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This is a manuscript version of an article published as: Haines, J.A., Nason, S.E., Skurdal, A.M.M., Bourchier, T., Boutin, S., Taylor, R.W., McAdam, A.G., Lane, J.E., Kelley, A.D., Humphries, M.M., Gorrell, J.C., Dantzer, B., Coltman, D.W., & Hämäläinen, A. (2020). Sex- and context-specific associations between personality and a measure of fitness but no link with life history traits. Animal Behaviour, 167, 23-29. https://doi.org/10.1016/j.anbehav.2020.06.013 The pace-of-life syndrome hypothesis posits that personality traits (i.e. consistent individual differences in behaviour) are linked to life history and fitness. Specifically, fast-paced individuals are predicted to be proactive (i.e. active and aggressive) with an earlier age at first reproduction, a shorter life span and higher fecundity than slow-paced individuals. Environmental conditions and sex differences may be important in maintaining behavioural and life history variation in populations and may influence the covariance of personality with life history or lifetime fitness. However, these effects are rarely tested together. We investigated whether the occurrence of a resource pulse (called a mast year) during adulthood altered the associations between personality and life history traits or lifetime offspring production in adult North American red squirrels, Tamiasciurus hudsonicus. Despite accounting for environmental context during adulthood, we found no evidence of an overall pace-of-life syndrome in this population as personality was not associated with age at first reproduction or longevity in either sex. Males and females had similar activity levels, but females were more aggressive, potentially due to the fitness benefits of protecting their offspring from predation. In all females, regardless of mast experience, there was no association between activity and lifetime pup production, but there was a positive association between aggression and lifetime pup production. In males that experienced a mast, there was a positive association between lifetime pup production and both activity and aggression. In males that did not experience a mast, there was no association between activity and lifetime pup production, but there was a negative association between aggression and lifetime pup production. Lifetime recruit production (number of adult offspring recruited into the breeding population) in either sex was not influenced by activity or aggression regardless of mast experience. Overall, our results suggest that the infrequent occurrence of mast years may contribute to maintaining variation in personality traits in red squirrels. Pre-print version This is a manuscript version of an article published as: Haines, J.A., Nason, S.E., Skurdal, A.M.M., Bourchier, T., Boutin, S., Taylor, R.W., McAdam, A.G., Lane, J.E., Kelley, A.D., Humphries, M.M., Gorrell, J.C., Dantzer, B., Coltman, D.W., & Hämäläinen, A. (2020). Sex- and context-specific associations between personality and a measure of fitness but no link with life history traits. Animal Behaviour, 167, 23-29. https://doi.org/10.1016/j.anbehav.2020.06.013 https://viurrspace.ca/bitstream/handle/10613/23323/GorrellAB.pdf?sequence=3

This document is a pre-copyedited, author-provided manuscript version of a published work that after revisions following peer review and technical editing by the publisher appeared in final form as: Hessels, E.W., Lofroth, E.C., Weir, R.D., & Gorrell, J.C. (2021). Characterizing the elusive Vancouver Island wolverine, Gulo gulo vancouverensis, using historical DNA. Journal of Mammalogy, 102(2), 530-540. To access the final edited and published work see: https://doi.org/10.1093/jmammal/gyaa175, The wolverine (Gulo gulo) is a Holarctic species found in North America primarily across the boreal forest, the subarctic, and along the Pacific coast, including Vancouver Island (VI), British Columbia. While wolverines on VI are rare and possibly extirpated, they have been previously described as a unique subspecies, G. g. vancouverensis, distinct from G. g. luscus from the mainland of North America. However, the validity of the VI subspecies is contentious, with conflicting results from studies of skull morphology. Here, we used molecular analyses to characterize the genetic diversity of the VI population and resolve this taxonomic debate to assist with conservation priorities. Historical DNA of VI wolverines was obtained from museum specimens, amplified at 16 nuclear microsatellite loci, and sequenced at the mitochondrial D-loop control region to compare with wolverines from mainland British Columbia. The VI population had lower allelic richness and was fixed for a single common mtDNA haplotype. Bayesian and non-Bayesian assignments using microsatellites generally revealed admixture across populations, implying allele frequencies between the VI and mainland populations were not significantly different. Hence, both types of genetic markers showed little evolutionary divergence between VI and the mainland population. Combined, these results do not provide evidence of significant genetic distinction for VI wolverines, nor support the subspecific classification. Immediate conservation efforts should focus on estimating population size, while future conservation planning can assume VI wolverines likely are not a unique genetic population and there remains the potential for natural recolonization of wolverines to VI.

Phenological shifts are the most widely reported ecological responses to climate change, but the requirements to distinguish their causes (i.e. phenotypic plasticity vs. microevolution) are rarely met. To do so, we analysed almost two decades of parturition data from a wild population of North American red squirrels (Tamiasciurus hudsonicus). Although an observed advance in parturition date during the first decade provided putative support for climate change-driven microevolution, a closer look revealed a more complex pattern. Parturition date was heritable [h2 = 0.14 (0.07-0.21 (HPD interval)] and under phenotypic selection [β = -0.14 ± 0.06 (SE)] across the full study duration. However, the early advance reversed in the second decade. Further, selection did not act on the genetic contribution to variation in parturition date, and observed changes in predicted breeding values did not exceed those expected due to genetic drift. Instead, individuals responded plastically to environmental variation, and high food [white spruce (Picea glauca) seed] production in the first decade appears to have produced a plastic advance. In addition, there was little evidence of climate change affecting the advance, as there was neither a significant influence of spring temperature on parturition date or evidence of a change in spring temperatures across the study duration. Heritable traits not responding to selection in accordance with quantitative genetic predictions have long presented a puzzle to evolutionary ecologists. Our results on red squirrels provide empirical support for one potential solution: phenotypic selection arising from an environmental, as opposed to genetic, covariance between the phenotypic trait and annual fitness.

Garry oak (Quercus garryana Douglas ex Hook.) is a deciduous tree whose ecosystem is listed “at risk” throughout its range in British Columbia (BC), Canada, under the Canadian Species at Risk Act. Garry oak ecosystems host the most diverse flora for coastal BC, yet they account for less than 0.3% of the province’s land coverage. Due to the loss and degradation of Garry oak habitat, many associated plant and animal species that rely on these sensitive ecosystems are endangered. Microsatellite markers were used to investigate temporal changes in fine-scale population genetic structure of 121 Garry oak trees from the Nanaimo region (Vancouver Island, BC) using diameter at breast height as a proxy for age. Overall, allelic diversity was moderate, ranging from 3.0 to 7.5 alleles per locus with a mean of 4.4 (±0.4 SE) across all loci. Global fixation indices (FST) of 0.06 and 0.09 suggest significant departures from Hardy–Weinberg equilibrium among all populations and age-classified subpopulations, respectively. We found no evidence for change in genetic diversity across generations. Our results indicate low levels of differentiation within populations and high levels of gene flow among populations, suggesting an adaptive potential for Garry oaks in response to future climate change events.

In many bird species, females mate with males other than their social partner, a behavior considered to be beneficial in terms of inbreeding avoidance, or to attract additional care of offspring from EP males. In noisy miners, we show that although 27% of broods contained EP offspring, EP males did not provide help to those broods. Moreover, we did not find evidence that this behavior functions as a mechanism to avoid inbreeding in this species.

Background Our understanding of gut microbiota has been limited primarily to findings from human and laboratory animals, but what shapes the gut microbiota in nature remains largely unknown. To fill this gap, we conducted a comprehensive study of gut microbiota of a well-studied North American red squirrel (Tamiasciurus hudsonicus) population. Red squirrels are territorial, solitary, and live in a highly seasonal environment and therefore represent a very attractive system to study factors that drive the temporal and spatial dynamics of gut microbiota. Result For the first time, this study revealed significant spatial patterns of gut microbiota within a host population, suggesting limited dispersal could play a role in shaping and maintaining the structure of gut microbial communities. We also found a remarkable seasonal rhythm in red squirrel’s gut microbial composition manifested by a tradeoff between relative abundance of two genera Oscillospira and Corpococcus and clearly associated with seasonal variation in diet availability. Our results show that in nature, environmental factors exert a much stronger influence on gut microbiota than host-associated factors including age and sex. Despite strong environmental effects, we found clear evidence of individuality and maternal effects, but host genetics did not seem to be a significant driver of the gut microbial communities in red squirrels. Conclusion Taken together, the results of this study emphasize the importance of external ecological factors rather than host attributes in driving temporal and spatial patterns of gut microbiota in natural environment. Electronic supplementary material The online version of this article (10.1186/s40168-017-0382-3) contains supplementary material, which is available to authorized users.

Interactions with conspecifics are an important aspect of an individual's environment. Although it is well known that the presence of conspecifics can have important effects on behaviour, in general it is also now acknowledged that the composition of the social environment can vary, and that this variation may have profound effects on individual behaviour and fitness. Using a wild population of North American red squirrels, Tamiasciurus hudsonicus, we investigated the importance of the composition of the social environment in a territorial species by assessing whether the risk of intrusion faced by territory owners varied with the degree of relatedness and familiarity in their social neighbourhoods. To test this, we conducted temporary removals of territory owners and observed the time until intrusion and the identity of intruding individuals. We found that individuals in neighbourhoods with low average familiarity faced a higher risk of intrusion and that unfamiliar neighbours were more likely to intrude. Surprisingly, we found that related neighbours also posed a higher risk of intrusion. The results from our study suggest that familiarity with neighbours may be an ecologically and evolutionarily relevant measure of the social environment, even in a species considered to be ‘asocial’. Future studies should consider the potential importance of the social environment, which has heretofore been mostly overlooked, as a relevant selective pressure in asocial, territorial species.

Assessments of population genetic structure have become an increasing focus as they can provide valuable insight into patterns of migration and gene flow. structure, the most highly cited of several clustering-based methods, was developed to provide robust estimates without the need for populations to be determined a priori. structure introduces the problem of selecting the optimal number of clusters, and as a result, the ΔK method was proposed to assist in the identification of the "true" number of clusters. In our review of 1,264 studies using structure to explore population subdivision, studies that used ΔK were more likely to identify K = 2 (54%, 443/822) than studies that did not use ΔK (21%, 82/386). A troubling finding was that very few studies performed the hierarchical analysis recommended by the authors of both ΔK and structure to fully explore population subdivision. Furthermore, extensions of earlier simulations indicate that, with a representative number of markers, ΔK frequently identifies K = 2 as the top level of hierarchical structure, even when more subpopulations are present. This review suggests that many studies may have been over- or underestimating population genetic structure; both scenarios have serious consequences, particularly with respect to conservation and management. We recommend publication standards for population structure results so that readers can assess the implications of the results given their own understanding of the species biology.

This is a manuscript version of an article published as: Haines, J.A., Nason, S.E., Skurdal, A.M.M., Bourchier, T., Boutin, S., Taylor, R.W., McAdam, A.G., Lane, J.E., Kelley, A.D., Humphries, M.M., Gorrell, J.C., Dantzer, B., Coltman, D.W., & Hämäläinen, A. (2020). Sex- and context-specific associations between personality and a measure of fitness but no link with life history traits. Animal Behaviour, 167, 23-29. https://doi.org/10.1016/j.anbehav.2020.06.013 The Canadian Rocky Mountains are one of the few places on Earth where the spatial genetic structure of wide-ranging species has been relatively unaffected by anthropogenic disturbance. We characterized the spatial genetic structure of Rocky Mountain bighorn sheep (Ovis canadensis canadensis Shaw, 1804) in the northern portion of their range. Using microsatellites from 1495 individuals and mitochondrial DNA sequences from 188 individuals, we examined both broad- and fine-scale spatial genetic structure, assessed sex-biased gene flow within the northern portion of the species range, and identified geographic patterns of genetic diversity. We found that broad-scale spatial genetic structure was consistent with barriers to movement created by major river valleys. The fine-scale spatial genetic structure was characterized by a strong isolation-by-distance pattern, and analysis of neighborhood size using spatial autocorrelation indicated gene flow frequently occurred over distances of up to 100 km. However, analysis of sex-specific spatial autocorrelation and analysis of mitochondrial haplotype distributions failed to detect any evidence of sex-biased gene flow. Finally, our analyses reveal decreasing genetic diversity with increasing latitude, consistent with patterns of post-glacial recolonization of the Rocky Mountains. Post-print version This is an author supplied, accepted manuscript version of an article published as: Deakin, S., Gorrell, J.C., Kneteman, J., Hik, D.S., Jobin, R.M., & Coltman, D.W. (2020). Spatial genetic structure of Rocky Mountain bighorn sheep (Ovis canadensis canadensis) at the northern limit of their native range. Canadian Journal of Zoology, 98(5), 317-330. DOI: 10.1139/cjz-2019-0183 Canadian Journal of Zoology is available online at: http://www.nrcresearchpress.com/journal/cjz, and this article is available at: http://dx.doi.org/10.1139/cjz-2019-0183. https://viurrspace.ca/bitstream/handle/10613/23294/Gorrell.CJZ.pdf?sequence=3

This is the peer reviewed version of the following article: Cullingham, C.I., Miller, J.M., Peery, R.M., Dupuis, J.R., Malenfant, R.M., Gorrell, J.C., & Janes, J.K. (2020). Confidently identifying the correct K value using the ΔK method: When does K=2? Molecular Ecology, 29(5), 862-869, which has been published in final form at http://doi.org/10.1111/mec.15374. This article may be used for non-commercial purposes in accordance with Wiley Terms and Conditions for Use of Self-Archived Versions., Populations delineated based on genetic data are commonly used for wildlife conservation and management. Many studies use the program structure combined with the ΔK method to identify the most probable number of populations (K). We recently found K = 2 was identified more often when studies used ΔK compared to studies that did not. We suggested two reasons for this: hierarchical population structure leads to underestimation, or the ΔK method does not evaluate K = 1 causing an overestimation. The present contribution aims to develop a better understanding of the limits of the method using one, two and three population simulations across migration scenarios. From these simulations we identified the “best K” using model likelihood and ΔK. Our findings show that mean probability plots and ΔK are unable to resolve the correct number of populations once migration rate exceeds 0.005. We also found a strong bias towards selecting K = 2 using the ΔK method. We used these data to identify the range of values where the ΔK statistic identifies a value of K that is not well supported. Finally, using the simulations and a review of empirical data, we found that the magnitude of ΔK corresponds to the level of divergence between populations. Based on our findings, we suggest researchers should use the ΔK method cautiously; they need to report all relevant data, including the magnitude of ΔK, and an estimate of connectivity for the research community to assess whether meaningful genetic structure exists within the context of management and conservation.

Kin recognition can facilitate kin selection and may have played a role in the evolution of sociality. Red squirrels (Tamiasciurus hudsonicus) defend territories using vocalizations known as rattles. They use rattles to discriminate kin, though the mechanism underlying this ability is unknown. Our objective was to distinguish between the mechanisms of prior association, where animals learn the phenotypes of kin they associate with early in life, and phenotype matching/recognition alleles, where animals use a template to match phenotypes, thereby allowing them to recognize kin without an association early in life. We used audio playbacks to measure the responses of squirrels to rattles from familiar kin, unfamiliar kin, and non-kin. Initial analyses revealed that red squirrels did not discriminate between familiar and unfamiliar kin, but also did not discriminate between kin and non-kin, despite previous evidence indicating this capability. Post hoc analyses showed that a squirrel’s propensity to rattle in response to playback depended on an interaction between relatedness and how the playback stimuli had been recorded. Red squirrels discriminated between rattles from close kin (r = 0.5) and rattles from non-kin (r < 0.125) when the rattles were recorded from provoked squirrels. Squirrels did not exhibit kin discrimination in response to unsolicited rattles. Once we accounted for how the stimuli had been recorded, we found no difference in the responses to familiar and unfamiliar kin. Our study suggests that kin discrimination by red squirrels may be context dependent.

North American red squirrels (Tamiasciurus hudsonicus) anticipate when white spruce (Picea glauca), their primary food source, will produce large amounts of cones in infrequent and irregular mast years (Boutin et al. 2006). Cones mature in autumn and are then available as food for red squirrels, but females produce larger, and often multiple, litters the preceding spring and summer in anticipation of the upcoming mast. Because this pulse of the cone food resource follows the birth of the litter, it cannot be a source of energy for the female to produce young; instead, there must be cues for increased reproductive investment by the females prior to mast cone production, perhaps through consumption of buds on the masting trees (Boutin et al. 2006, 2013). Boutin et al. (2006) only studied females; whether male behavior anticipates mast years is still unstudied. At our study area in the Yukon, 2014 was a mast year for spruce cone production in late summer (Lamontagne et al. 2005). J. A. Haines was observing male red squirrel mating behavior during spring 2014, giving her an unanticipated opportunity to document the previously unstudied effects of a mast year on male red squirrels., Data collection was supported by grants provided to S. Boutin, A. G. McAdam, M. M. Humphries, J. E. Lane, and D. W. Coltman from the Natural Sciences and Engineering Research Council of Canada, as well as National Science Foundation grants to A. G. McAdam and B. Dantzer. J. A. Haines was supported by an NSERC Doctoral Postgraduate Scholarship, University of Alberta Canadian Circumpolar Institute Circumpolar/Boreal Alberta Research Funds and Aboriginal Affairs and Northern Development Canada Northern Scientific Training Program Grants, and University of Alberta Faculty of Graduate Studies and Research scholarships., "The Scientific Naturalist", Copyright by the Ecological Society of America. This is an electronic version of an article published as: Haines, J.A., Coltman, D.W., Dantzer, B., Gorrell, J.C., Humphries, M.M., Lane, J.E., McAdam, A.G., & Boutin, S. (2018). Sexually selected infanticide by male red squirrels in advance of a mast year. Ecology, 99(5), 1242-1244. DOI: 10.1002/ecy.2158 Ecology is published by the Ecological Society of America and is available online at: https://esajournals.onlinelibrary.wiley.com/journal/19399170/, and this article is available at: https://esajournals.onlinelibrary.wiley.com/doi/full/10.1002/ecy.2158., https://viurrspace.ca/bitstream/handle/10613/6527/Gorrell.Ecology.99.5.pdf?sequence=3

Pollen DNA metabarcoding-marker-based genetic identification of potentially mixed-species pollen samples-has applications across a variety of fields. While basic species-level pollen identification using standard DNA barcode markers is established, the extent to which metabarcoding (a) correctly assigns species identities to mixes (qualitative matching) and (b) generates sequence reads proportionally to their relative abundance in a sample (quantitative matching) is unclear, as these have not been assessed relative to known standards. We tested the quantitative and qualitative robustness of metabarcoding in constructed pollen mixtures varying in species richness (1-9 species), taxonomic relatedness (within genera to across class) and rarity (5%-100% of grains), using Illumina MiSeq with the markers rbcL and ITS2. Qualitatively, species composition determinations were largely correct, but false positives and negatives occurred. False negatives were typically driven by lack of a barcode gap or rarity in a sample. Species richness and taxonomic relatedness, however, did not strongly impact correct determinations. False positives were likely driven by contamination, chimeric sequences and/or misidentification by the bioinformatics pipeline. Quantitatively, the proportion of reads for each species was only weakly correlated with its relative abundance, in contrast to suggestions from some other studies. Quantitative mismatches are not correctable by consistent scaling factors, but instead are context-dependent on the other species present in a sample. Together, our results show that metabarcoding is largely robust for determining pollen presence/absence but that sequence reads should not be used to infer relative abundance of pollen grains.

The objectives of this study were to describe and evaluate potential drivers of genetic structure in Canadian breeding populations of the Ovenbird, Seiurus aurocapilla. We performed genetic analyses on feather samples of individuals from six study sites using nuclear microsatellites. We also assessed species identity and population genetic structure of quill mites (Acariformes, Syringophilidae). For male Ovenbirds breeding in three study sites, we collected light-level geolocator data to document migratory paths and identify the wintering grounds. We also generated paleohindcast projections from bioclimatic models of Ovenbird distribution to identify potential refugia during the last glacial maximum (LGM, 21,000 years before present) as a factor explaining population genetic structure. Birds breeding in the Cypress Hills (Alberta/Saskatchewan) may be considered a distinct genetic unit, but there was no evidence for genetic differentiation among any other populations. We found relatively strong migratory connectivity in both western and eastern populations, but some evidence of mixing among populations on the wintering grounds. There was also little genetic variation among syringophilid mites from the different Ovenbird populations. These results are consistent with paleohindcast distribution predictions derived from two different global climate models indicating a continuous single LGM refugium, with the possibility of two refugia. Our results suggest that Ovenbird populations breeding in boreal and hemiboreal regions are panmictic, whereas the population breeding in Cypress Hills should be considered a distinct management unit. Natural Sciences and Engineering Research Council of Canada (NSERC); Alberta Conservation Association; NSERC Postgraduate Scholarship; Queen Elizabeth II Graduate Scholarship; University of Alberta; Canadian Circumpolar Institute; Northern Scientific Program This is an electronic version of an article which was originally published as: Haché, S., Bayne, E.M., Villard, M.-A., Proctor, H., Davis, C.S., Stralberg, D., … Krikun, R. (2017). Phylogeography of a migratory songbird across its Canadian breeding range: Implication for conservation units. Ecology and Evolution, 1-11, which has been published at http://dx.doi.org/10.1002/ece3.3170. https://viuspace.viu.ca/bitstream/handle/10613/5027/Gorrell.EE.Article.pdf?sequence=5

A trait must genetically correlate with fitness in order to evolve in response to natural selection, but theory suggests that strong directional selection should erode additive genetic variance in fitness and limit future evolutionary potential. Balancing selection has been proposed as a mechanism that could maintain genetic variance if fitness components trade off with one another and has been invoked to account for empirical observations of higher levels of additive genetic variance in fitness components than would be expected from mutation– selection balance. Here, we used a long-term study of an individually marked population of North American red squirrels (Tamiasciurus hudsonicus) to look for evidence of (1) additive genetic variance in lifetime reproductive success and (2) fitness trade-offs between fitness components, such as male and female fitness or fitness in high- and low-resource environments. “Animal model” analyses of a multigenerational pedigree revealed modest maternal effects on fitness, but very low levels of additive genetic variance in lifetime reproductive success overall as well as fitness measures within each sex and environment. It therefore appears that there are very low levels of direct genetic variance in fitness and fitness components in red squirrels to facilitate contemporary adaptation in this population. This is an electronic version of an article that was originally published as: McFarlane, S.E., Gorrell, J.C., Coltman, D.W., Humphries, M.M., Boutin, S., & McAdam, A.G. (2014). Very low levels of direct additive genetic variance in fitness and fitness components in a red squirrel population. Ecology and Evolution, 4(10), 1729-1738. DOI: 10.1002/ece3.982. Ecology and Evolution is an open access journal published by Wiley-Blackwell. More information about the journal can be found at: http://onlinelibrary.wiley.com/journal/10.1002/(ISSN)2045-7758. This article can be accessed at: http://onlinelibrary.wiley.com/doi/10.1002/ece3.982/full. Ecology and Evolution Journal Cover Image (Volume 4, Issue 10) : A grizzly bear (Ursus arctos) of the Greater Yellowstone Ecosystem, which encompasses portions of Wyoming, Montana, and Idaho, USA. Photo by Ray Paunovick, Wild Planet Films, Bozeman, Montana, USA. https://viuspace.viu.ca/bitstream/handle/10613/3208/Gorrell.EE.pdf?sequence=4

In polyandrous and polygynandrous species where females mate with multiple partners, males are expected to maximize their fitness by exhibiting an array of reproductive behaviors to ensure fertilization success, such as competing for the best mating order within a mating sequence, optimizing their investment in copulation, and mate guarding. Though there is genetic evidence of a first-male precedence in siring success for many mammalian species, the causes of this effect are poorly understood. We studied influences on first-male precedence in Columbian ground squirrels (Urocitellus columbianus). We found that the time a male spent consorting and mate guarding declined with his mating order (both the highest for the first male to mate). Mate guarding by the first male significantly reduced, but did not exclude, the number of additional males a female accepted. Later mating males reduced the time spent consorting, suggesting a perceived decreased chance of fertilization success. Consortship and mate guarding durations were positively related to the male’s siring success and to each other, suggesting that males adjusted these behaviors strategically to increase their chances of fertilization success. Our results suggest that besides being the first male to consort, first-male sperm precedence is further enhanced through longer mating bouts and by suppressing the chances and/or efforts of later mating males.

Multiple mating by females is common in many mammalian species, often resulting in mixed paternity litters. In such mating systems, mating order, male age, and male body mass frequently play an important role in determining male reproductive success. We tested for these effects on male reproductive success in Columbian ground squirrels (Urocitellus columbianus). The mating activity of estrous females was observed, and the occurrence of sperm precedence was tested using microsatellites to determine paternity in a total of 147 litters (434 offspring), including 110 litters (334 offspring) where the mating position of individual males was determined. Females mated with up to 8 males per litter, whereas paternity analyses revealed that only the first 5 males to mate actually sired offspring. The number of offspring sired significantly decreased with position in the mating sequence, showing a strong first male advantage. The extent of this advantage diminished with an increasing number of male mating partners, indicating that sperm competition plays an important role. A male’s position in the females’ mating sequences was not consistent within and across seasons, suggesting that individual males did not follow distinct reproductive strategies. Rather, males of intermediate age were more successful than young and old males, when corrected for age effects; heavier males were more likely to mate first. We conclude that males gain the largest part of their seasonal reproductive output from mating first with a female due to a pronounced first male advantage but gain considerable additional fitness from mating with additional, already mated females. Key words: Columbian ground squirrel, first male advantage, mating strategy, multiple paternity, senescence, Urocitellus columbianus. [Behav Ecol]

Most empirical attempts to explain the evolution of parental care have focused on its costs and benefits (i.e. fitness consequences). In contrast, few investigations have been made of the other necessary prerequisite for evolutionary change, inheritance. Here, we examine the fitness consequences and heritability (h(2)) of a post-weaning parental care behaviour (territory bequeathal) in a wild population of North American red squirrels. Each year, a subset (average across all years = 19%) of reproductive females bequeathed their territory to a dependent offspring. Bequeathing females experienced higher annual reproductive success and did not suffer a survival cost to themselves relative to those females retaining their territory. Bequeathing females thus realized higher relative annual fitness [ω = 1.18 ± 0.03 (SE)] than nonbequeathing females [ω = 0.96 ± 0.02 (SE)]. Additive genetic influences on bequeathal behaviour, however, were not significantly different from 0 (h(2) = 1.9 × 10(-3); 95% highest posterior density interval = 3.04 × 10(-8) to 0.37) and, in fact, bequeathal behaviour was not significantly repeatable (R = 2.0 × 10(-3); 95% HPD interval = 0-0.27). In contrast, directional environmental influences were apparent. Females were more likely to bequeath in years following low food abundance and when food availability in the upcoming autumn was high. Despite an evident fitness benefit, a lack of heritable genetic variance will constrain evolution of this trait.

Orphaned animals benefit from being adopted, but it is unclear why an adopting parent should incur the costs of rearing extra young. Such altruistic parental behaviour could be favoured if it is directed towards kin and the inclusive benefits of adoption exceed the costs. Here, we report the occurrence of adoption (five occurrences among 2,230 litters over 19 years) in asocial red squirrels (Tamiasciurus hudsonicus). Adoptions were always between kin, while orphans without nearby kin were never adopted. Adoptions were confined exclusively to circumstances in which the benefits to the adopted juvenile (b), discounted by the degree of relatedness between the surrogate and the orphan (r), exceeded the fitness costs of adding an extra juvenile to her litter (c), as predicted by Hamilton's rule (rb>c) for the evolution of altruism. By focusing on adoption in an asocial species, our study provides a clear test of Hamilton's rule that explains the persistence of occasional altruism in a natural mammal population. This is an electronic version of an article that was originally published as: Gorrell, J.C., McAdam, A.G., Coltman, D.W., Humphries, M.M., & Boutin, S. (2010). Adopting kin enhances inclusive fitness in asocial red squirrels. Nature Communications, 1, 1-4. DOI: 10.1038/ncomss1022. Nature Communications is published by Macmillan Publishers, part of Springer Nature. More information about the journal can be found at: http://www.nature.com/ncomms/. This article can be accessed on the Nature.com website at: http://www.nature.com/articles/ncomms1022. https://viuspace.viu.ca/bitstream/handle/10613/3207/Gorrell.NC.pdf?sequence=4

An understanding of mating systems and fine-scale spatial genetic structure is required to effectively manage forest pest species such as Dendroctonus ponderosae (mountain pine beetle). Here we used genome-wide single-nucleotide polymorphisms to assess the fine-scale genetic structure and mating system of D. ponderosae collected from a single stand in Alberta, Canada. Fine-scale spatial genetic structure was absent within the stand and the majority of genetic variation was best explained at the individual level. Relatedness estimates support previous reports of pre-emergence mating. Parentage assignment tests indicate that a polygamous mating system better explains the relationships among individuals within a gallery than the previously reported female monogamous/male polygynous system. Furthermore, there is some evidence to suggest that females may exploit the galleries of other females, at least under epidemic conditions. Our results suggest that current management models are likely to be effective across large geographic areas based on the absence of fine-scale genetic structure.