves/100 cows). We predicted that populations with high calf recruitment were nutritionally buffered against the effects of unfavorable weather, and thus were below nutritional carrying capacity. We applied a suite of tools to quantify habitat and nutritional condition of each population and found that increased browse condition, forage quality, and body fat were associated with increased pregnancy and calf recruitment, thereby providing multiple lines of evidence that declines in calf recruitment were underpinned by resource limitation. From 2001 to 2015, recruitment was more sensitive to interannual variation in weather (e.g., winter severity, drought) and plant phenology (e.g., duration of spring) for populations with reduced browse condition, forage quality, and body fat, suggesting these populations lacked the nutritional reserves necessary to buffer demographic performance against the effects of unfavorable weather. Further, average within-population calf recruitment was determined by regional climatic variation, suggesting that the pattern of reduced recruitment near the southern range boundary of moose stems from an interaction between climate and resource limitation. When coupled with information on habitat, nutrition, weather, and climate, life-history theory provides a framework to estimate nutritional limitation, proximity to nutritional carrying capacity, and impacts of climate change for ungulates.

Colorado, Utah, Wyoming↗

Drivers of site fidelity in ungulates

While the tendency to return to previously visited locations—termed ‘site fidelity’—is common in animals, the cause of this behaviour is not well understood. One hypothesis is that site fidelity is shaped by an animal's environment, such that animals living in landscapes with predictable resources have stronger site fidelity. Site fidelity may also be conditional on the success of animals’ recent visits to that location, and it may become stronger with age as the animal accumulates experience in their landscape. Finally, differences between species, such as the way memory shapes site attractiveness, may interact with environmental drivers to modulate the strength of site fidelity. We compared inter-year site fidelity in 669 individuals across eight ungulate species fitted with GPS collars and occupying a range of environmental conditions in North America and Africa. We used a distance-based index of site fidelity and tested hypothesized drivers of site fidelity using linear mixed effects models, while accounting for variation in annual range size. Mule deer Odocoileus hemionus and moose Alces alces exhibited relatively strong site fidelity, while wildebeest Connochaetes taurinus and barren-ground caribou Rangifer tarandus granti had relatively weak fidelity. Site fidelity was strongest in predictable landscapes where vegetative greening occurred at regular intervals over time (i.e. high temporal contingency). Species differed in their response to spatial heterogeneity in greenness (i.e. spatial constancy). Site fidelity varied seasonally in some species, but remained constant over time in others. Elk employed a ‘win-stay, lose-switch’ strategy, in which successful resource tracking in the springtime resulted in strong site fidelity the following spring. Site fidelity did not vary with age in any species tested. Our results provide support for the environmental hypothesis, particularly that regularity in vegetative phenology shapes the strength of site fidelity at the inter-annual scale. Large unexplained differences in site fidelity suggest that other factors, possibly species-specific differences in attraction to known sites, contribute to variation in the expression of this behaviour. Understanding drivers of variation in site fidelity across groups of organisms living in different environments provides important behavioural context for predicting how animals will respond to environmental change.

Journal of Animal Ecology↗