rKIN: Kernel-based method for estimating isotopic niche size and overlap.

The isotopic niche of consumers represents biologically relevant information on resource and habitat use. Several tools have been developed to quantify niche size and overlap. Nonetheless, methods adapted by spatial ecologists to quantify animal home ranges can be modified for use in stable isotope ecology when data are not normally distributed in bivariate space. We offer a tool that draws on existing spatial metrics, such as minimum convex polygon (MCP) and standard ellipse area (SEA), and add novel metrics using kernel utilization density (KUD) estimators to measure isotopic niche size and overlap. We present examples using empirical and simulated data to demonstrate the performance of the package kernel isotopic niches in r (rKIN) under various scenarios. Results of niche size from MCP, SEA and KUD were highly correlated but divergent among datasets. Overall, the KUD method produced the largest niche sizes and was more sensitive to the distribution of the isotopic data. Pairwise estimates of overlap were highly variable, likely because MCP and SEA inherently include or exclude unused areas in the resulting niche estimate. Four bandwidth methods (reference, normal scale, plug-in and biased cross-validation) produced comparable estimates of niche size and overlap at various sample sizes (10-40). Niche size and overlap were consistent across sample sizes >15. Use of rKIN will allow isotope ecologists to quantify niche shifts, expansions or contractions, as well as assess the performance of several estimation methods. The package also can be applied to other data types (e.g. principal component analysis, multi-dimensional scaling) so long as axes and measurement units are identical and can be converted to Cartesian coordinates.

Risk-sensitive allocation in seasonal dynamics of fat and protein reserves in a long-lived mammal.

Body reserves of numerous taxa follow seasonal rhythms that are a function of temporal patterns in food availability and life-history events; however, tests of the theory underlying the allocation of somatic reserves for long-lived organisms are rare, especially for free-ranging mammals. We evaluated the hypothesis that allocation of somatic reserves to survival (i.e., metabolic processes) and reproduction should be sensitive to current nutritional state relative to seasonal thresholds in those reserves. Our goal was to reveal the linkages between nutrition and life-history traits to understand how long-lived, iteroparous organisms balance the allocation of somatic reserves to reproduction, while retaining reserves as insurance for survival in unpredictable environments. Our evaluation was based on seasonal dynamics in fat (measured as ingesta-free body fat; IFBFat) and protein reserves (measured as ingesta-free, fat-free body mass; IFFFBMass) of 136 female mule deer (Odocoileus hemionus) over 8 years. Although mean changes in fat and protein reserves were positive over summer and negative over winter, accretion and catabolism of those reserves was not consistent among individuals. Over winter, both lipid and protein stores available in autumn were catabolized in proportion to their availability above a post-winter threshold (5·8% IFBFat, 33 kg IFFFBMass); however, lean body tissue was spared at the expense of lipid reserves. Female deer mostly synthesized lean body tissue over summer and committed post-winter fat reserves to reproduction relative to their availability above an autumn threshold (>8·6% IFBFat), which was lowered by 2·8 percentage points (pp) for each additional young recruited. Mothers reduced their autumn fat threshold to secure current reproductive investment and, thereby, endured a cost of reproduction at the expense of fat accumulation. Allocation of somatic reserves occurred in a risk-sensitive framework; females allocated reserves relative to their availability above seasonal thresholds. In contrast to current notions of summer accretion and winter catabolism of body reserves, some individuals deposited reserves over winter and catabolized reserves over summer, mainly because regulation of individual condition was state-dependent. Consequently, behaviour and life-history strategies may be as much a function of nutritional contributions of the previous season as of the current one.

Evolution of ungulate mating systems: Integrating social and environmental factors.

Ungulates exhibit diverse mating systems that range from monogamous pair territories to highly polygynous leks. We review mating systems and behaviors across ungulates and offer a new approach synthesizing how interacting factors may shape those mating systems. Variability exists in mating systems among and within species of ungulates and likely is affected by predation risk, availability of resources (food and mates), habitat structure, and sociality. Ungulate mating systems may be labile as a consequence of the varying strength of those interacting factors. In addition, degree of polygyny and sexual dimorphism in size are associated with the evolution of mating systems. Neither male-male combat nor paternal care, however, can completely explain differences in sexual size dimorphism for ungulates, a necessary component in understanding the development of some mating systems. Whatever the evolutionary pathway, sexual segregation limits paternal care allowing more intense male-male competition. Selection of habitat structure, because it modifies risk of predation, is a major determinant of sociality for ungulates. Likewise, ruggedness and steepness of terrain limit the types of mating systems that can occur because of limitations in group size and cohesiveness, as well as the ability of males to herd even small groups of females effectively. The quality and defensibility of resources affect mating systems, as does the defensibility of females. Population density of females also may be a critical determinant of the types of mating systems that develop. Size of groups likewise constrains the types of mating tactics that males can employ. Our aim was to use those relationships to create a broad conceptual model that predicts how various environmental and social factors interact to structure mating systems in ungulates. This model provides a useful framework for future tests of the roles of both ecological and social conditions in influencing the social systems of ungulates.

Population density of North American elk: effects on plant diversity.

Large, herbivorous mammals have profound effects on ecosystem structure and function and often act as keystone species in ecosystems they inhabit. Density-dependent processes associated with population structure of large mammals may interact with ecosystem functioning to increase or decrease biodiversity, depending on the relationship of herbivore populations relative to the carrying capacity (K) of the ecosystem. We tested for indirect effects of population density of large herbivores on plant species richness and diversity in a montane ecosystem, where increased net aboveground primary productivity (NAPP) in response to low levels of herbivory has been reported. We documented a positive, linear relationship between plant-species diversity and richness with NAPP. Structural equation modeling revealed significant indirect relationships between population density of herbivores, NAPP, and species diversity. We observed an indirect effect of density-dependent processes in large, herbivorous mammals and species diversity of plants through changes in NAPP in this montane ecosystem. Changes in species diversity of plants in response to herbivory may be more indirect in ecosystems with long histories of herbivory. Those subtle or indirect effects of herbivory may have strong effects on ecosystem functioning, but may be overlooked in plant communities that are relatively resilient to herbivory.

Population structure and genetic diversity of moose in Alaska.

Moose (Alces alces) are highly mobile mammals that occur across arboreal regions of North America, Europe, and Asia. Alaskan moose (Alces alces gigas) range across much of Alaska and are primary herbivore consumers, exerting a prominent influence on ecosystem structure and functioning. Increased knowledge gained from population genetics provides insights into their population dynamics, history, and dispersal of these unique large herbivores and can aid in conservation efforts. We examined the genetic diversity and population structure of moose (n = 141) with 8 polymorphic microsatellites from 6 regions spanning much of Alaska. Expected heterozygosity was moderate (H(E) = 0.483-0.612), and private alleles ranged from 0 to 6. Both F(ST) and R(ST) indicated significant population structure (P < 0.001) with F(ST) < 0.109 and R(ST) < 0.125. Results of analyses from STRUCTURE indicated 2 prominent population groups, a mix of moose from the Yakutat and Tetlin regions versus all other moose, with slight substructure observed among the second population. Estimates of dispersal differed between analytical approaches, indicating a high level of historical or current gene flow. Mantel tests indicated that isolation-by-distance partially explained observed structure among moose populations (R(2) = 0.45, P < 0.01). Finally, there was no evidence of bottlenecks either at the population level or overall. We conclude that weak population structure occurs among moose in Alaska with population expansion from interior Alaska westward toward the coast.

Habitat use and social mixing between groups of resident and augmented bighorn sheep.

Monitoring dispersal, habitat use, and social mixing of released ungulates is crucial for successful translocation and species conservation. We monitored 127 female bighorn sheep (Ovis canadensis) released in three populations from 2000 to 2009 to investigate if augmented bighorns expanded and shifted seasonal ranges, used different habitat compared with resident females, and if animals mixed socially. Augmented bighorns in all populations expanded range use compared with residents by shifting utilization distributions. Size of utilization distributions, however, were smaller for augmented females compared with residents in all areas except one. Overlap of seasonal utilization distributions between augmented and resident bighorns and use of slope and elevation differed across populations. In two populations, differences in size and overlap of seasonal utilization distributions and use of slope and elevation supported the hypothesis that habitat use of bighorns in their source area influenced their habitat use after release. Mixing between resident and augmented adult females occurred on average during only 21% of sightings and was similar across populations. Our results clarify how augmented bighorns mix with resident animals and how habitat use is modified following augmentations. Such information is needed to improve bighorn sheep augmentations and can be applied to augmentations of other ungulates.

Hunting and mountain sheep: Do current harvest practices affect horn growth?

The influence of human harvest on evolution of secondary sexual characteristics has implications for sustainable management of wildlife populations. The phenotypic consequences of selectively removing males with large horns or antlers from ungulate populations have been a topic of heightened concern in recent years. Harvest can affect size of horn-like structures in two ways: (a) shifting age structure toward younger age classes, which can reduce the mean size of horn-like structures, or (b) selecting against genes that produce large, fast-growing males. We evaluated effects of age, climatic and forage conditions, and metrics of harvest on horn size and growth of mountain sheep (Ovis canadensis ssp.) in 72 hunt areas across North America from 1981 to 2016. In 50% of hunt areas, changes in mean horn size during the study period were related to changes in age structure of harvested sheep. Environmental conditions explained directional changes in horn growth in 28% of hunt areas, 7% of which did not exhibit change before accounting for effects of the environment. After accounting for age and environment, horn size of mountain sheep was stable or increasing in the majority (~78%) of hunt areas. Age-specific horn size declined in 44% of hunt areas where harvest was regulated solely by morphological criteria, which supports the notion that harvest practices that are simultaneously selective and intensive might lead to changes in horn growth. Nevertheless, phenotypic consequences are not a foregone conclusion in the face of selective harvest; over half of the hunt areas with highly selective and intensive harvest did not exhibit age-specific declines in horn size. Our results demonstrate that while harvest regimes are an important consideration, horn growth of harvested male mountain sheep has remained largely stable, indicating that changes in horn growth patterns are an unlikely consequence of harvest across most of North America.

Sexual segregation in North American elk: the role of density dependence.

We investigated how density-dependent processes and subsequent variation in nutritional condition of individuals influenced both timing and duration of sexual segregation and selection of resources. During 1999-2001, we experimentally created two population densities of North American elk (Cervus elaphus), a high-density population at 20 elk/km(2), and a low-density population at 4 elk/km(2) to test hypotheses relative to timing and duration of sexual segregation and variation in selection of resources. We used multi-response permutation procedures to investigate patterns of sexual segregation, and resource selection functions to document differences in selection of resources by individuals in high- and low-density populations during sexual segregation and aggregation. The duration of sexual segregation was 2 months longer in the high-density population and likely was influenced by individuals in poorer nutritional condition, which corresponded with later conception and parturition, than at low density. Males and females in the high-density population overlapped in selection of resources to a greater extent than in the low-density population, probably resulting from density-dependent effects of increased intraspecific competition and lower availability of resources.

Moose herbivory, browse quality, and nutrient cycling in an Alaskan treeline community.

Moose (Alces alces) browsing on diamondleaf willow (Salix planifolia pulchra) caused significant increases in subsequent growth of stems and leaves in treeline plant communities in central Alaska, USA. Willows growing in the shade were significantly more palatable for moose than those growing in the sun. Moose density had strong effects on rates of nutrient cycling, ostensibly through effects of browsing and inputs from fecal and urinary nitrogen. Moose are a keystone herbivore that likely mediate rates of nutrient cycling in northern ecosystems.

Nutritional development of feeding strategies in arctic ruminants: digestive morphometry of reindeer, Rangifer tarandus, and muskoxen, Ovibos moschatus.

Reindeer have been classified as intermediate feeders and muskoxen as grazers based on differences in digestive morphology and consumption of fibrous plants. We hypothesized that the digestive morphology of young (< 2 months) reindeer and muskoxen anticipates transitions in diet and determines the feeding strategy of each species at adulthood. We compared structural morphology and rates of cell division in the rumen, abomasum, duodenum and liver of reindeer and muskoxen as neonates (1 day old), during the transition from milk to forage (30-60 days old) and in adults (> 7 yr). Development in utero provided the neonate with a functioning mucosa of the gastric abomasum and duodenal mucosa with high surface enlargement for digestion and absorption of concentrated milks. Transition to forage was preceded by changes in ruminal papillae structure that increased surface area and likely contributed to active fermentation by 60 days of age. The abomasum also increased in acid-secreting parietal cells during the transition to forage, which may enhance digestion of plant and microbial proteins. Rates of cell division also indicated a sustained differentiation of tissue structure during the transitional period. Young arctic ruminants expressed digestive structures that preceded full function, which indicated the ultimate feeding strategy of each species. For example, the rumen of young muskoxen had thick cornified epithelia and muscle layers that would provide ruminal mucosa with better protection from fibrous abrasion and enhance motility of bulky diets. Conversely, young reindeer had more complex papillary shapes in the rumen and more foliate villi in the duodenum, indicating a greater absorptive capacity of these structures than in muskoxen. Ontogenetic programs, therefore, play the primary role for digestive development of reindeer and muskoxen and determine the nutritional strategies of adults.

Migrating mule deer: effects of anthropogenically altered landscapes.

BACKGROUND: Migration is an adaptive strategy that enables animals to enhance resource availability and reduce risk of predation at a broad geographic scale. Ungulate migrations generally occur along traditional routes, many of which have been disrupted by anthropogenic disturbances. Spring migration in ungulates is of particular importance for conservation planning, because it is closely coupled with timing of parturition. The degree to which oil and gas development affects migratory patterns, and whether ungulate migration is sufficiently plastic to compensate for such changes, warrants additional study to better understand this critical conservation issue. METHODOLOGY/PRINCIPAL FINDINGS: We studied timing and synchrony of departure from winter range and arrival to summer range of female mule deer (Odocoileus hemionus) in northwestern Colorado, USA, which has one of the largest natural-gas reserves currently under development in North America. We hypothesized that in addition to local weather, plant phenology, and individual life-history characteristics, patterns of spring migration would be modified by disturbances associated with natural-gas extraction. We captured 205 adult female mule deer, equipped them with GPS collars, and observed patterns of spring migration during 2008-2010. CONCLUSIONS/SIGNIFICANCE: Timing of spring migration was related to winter weather (particularly snow depth) and access to emerging vegetation, which varied among years, but was highly synchronous across study areas within years. Additionally, timing of migration was influenced by the collective effects of anthropogenic disturbance, rate of travel, distance traveled, and body condition of adult females. Rates of travel were more rapid over shorter migration distances in areas of high natural-gas development resulting in the delayed departure, but early arrival for females migrating in areas with high development compared with less-developed areas. Such shifts in behavior could have consequences for timing of arrival on birthing areas, especially where mule deer migrate over longer distances or for greater durations.

Density-dependent effects on physical condition and reproduction in North American elk: an experimental test.

Density dependence plays a key role in life-history characteristics and population ecology of large, herbivorous mammals. We designed a manipulative experiment to test hypotheses relating effects of density-dependent mechanisms on physical condition and fecundity of North American elk (Cervus elaphus) by creating populations at low and high density. We hypothesized that if density-dependent effects were manifested principally through intraspecific competition, body condition and fecundity of females would be lower in an area of high population density than in a low-density area. Thus, we collected data on physical condition and rates of pregnancy in each experimental population. Our manipulative experiment indicated that density-dependent feedbacks affected physical condition and reproduction of adult female elk. Age-specific pregnancy rates were lower in the high-density area, although there were no differences in pregnancy of yearlings or in age at peak reproduction between areas. Age-specific rates of pregnancy began to diverge at 2 years of age between the two populations and peaked at 6 years old. Pregnancy rates were most affected by body condition and mass, although successful reproduction the previous year also reduced pregnancy rates during the current year. Our results indicated that while holding effects of winter constant, density-dependent mechanisms had a much greater effect on physical condition and fecundity than density-independent factors (e.g., precipitation and temperature). Moreover, our results demonstrated effects of differing nutrition resulting from population density during summer on body condition and reproduction. Thus, summer is a critical period for accumulation of body stores to buffer animals against winter; more emphasis should be placed on the role of spring and summer nutrition on population regulation in large, northern herbivores.

Mitochondrial phylogeography of moose (Alces alces): late pleistocene divergence and population expansion.

We examined phylogeographic relationships of moose (Alces alces) worldwide to test the proposed existence of two geographic races and to infer the timing and extent of demographic processes underpinning the expansion of this species across the Northern Hemisphere in the late Pleistocene. Sequence variation within the left hypervariable domain of the control region occurred at low or moderate levels worldwide and was structured geographically. Partitioning of genetic variance among regions indicated that isolation by distance was the primary agent for differentiation of moose populations but does not support the existence of distinct eastern and western races. Levels of genetic variation and structure of phylogenetic trees identify Asia as the origin of all extant mitochondrial lineages. A recent coalescence is indicated, with the most recent common ancestor dating to the last ice age. Moose have undergone two episodes of population expansion, likely corresponding to the final interstade of the most recent ice age and the onset of the current interglacial. Timing of expansion for the population in the Yakutia--Manchuria region of eastern Asia indicates that it is one of the oldest populations of moose and may represent the source of founders of extant populations in North America, which were colonized within the last 15,000 years. Our data suggest an extended period of low population size or a severe bottleneck prior to the divergence and expansion of extant lineages and a recent, less-severe bottleneck among European lineages. Climate change during the last ice age, acting through contraction and expansion of moose habitat and the flooding of the Bering land bridge, undoubtedly was a key factor influencing the divergence and expansion of moose populations.