Geology ReportsSearch

Geology topics

Lisette P. Waits

Publications and source records attributed to Lisette P. Waits.

At least 19 recordsLinked to original sources

Genetic structure in a previously extirpated population of gray wolves following reintroduction and natural recolonization

Genetic structuring in wildlife populations is driven by barriers that restrict gene flow as well as the history of population demography. Mechanisms driving genetic structuring can be nuanced in group-living species, such as gray wolves ( Canis lupus ). Behavioral factors, such as social affiliation and resistance, natal habitat imprinting, and trade-offs between dispersal from natal packs and territorial biding, affect habitat selection of wolves despite landscape barriers providing little resistance to their extensive dispersal capabilities. Wolves were previously extirpated from Idaho, USA, and current populations are the result of both reintroductions in 1995 and 1996 and natural dispersal from Canada. In this context we examined genetic structure of wolves in Idaho using 101 individuals genotyped at 18 nuclear DNA microsatellite loci and a subset of 38 individuals genotyped at 1019 single nucleotide polymorphism markers. We hypothesized panmictic (i.e., random mating) genetic structure in Idaho due to the long-distance dispersal abilities of gray wolves. Contrary to our hypothesis, we found three genetic clusters of gray wolves in Idaho, primarily supported by SNP markers. Microsatellite data suggested similar patterns, but permutation tests indicated these differences were not statistically significant. The extent of differentiation and evidence of gene flow, however, suggests that the three genetic clusters are not wholly isolated from one another. The distinctions between clusters spatially align with areas of reintroduction into central Idaho and Yellowstone National Park, as well ongoing natural recolonization from adjacent populations in Canada and Montana. Wolves at the periphery of analysis areas showed more admixture than those in the core, consistent with territoriality and mating behaviors contributing to genetic structuring. We demonstrate how management history, including reintroduction efforts, and animal behavior may interact and contribute to patterns of genetic structure in wild populations.

Idaho, Montana, Wyoming

Landscape associations and population genetics of a generalist carnivore at a range limit

American black bear ( Ursus americanus ) sightings have increased in the Oklahoma Panhandle, an area outside of the species’ historical range, prompting an assessment of bears in the region. We used camera traps and an occupancy modeling framework to identify factors influencing bear detection and space-use patterns. We used noninvasive genetic sampling techniques to evaluate genetic diversity, population structure, and bear abundance in the region. During the summers of 2022–2023, we deployed cameras at 160 sites across western Oklahoma (USA) and detected ≥1 bear at 20 sites. The most-supported model from our single-season single-species analysis indicated that bear detection was positively associated with temperature and precipitation, negatively associated with day of year, and differed between years. The most-supported model indicated that bear space use was negatively associated with elevation (β = −0.013, 85% CI = −0.025, 0.000), and positively associated with slope (β = 0.645, 85% CI = 0.305, 0.984) and coarse woody debris counts (β = 1.539, 85% CI = 0.314, 2.765). We deployed 41 hair snares in Oklahoma resulting in the collection of 153 hair samples and received 69 tissue samples from black bears harvested in northeastern New Mexico. Using 11 microsatellite markers, we identified 21 (12M:9F) bears in western Oklahoma, and 69 (40M:29F) in New Mexico. We found evidence that bears occurring in Oklahoma were an extension of a previously documented population that occurred in northcentral New Mexico. We detected significant population-level heterozygote deficiency ( P = 0.013) compared to expectations under Hardy-Weinberg equilibrium. Using capture with replacement models, we estimated 26 (95% CI = 19–43) bears in western Oklahoma during 2022–2023. Our results provide baseline data on population distribution, abundance, and genetic health of bears in the region and identify factors that may drive human-bear conflicts as the bear population increases in western Oklahoma.

New Mexico, Oklahoma

Genetic variation and metapopulation structure inform recovery goals in a threatened species

Background: Monitoring genetic parameters is important for setting effective conservation and management strategies, particularly for small, fragmented, and isolated populations. Small, isolated populations face increased rates of genetic drift and inbreeding, which increase extinction risk especially when gene flow is limited. Methods: Here, we applied a Genotyping-in-Thousands by sequencing (GT-seq) panel to inform recovery action for the federally threatened northern Idaho ground squirrel ( Urocitellus brunneus ). We evaluated genetic diversity, structure, connectivity, and effective population size to address species recovery goals. Results: We delineated three types of conservation units: (1) three evolutionarily significant units that represent long-term population structure and variation, (2) nine management units that reflect current demographic connectivity and restrictions to gene flow, and (3) three adaptive units that capture adaptive differentiation across the species range. Effective population sizes per management unit were small overall (mean 38.16, range 2.3–220.9), indicating that recovery goals of 10 subpopulations with N e > 500 have not been reached. Conclusions: Our results support the maintenance of connectivity within evolutionarily significant units through the restoration of dispersal corridors. Next steps could include further sampling of some subpopulations with low sample sizes, unsampled subpopulations, and subpopulations that are geographically isolated. Genotyping future samples with the same GT-seq panel would help to detect dispersal, assess effective population size, monitor the effects of inbreeding, and evaluate adaptive differentiation to monitor the effects of management action and environmental change.

Idaho

Genetic connectivity in a cooperatively breeding carnivore between two protected areas

Wildlife populations are increasingly threatened by human activities. Most studies, however, are often short in duration or do not encompass the large spatial extent necessary to measure the potential effects of human activities on population vital rates. Furthermore, the life history features of species with high fecundity and excellent dispersal capabilities can act as buffers against the potential negative effects of human activities on their populations. We used a 30-year dataset of genetic samples from gray wolves ( Canis lupus ) in Alaska, USA, to examine genetic connectivity and diversity between National Park units separated by a region with recurrent human-caused mortality. We found that the two protected populations were genetically similar and that dispersal events occurred between them even though they are > 450 km apart. We posit that intact ecosystems and a history of continuous distribution of wolves surrounding the affected regions likely maintained the genetic connectivity of wolves in the two protected areas.

Alaska

Linking age and social status of cooperative breeders to vulnerability throughout the harvest season

Individual behaviors are influenced by environmental, genetic, and demographic factors. Some animals choose to live in groups and cooperatively breed, and their behaviors can change depending on dynamic factors such as group size and composition that affect group persistence. In Idaho, USA, gray wolves ( Canis lupus ) are harvested annually, providing an opportunity to investigate the effects of harvest and seasonal behaviors on a population of cooperative breeders. These annual hunting and trapping seasons overlap with the dispersal and breeding periods for wolves and we do not know how harvest affects the vulnerability of different sex and age classes during these important biological periods. We applied 9 years (2009–2018) of genetic, age, and harvest data from harvested wolves to investigate how behaviors (dispersal and breeding) and biological drivers might influence the vulnerability of wolves to harvest. We created pedigrees from genotypes of non-invasively collected scats to estimate the expected proportion of the wolf population composed of 3 different age classes (pup, yearling, and sexually mature or ≥2 years old) and compared them to the observed number of each age class harvested during biologically significant periods (i.e., dispersal and breeding). We found that pups were more vulnerable to harvest in December when wolf harvest transitioned largely to trapping (accounts for 66% of harvest), and found evidence that adults were more vulnerable to harvest during their breeding season in January and February. In contrast, we found no difference in the expected versus observed number of wolves ≥2 years old in the harvest during peak dispersal season (December), or in the expected versus observed number of yearlings in the harvest during September and October when pups are mobile and groups of wolves abandon the use of pup-rearing sites. Some age classes were disproportionally harvested during certain periods for specific years, but this was not consistent across all years, suggesting there is more to learn about the vulnerability of different age classes to harvest. We found harvest can disproportionally affect some demographic classes of individuals depending on year, biological period, and harvest type. With wolves continuing to recolonize historical ranges, our approach can benefit managers and future studies with the goal of identifying how interannual harvest affects groups of wolves.

Idaho

Spatial close-kin mark-recapture models applied to terrestrial species with continuous natal dispersal

Close-kin mark–recapture (CKMR) methods use information on genetic relatedness among individuals to estimate demographic parameters. An individual's genotype can be considered a ‘recapture’ of each of its parent's genotype, and the frequency of kin-pair matches detected in a population sample can directly inform estimates of abundance. CKMR inference procedures require analysts to define kinship probabilities in functional forms, which inevitably involve simplifying assumptions. Among others, population structure can have a strong influence on how kinship probabilities are formulated. Many terrestrial species are philopatric or face barriers to dispersal, and not accounting for dispersal limitation in kinship probabilities, can create substantial bias if sampling is also spatially structured (e.g. via harvest). We present a spatially explicit formulation of CKMR that corrects for incomplete mixing by incorporating natal dispersal distances and spatial distribution of individuals into the kinship probabilities. We used individual-based simulations to evaluate the accuracy of abundance estimates obtained with one spatially naïve and two spatially explicit CKMR models across six scenarios with distinct spatial patterns of relative abundance and sampling probability. Estimates of abundance obtained with a CKMR model naïve to spatial structure were negatively biased when sampling was spatially biased. Incorporating patterns of natal dispersal in the kinship probabilities helped address this bias, but estimates were not always accurate depending on the model used and the scenario considered. Incorporating natal dispersal into spatially structured CKMR models can address the bias created by population structure and heterogeneous sampling but will often require additional assumptions and auxiliary data (e.g. relative abundance indices). The models shown here were designed for terrestrial species with continuous patterns of natal dispersal and high year-to-year site fidelity but could be extended to other species.

Methods in Ecology and Evolution

Gene flow prevents genetic diversity loss despite small effective population size in fragmented grizzly bear (Ursus arctos) populations

Genetic monitoring is important in small, fragmented populations that rely on gene flow to maintain genetic diversity. The Selkirk, Yaak, and Cabinet grizzly bear ( Ursus arctos ) populations are among the smallest in North America and are near the southernmost extent of the species’ range. These populations received little to no effective migration for generations but have recently experienced increased gene flow through natural migration and a population augmentation program. A long-term dataset of grizzly bear microsatellite genotypes from 1973 to 2021 presented a unique opportunity to examine genetic trends in these populations over time. We used this dataset of 464 bears to evaluate if gene flow affected observed heterozygosity ( H O ), expected heterozygosity ( H E ), allelic richness ( A R ), and average pairwise relatedness ( r ) in each of these populations. We also estimated effective population size ( N e ) using the temporal and linkage disequilibrium (LD) methods. Post gene flow, A R increased in the Selkirk and Cabinet populations and r decreased in all three populations. We did not observe any significant changes in H E or H O , but H E values in our populations were significantly higher than those estimated using a model without gene flow. Our N e estimates were consistent between the temporal and LD methods and ranged from 15.2 to 15.8, 15.4–17.5, and 5.6–8.9 for the Selkirk, Yaak, and Cabinet populations, respectively. Overall, our findings indicate that gene flow is increasing or maintaining genetic diversity in these populations. However, N e remains low and additional connectivity or augmentation may be needed, particularly in the Cabinet population.

Idaho, Montana, Washington

Gray wolf breeders are more vulnerable to harvest during the breeding season

In cooperatively breeding carnivores, breeders are vital to perpetuating the group; the death or removal of an individual breeder can greatly affect group composition, genetic content, and short-term population growth. Understanding the number of breeders harvested and timing of harvest can increase our knowledge of how mortality affects groups of cooperative breeders. Gray wolves ( Canis lupus ) in Idaho, USA, are exposed to annual harvest and are an ideal species for studying the effects of harvest on breeder turnover. We combined genotypes from tissue samples of harvested wolves with parentage analyses and cementum annuli ages and estimated when and how many breeding wolves were harvested. We genotyped and aged 229 adults and 203 pups using tissue and tooth samples from wolves harvested between 2014 and 2016. We identified a minimum count of 33 breeders in the harvest and found that they were disproportionately harvested more during the breeding season. We estimated that a minimum of ~14.5% of adult wolves harvested annually, or approximately 1 in 7, were breeders. We posit their behavior during breeding season may increase their vulnerability to harvest. By linking animal life history with vulnerability to human-caused mortality we show that managers could structure harvest seasons so there is less overlap with wolves’ breeding season if there is concern about the demographic consequences of harvesting breeders.

Idaho

Predator-specific mortality of sage-grouse nests based on predator DNA on eggshells

Greater sage-grouse (hereafter sage-grouse; Centrocercus urophasianus ) populations have declined across their range. Increased nest predation as a result of anthropogenic land use is one mechanism proposed to explain these declines. However, sage-grouse contend with a diverse suite of nest predators that vary in functional traits (e.g., search tactics or hunting mode) and abundance. Consequently, generalizing about factors influencing nest fate is challenging. Identifying the explicit predator species responsible for nest predation events is, therefore, critical to understanding causal mechanisms linking land use to patterns of sage-grouse nest success. Cattle grazing is often assumed to adversely affect sage-grouse recruitment by reducing grass height (and hence cover), thereby facilitating nest detection by predators. However, recent evidence found little support for the hypothesized effect of grazing on nest fate at the pasture scale. Rather, nest success appears to be similar on pastures grazed at varying intensities. One possible explanation for the lack of observed effect involves a localized response by one or more nest predators. The presence of cattle may cause a temporary reduction in predator density and/or use within a pasture (the cattle avoidance hypothesis). The cattle avoidance hypothesis predicts a decreased probability of at least one sage-grouse nest predator predating sage-grouse nests in pastures with livestock relative to pastures without livestock present during the nesting season. To test the cattle avoidance hypothesis, we collected predator DNA from eggshells from predated nests and used genetic methods to identify the sage-grouse nest predator(s) responsible for the predation event. We evaluated the influence of habitat and grazing on predator-specific nest predation. We evaluated the efficacy of our genetic method by deploying artificial nests with trail cameras and compared the results of our genetic method to the species captured via trail camera. Our molecular methods identified at least one nest predator captured predating artificial nests via trail camera for 33 of 35 (94%) artificial nests. We detected nest predators via our molecular analysis at 76 of 114 (67%) predated sage-grouse nests. The primary predators detected at sage-grouse nests were coyotes ( Canis latrans ) and corvids ( Corvidea ). Grazing did not influence the probability of nest predation by either coyotes or corvids. Sagebrush canopy cover was negatively associated with the probability a coyote predated a nest, distance to water was positively associated with the probability a corvid predated a nest, and average minimum temperature was negatively associated with the probability that either a coyote or a corvid predated a nest. Our study provides a framework for implementing an effective, non-invasive method for identifying sage-grouse nest predators that can be used to better understand how management actions at local and regional scales may impact an important component of sage-grouse recruitment.

Idaho

Genetic analysis of harvest samples reveals population structure in a highly mobile generalist carnivore

Delineating wildlife population boundaries is important for effective population monitoring and management. The bobcat ( Lynx rufus ) is a highly mobile generalist carnivore that is ecologically and economically important. We sampled 1225 bobcats harvested in South Dakota, USA (2014–2019), of which 878 were retained to assess genetic diversity and infer population genetic structure using 17 microsatellite loci. We assigned individuals to genetic clusters ( K ) using spatial and nonspatial Bayesian clustering algorithms and quantified differentiation ( F ST and GST″ ) among clusters. We found support for population genetic structure at K = 2 and K = 4, with pairwise F ST and GST″ values indicating weak to moderate differentiation, respectively, among clusters. For K = 2, eastern and western clusters aligned closely with historical bobcat management units and were consistent with a longitudinal suture zone for bobcats previously identified in the Great Plains. We did not observe patterns of population genetic structure aligning with major rivers or highways. Genetic divergence observed at K = 4 aligned roughly with ecoregion breaks and may be associated with environmental gradients, but additional sampling with more precise locational data may be necessary to validate these patterns. Our findings reveal that cryptic population structure may occur in highly mobile and broadly distributed generalist carnivores, highlighting the importance of considering population structure when establishing population monitoring programs or harvest regulations. Our study further demonstrates that for elusive furbearers, harvest can provide an efficient, broad-scale sampling approach for genetic population assessments.

South Dakota

Development and validation of a GT-seq panel for genetic monitoring in a threatened species using minimally invasive sampling

Minimally invasive samples are often the best option for collecting genetic material from species of conservation concern, but they perform poorly in many genomic sequencing methods due to their tendency to yield low DNA quality and quantity. Genotyping-in-thousands by sequencing (GT-seq) is a powerful amplicon sequencing method that can genotype large numbers of variable-quality samples at a standardized set of single nucleotide polymorphism (SNP) loci. Here, we develop, optimize, and validate a GT-seq panel for the federally threatened northern Idaho ground squirrel ( Urocitellus brunneus ) to provide a standardized approach for future genetic monitoring and assessment of recovery goals using minimally invasive samples. The optimized panel consists of 224 neutral and 81 putatively adaptive SNPs. DNA collected from buccal swabs from 2016 to 2020 had 73% genotyping success, while samples collected from hair from 2002 to 2006 had little to no DNA remaining and did not genotype successfully. We evaluated our GT-seq panel by measuring genotype discordance rates compared to RADseq and whole-genome sequencing. GT-seq and other sequencing methods had similar population diversity and F ST estimates, but GT-seq consistently called more heterozygotes than expected, resulting in negative F IS values at the population level. Genetic ancestry assignment was consistent when estimated with different sequencing methods and numbers of loci. Our GT-seq panel is an effective and efficient genotyping tool that will aid in the monitoring and recovery of this threatened species, and our results provide insights for applying GT-seq for minimally invasive DNA sampling techniques in other rare animals.

Ecology and Evolution

The influence of grazing on the spatiotemporal activity patterns of a primary sage-grouse nest predator

Perturbations in ecological processes can occur when wildlife alter their spatiotemporal activity patterns to avoid human activities that they perceive as a risk. Such perturbations can have cascading effects throughout wildlife communities. For greater sage-grouse ( Centrocercus urophasianus ; hereafter sage-grouse), nest predation plays an important role in population dynamics. Domestic cattle ( Bos taurus ) grazing has been hypothesized to increase nest predation by reducing grass height, and therefore reducing nest concealment, which may facilitate nest detection by predators. Grass height is lower on grazed pastures, but sage-grouse nest success appears similar on pastures grazed at varying intensities in several recent studies. Any reductions in nest concealment caused by grazing could potentially be offset by a localized response of one or more nest predators to the presence of cattle (i.e., the cattle avoidance hypothesis). A reduction in nest predator density or relative use within pastures could explain similar patterns of nest success on pastures grazed at varying intensities. Also, wildlife can potentially partition themselves temporally to avoid risks associated with human activities. For example, a shift in diel activity patterns by nest predators in response to cattle could result in predators being active during portions of the day when they are less efficient at locating sage-grouse nests. Thus, the effects of grazing could be offset by a temporal avoidance of cattle by predators. We deployed motion sensor cameras across six pastures to evaluate whether coyotes ( Canis latrans ; a primary sage-grouse nest predator) altered spatiotemporal activity patterns in response to cattle. We found that the probability of detecting coyotes had a positive relationship with cattle detections at camera sites (β = 0.22; 95% CI = 0.14,0.30). We also found that coyotes did not shift their diel activity patterns in response to cattle being in the pastures. Thus, in our system, similar sage-grouse nest success among pastures with different grazing intensities cannot be explained by the cattle avoidance hypothesis, at least for coyotes.

Idaho

The role of neutral and adaptive genomic variation in population diversification and speciation in two ground squirrel species of conservation concern

Understanding the neutral (demographic) and adaptive processes leading to the differentiation of species and populations is a critical component of evolutionary and conservation biology. In this context, recently diverged taxa represent a unique opportunity to study the process of genetic differentiation. Northern and southern Idaho ground squirrels ( Urocitellus brunneus —NIDGS, and U . endemicus —SIDGS, respectively) are a recently diverged pair of sister species that have undergone dramatic declines in the last 50 years and are currently found in metapopulations across restricted spatial areas with distinct environmental pressures. Here we genotyped single-nucleotide polymorphisms (SNPs) from buccal swabs with restriction site-associated DNA sequencing (RADseq). With these data we evaluated neutral genetic structure at both the inter- and intraspecific level, and identified putatively adaptive SNPs using population structure outlier detection and genotype–environment association (GEA) analyses. At the interspecific level, we detected a clear separation between NIDGS and SIDGS, and evidence for adaptive differentiation putatively linked to torpor patterns. At the intraspecific level, we found evidence of both neutral and adaptive differentiation. For NIDGS, elevation appears to be the main driver of adaptive differentiation, while neutral variation patterns match and expand information on the low connectivity between some populations identified in previous studies using microsatellite markers. For SIDGS, neutral substructure generally reflected natural geographical barriers, while adaptive variation reflected differences in land cover and temperature, as well as elevation. These results clearly highlight the roles of neutral and adaptive processes for understanding the complexity of the processes leading to species and population differentiation, which can have important conservation implications in susceptible and threatened species.

Idaho

Genomic association with pathogen carriage in bighorn sheep (Ovis canadensis)

Genetic composition can influence host susceptibility to, and transmission of, pathogens, with potential population‐level consequences. In bighorn sheep ( Ovis canadensis ), pneumonia epidemics caused by Mycoplasma ovipneumoniae have been associated with severe population declines and limited recovery across North America. Adult survivors either clear the infection or act as carriers that continually shed M. ovipneumoniae and expose their susceptible offspring, resulting in high rates of lamb mortality for years following the outbreak event. Here, we investigated the influence of genomic composition on persistent carriage of M. ovipneumoniae in a well‐studied bighorn sheep herd in the Wallowa Mountains of Oregon, USA. Using 10,605 SNPs generated using RADseq technology for 25 female bighorn sheep, we assessed genomic diversity metrics and employed family‐based genome‐wide association methodologies to understand variant association and genetic architecture underlying chronic carriage. We observed no differences among genome‐wide diversity metrics (heterozygosity and allelic richness) between groups. However, we identified two variant loci of interest and seven associated candidate genes, which may influence carriage status. Further, we found that the SNP panel explained ~55% of the phenotypic variance (SNP‐based heritability) for M. ovipneumoniae carriage, though there was considerable uncertainty in these estimates. While small sample sizes limit conclusions drawn here, our study represents one of the first to assess the genomic factors influencing chronic carriage of a pathogen in a wild population and lays a foundation for understanding genomic influence on pathogen persistence in bighorn sheep and other wildlife populations. Future research should incorporate additional individuals as well as distinct herds to further explore the genomic basis of chronic carriage.

Oregon

Diet of a rare herbivore based on DNA metabarcoding of feces: Selection, seasonality, and survival

In herbivores, survival and reproduction are influenced by quality and quantity of forage, and hence, diet and foraging behavior are the foundation of an herbivore's life history strategy. Given the importance of diet to most herbivores, it is imperative that we know the species of plants they prefer, especially for herbivorous species that are at risk for extinction. However, it is often difficult to identify the diet of small herbivores because: (a) They are difficult to observe, (b) collecting stomach contents requires sacrificing animals, and (c) microhistology requires accurately identifying taxa from partially digested plant fragments and likely overemphasizes less-digestible taxa. The northern Idaho ground squirrel ( Urocitellus brunneus ) is federally threatened in the United States under the Endangered Species Act. We used DNA metabarcoding techniques to identify the diet of 188 squirrels at 11 study sites from fecal samples. We identified 42 families, 126 genera, and 120 species of plants in the squirrel's diet. Our use of three gene regions was beneficial because reliance on only one gene region (e.g., only trnL ) would have caused us to miss >30% of the taxa in their diet. Northern Idaho ground squirrel diet differed between spring and summer, frequency of many plants in the diet differed from their frequency within their foraging areas (evidence of selective foraging), and several plant genera in their diet were associated with survival. Our results suggest that while these squirrels are generalists (they consume a wide variety of plant species), they are also selective and do not eat plants relative to availability. Consumption of particular genera such as Perideridia may be associated with higher overwinter survival.

Idaho

Examining spatial patterns of selection and use for an altered predator guild

Anthropogenic disturbances have altered species’ distributions potentially impacting interspecific interactions. Interference competition is when one species denies a competing species access to a resource. One mechanism of interference competition is aggression, which can result in altered space-use of a subordinate species due to the threat of harm, otherwise known as a ‘landscape of fear’. Alternatively, subordinates might outcompete dominant species in resource-poor environments via a superior ability to extract resources. Our goal was to evaluate spatial predictions of the ‘landscape of fear’ hypothesis for a carnivore guild in Newfoundland, Canada, where coyotes recently immigrated. Native Newfoundland carnivores include red foxes, Canada lynx, and black bears. We predicted foxes and lynx would avoid coyotes because of their larger size and similar dietary niches. We used scat-detecting dogs and genetic techniques to locate and identify predator scats. We then built resource selection functions and tested for avoidance by incorporating predicted values of selection for the alternative species into the best supported models of each species. We found multiple negative relationships, but notably did not find avoidance by foxes of areas selected by coyotes. While we did find that lynx avoided coyotes, we also found a reciprocal relationship. The observed patterns suggest spatial partitioning and not coyote avoidance, although avoidance could still be occurring at different spatial or temporal scales. Furthermore, Newfoundland’s harsh climate and poor soils may swing the pendulum of interspecific interactions from interference competition to exploitative competition, where subordinates outcompete dominant competitors through a superior ability to extract resources.

Newfoundland

Effects of breeder turnover and harvest on group composition and recruitment in a social carnivore

Breeder turnover can influence population growth in social carnivores through changes to group size, composition and recruitment. Studies that possess detailed group composition data that can provide insights about the effects of breeder turnover on groups have generally been conducted on species that are not subject to recurrent annual human harvest. We wanted to know how breeder turnover affects group composition and how harvest, in turn, affects breeder turnover in cooperatively breeding grey wolves ( Canis lupus Linnaeus 1758). We used noninvasive genetic sampling at wolf rendezvous sites to construct pedigrees and estimate recruitment in groups of wolves before and after harvest in Idaho, USA. Turnover of breeding females increased polygamy and potential recruits per group by providing breeding opportunities for subordinates although resultant group size was unaffected 1 year after the turnover. Breeder turnover had no effect on the number of nonbreeding helpers per group. After breeding male turnover, fewer female pups were recruited in the new males’ litters. Harvest had no effect on the frequency of breeder turnover. We found that breeder turnover led to shifts in the reproductive hierarchies within groups and the resulting changes to group composition were quite variable and depended on the sex of the breeder lost. We hypothesize that nonbreeding females direct help away from non-kin female pups to preserve future breeding opportunities for themselves. Breeder turnover had marked effects on the breeding opportunities of subordinates and the number and sex ratios of subsequent litters of pups. Seemingly subtle changes to groups, such as the loss of one individual, can greatly affect group composition, genetic content, and short-term population growth when the individual lost is a breeder.

Idaho

Harvest and group effects on pup survival in a cooperative breeder

Recruitment in cooperative breeders can be negatively affected by changes in group size and composition. The majority of cooperative breeding studies have not evaluated human harvest; therefore, the effects of recurring annual harvest and group characteristics on survival of young are poorly understood. We evaluated how harvest and groups affect pup survival using genetic sampling and pedigrees for grey wolves in North America. We hypothesized that harvest reduces pup survival because of (i) reduced group size, (ii) increased breeder turnover and/or (iii) reduced number of female helpers. Alternatively, harvest may increase pup survival possibly due to increased per capita food availability or it could be compensatory with other forms of mortality. Harvest appeared to be additive because it reduced both pup survival and group size. In addition to harvest, turnover of breeding males and the presence of older, non-breeding males also reduced pup survival. Large groups and breeder stability increased pup survival when there was harvest, however. Inferences about the effect of harvest on recruitment require knowledge of harvest rate of young as well as the indirect effects associated with changes in group size and composition, as we show. The number of young harvested is a poor measure of the effect of harvest on recruitment in cooperative breeders.

Proceedings of the Royal Society B: Biological Sci