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Predicting the geographic range of species and their response to climatic variation and change are entwined goals in conservation and evolutionary ecology. Species distribution models (SDMs) are foundational in this effort and used to visualize the geographic range of species as the spatial representation of its realized niche. SDMs are also used to forecast range shifts under climate change, but often in the absence of empirical evidence that climate limits population growth. We explored the influence of climate on demography, seasonal migration, and the extent of the geographic range in song sparrows (Melospiza melodia), a species thought to display marked local adaptation to regional climate. To do so, we developed SDMs to predict the demographic and climate niches of migratory and resident song sparrows across our study area in western North America from California to Alaska, using 48 years of demographic data from a focal population in British Columbia and 1.2 million continental-scale citizen science observations. Spatial agreement of our demographic and climate niche models in the region of our focal population was strong (76%), supporting the hypothesis that demographic performance and the occurrence of seasonal migration varied predictably with climatic conditions. In contrast, agreement at the northern (58%) and southern (40%) extents of our study area was lower, as expected if the factors limiting population growth vary regionally. Our results support the hypothesis that local climate drives spatial variation in the occurrence of seasonal migration in song sparrows by limiting the fitness of year-round residents, and suggest that climate warming has favored range expansions and facilitated an upward shift in elevational range song sparrows that forgo seasonal migration. Our work highlights the potential role of seasonal migration in climate adaptation and limits on the reliability of climate niche models not validated with demographic data.

Individuals vary in their immune genotype, inbreeding coefficient f, immune responses, survival to adulthood, and adult longevity. However, whether immune genes predict survival or longevity, whether such relationships are mediated through immune responses, and how f affects immune genotype remain unclear. We use a wild song sparrow (Melospiza melodia) population in which survival to adulthood, adult longevity, and f were measured precisely, and in which immune responses have previously been assessed. We investigate four toll-like receptor (TLR) and the major histocompatibility complex (MHC) class IIB exon 2 genes. We test: a) whether immune genes predict fitness (survival to adulthood or adult longevity); b) whether immune genes predict immune response; c) whether immune response predicts fitness; and d) whether fitness, immune responses, or immune genotypes are correlated with f. We find that survival to adulthood is not associated with immune gene variation, but adult longevity is decreased by high MHC allele diversity (especially in birds that were relatively outbred), and by the presence of a specific MHC supertype. Immune responses were affected by specific immune genotypes. Survival to adulthood and adult longevity were not predicted by immune response, implying caution in the use of immune response as a predictor for fitness. We also found no relationship between f and immune genotype. This finding indicates that immune gene associations with longevity and immune response are not artefacts of f, and suggests that pathogen-mediated selection at functional loci can slow the loss of genetic variation arising from genetic drift and small population size.

Non-native earthworms can alter ecosystems by modifying soil structure, depredating seeds and seedlings, and consuming soil organic matter, yet the initial responses of plant communities to earthworm invasions remain poorly understood. We assessed the effect of non-native earthworms on seedling survival during germination and after establishment using six native and six non-native plant species grown from seed in single- and multi-species experimental mesocosms. We examined the extent to which earthworms (1) influenced seedling survival, (2) selectively depredated native versus non-native plants, (3) impacted establishment based on seed size and/or root morphology, and (4) shaped community assembly. The effect of earthworms on seedling survival varied temporally and among species but inconsistently with respect to species origin. Differences in seed/seedling survival translated to changes in community assembly. Earthworms tended to reduce species abundance, richness, evenness, and diversity in multi-species mesocosms and led to the divergence of communities by treatment. In general, species with large seeds and fibrous roots dominated communities with earthworms present, whereas species with small seeds and taproots only persisted in multi-species mesocosms without earthworms. Our findings suggest that earthworms act as ecological filters in the early stages of invasion to shape community composition based on plant morphological traits.

Funding: Hoge Veluwe great tits: the NIOO-KNAW, ERC, and numerous funding agencies; Wytham great tits: Biotechnology and Biological Sciences Research Council, ERC, and the UK Natural Environment Research Council (NERC). The rate of adaptive evolution, the contribution of selection to genetic changes that increase mean fitness, is determined by the additive genetic variance in individual relative fitness. To date, there are few robust estimates of this parameter for natural populations, and it is therefore unclear whether adaptive evolution can play a meaningful role in short-term population dynamics. We developed and applied quantitative genetic methods to long-term datasets from 19 wild bird and mammal populations and found that, while estimates vary between populations, additive genetic variance in relative fitness is often substantial and, on average, twice that of previous estimates. We show that these rates of contemporary adaptive evolution can affect population dynamics and hence that natural selection has the potential to partly mitigate effects of current environmental change. Postprint

Ongoing adaptive evolution, and resulting “evolutionary rescue” of declining populations, requires additive genetic variation in fitness. Such variation can be increased by gene flow resulting from immigration, potentially facilitating evolution. But, gene flow could in fact constrain rather than facilitate local adaptive evolution if immigrants have low additive genetic values for local fitness. Local migration-selection balance and micro-evolutionary stasis could then result. However, key quantitative genetic effects of natural immigration, comprising the degrees to which gene flow increases the total local additive genetic variance yet counteracts local adaptive evolutionary change, have not been explicitly quantified in wild populations. Key implications of gene flow for population and evolutionary dynamics consequently remain unclear. Our quantitative genetic analyses of long-term data from free-living song sparrows (Melospiza melodia) show that mean breeding value for local juvenile survival to adulthood, a major component of fitness, increased across cohorts more than expected solely due to drift. Such micro-evolutionary change should be expected given nonzero additive genetic variance and consistent directional selection. However, this evolutionary increase was counteracted by negative additive genetic effects of recent immigrants, which increased total additive genetic variance but prevented a net directional evolutionary increase in total additive genetic value. These analyses imply an approximate quantitative genetic migration-selection balance in a major fitness component, and hence demonstrate a key mechanism by which substantial additive genetic variation can be maintained yet decoupled from local adaptive evolutionary change.

Elucidating forces capable of driving species diversification in the face of gene flow remains a key goal in evolutionary biology. Song sparrows, Melospiza melodia, occur as 25 subspecies in diverse habitats across North America, are among the continent's most widespread vertebrate species, and are exemplary of many highly variable species for which the conservation of locally adapted populations may be critical to their range-wide persistence. We focus here on six morphologically distinct subspecies resident in the San Francisco Bay region, including three salt-marsh endemics and three residents in upland and riparian habitats adjacent to the Bay. We used reduced-representation sequencing to generate 2,773 SNPs to explore genetic differentiation, spatial population structure, and demographic history. Clustering separated individuals from each of the six subspecies, indicating subtle differentiation at microgeographic scales. Evidence of limited gene flow and low nucleotide diversity across all six subspecies further supports a hypothesis of isolation among locally adapted populations. We suggest that natural selection for genotypes adapted to salt marsh environments and changes in demography over the past century have acted in concert to drive the patterns of diversification reported here. Our results offer evidence of microgeographic specialization in a highly polytypic bird species long discussed as a model of sympatric speciation and rapid adaptation, and they support the hypothesis that conserving locally adapted populations may be critical to the range-wide persistence of similarly highly variable species.

Declines in global biodiversity due to land conversion and habitat loss are driving a ‘Sixth Mass Extinction’ and many countries fall short of meeting even nominal targets for land protection. We explored how such shortfalls in Australia, Brazil and Canada might be addressed by enhancing partnerships between Indigenous communities and other government agencies that recognize and reward the existing contributions of Indigenous-managed lands to global biodiversity conservation, and their potential contribution to meeting international treaty targets. We found that Indigenous-managed lands were slightly more vertebrate species rich than existing protected areas in all three countries, and in Brazil and Canada, that they supported more threatened vertebrate species than existing protected areas or randomly selected non-protected areas. Our results suggest that overall, Indigenous-managed lands and existing protected areas host similar levels of vertebrate biodiversity in Brazil, Canada, and Australia. Partnerships with Indigenous communities that seek to maintain or enhance Indigenous land tenure practices on Indigenous-managed lands may therefore have some potential to ameliorate national and global shortfalls in land protection for biodiversity conservation using a mix of conventional protected areas and Indigenous-managed lands.

Theory suggests that different taxa having colonized a similar, challenging environment will show parallel or lineage-specific adaptations to shared selection pressures, but empirical examples of parallel evolution in independent taxa are exceedingly rare. We employed comparative genomics to identify parallel and lineage-specific responses to selection within and among four species of North American sparrows that represent four independent, post-Pleistocene colonization events by an ancestral, upland subspecies and a derived salt marsh specialist. We identified multiple cases of parallel adaptation in these independent comparisons following salt marsh colonization, including selection of 12 candidate genes linked to osmoregulation. In addition to detecting shared genetic targets of selection across multiple comparisons, we found many novel, species-specific signatures of selection, including evidence of selection of loci associated with both physiological and behavioral mechanisms of osmoregulation. Demographic reconstructions of all four species highlighted their recent divergence and small effective population sizes, as expected given their rapid radiation into saline environments. Our results highlight the interplay of both shared and lineage-specific selection pressures in the colonization of a biotically and abiotically challenging habitat and confirm theoretical expectations that steep environmental clines can drive repeated and rapid evolutionary diversification in birds.

Spatial planning and decision-support tools facilitate the consideration of socioecological tradeoffs associated with extractive activities, but insufficient data resources often limit their application. Focusing on birds and mining concessions in the Northern Andes, we illustrate how publicly-available data can be used in spatial prioritization tools to identify where concessions have the potential to impact 22 species of Neotropical migratory birds. Concessions covered 11% of the land area of Colombia, Ecuador, and Peru, and disproportionately occurred within important areas for migratory birds in Colombia and Ecuador, but not Peru. eBird data showed that one-quarter of avian species, including the Olive-sided Flycatcher, Cerulean Warbler, and Canada Warbler, had >10% of their global populations on concession land during the non-breeding season. More worrisome, species of greater conservation importance and with larger population declines (1974–2014) were most likely to co-occur with mining. Our approach highlights how public biodiversity data can be used to predict, avoid, or mitigate potential ecological impacts from extractive activities.

Life‐history theory postulates that physiological traits, such as energy metabolism, can be understood in terms of allocation trade‐offs between self‐maintenance and reproduction over an organism's life span, and data show that metabolic intensity and survival vary inversely with latitude, with tropical birds exhibiting a “slow” pace of life relative to temperature species. However, tropical regions harbour strong environmental gradients of their own, and it remains to be shown whether similar life‐history trade‐offs between metabolism and longevity are reflected among tropical birds of the same latitude. We estimated apparent annual survival in 37 species of tropical passerine birds along an elevational gradient (400–3,000 m) in Peru to test whether variation in survival was influenced by basal metabolic rate (BMR; estimated at the same sites), elevation or both factors. We used path analysis to test our prediction that survival would decline as BMR increased, while accounting for the potential direct effects of elevation on survival due to differences in predation pressure or environmental conditions as well as potential indirect effects of elevation on BMR via temperature and the costs of thermoregulation. Higher BMR in tropical passerine birds predicted lower apparent survival, regardless of the elevation at which species occurred. In addition, elevation had a direct negative effect on apparent survival, perhaps due to harsher abiotic conditions, low site fidelity or both at high elevations. We provide evidence of a link between metabolic rate and longevity previously undescribed in populations of free‐living birds. Our results illustrate that tropical montane species may be characterized by a unique suite of traits in their pace of life, in which BMR does not differ from lowland birds, but survival does. A plain language summary is available for this article.

Non-native earthworms can drive ecosystem change, simplify native plant communities, and promote invasion by non-native plants, but we know little about their pathways into island archipelagos, which currently support about 40% of the worlds threatened species. We studied links among non-native earthworms, human settlement, deer, and plant communities on 26 islands in the San Juan and Southern Gulf Island archipelagos of the Georgia Basin Ecoregion of western North America. We evaluated the (1) invasion pathways and occurrence of non-native earthworms on islands, (2) influence of non-native earthworms on herbaceous and woody plant cover, and (3) potential for synergistic interactions among deer, earthworms and non-native plants. Human settlement was a pre-condition to detecting non-native earthworms on islands. Non-native earthworm abundance was related positively to the cover of non-native herbaceous and woody plants, effects which may be exacerbated by high deer density. Our findings suggest that the absence of non-native earthworms on many small islands makes their protection crucial to the conservation of intact examples of native ecosystems, including critically endangered Garry oak and maritime meadows in Canada.

A capacity for rapid adaptation should enhance the persistence of populations subject to temporal and spatial heterogeneity in natural selection, but examples from nature remain scarce.

Global declines in large-bodied terrestrial vertebrates are widely linked to human disturbance and habitat loss. Identifying remaining opportunities to conserve areas wherein such species may persist or might be reintroduced has thus become a key challenge in biodiversity conservation globally. We used multi-year camera trap data and spatial capture-recapture models to test how individually-identified Andean bears (Tremarctos ornatus) responded to roads and a suite of forest cover and topographic variables that may be indicative of food resources in critically-threatened, equatorial dry forest habitats of northern Peru. Bear density averaged 4 individuals per 100 km2 in our 668 km2 study area, but proximity to roads significantly reduced density, possibly because they are avoided and/or facilitate demographic sinks or human hunting. In contrast, the effects of forest cover and topographic variables on bear density varied seasonally, reflecting documented variation in mating behavior that may result from seasonal food resources. Despite large uncertainties in model predictions on the occurrence of bear habitat outside our study area, extrapolating our results suggests that a substantial area of high-quality habitat for Andean bears exists outside the recognized IUCN range, including equatorial dry forest habitats known to harbor many threatened species. Because Andean bears are vulnerable to extinction and are an established flagship species in much of South America, the expanded range and potential habitats that we identify here require urgent validation for possible inclusion in landscape-level plans to conserve equatorial dry forests of the Andes.

Biological invasions by cryptic ecosystem engineers can alter the ecological and socio‐economic values of ecosystems in ways that may take decades to detect. The invasion of North American glacial refuges by non‐native earthworms offers a prominent but understudied example of a cryptic invasion. Non‐native earthworms are known to alter carbon sequestration, disrupt mycorrhizal networks and homogenize plant communities, but natural resource managers still lack robust protocols to identify and safeguard high conservation value communities (HCVCs) from such invasions. In the absence of such protocols, or reliable methods to eradicate non‐native earthworm populations once established, there is an urgent need for methods to identify HCVCs at risk of or potentially shielded from earthworm invasion by the existence of abiotic barriers to their dispersal or persistence in such sites. We developed species distribution models (SDMs) using in situ field surveys and remotely sensed data to (1) identify factors limiting the occurrence of non‐native Lumbricus earthworms in imperilled coastal Douglas‐fir (Pseudotsuga menziesii) forest and Garry oak (Quercus garryana) and maritime meadow ecosystems endemic to glacial refugia of the Pacific Northwest of North America and to (2) evaluate their influence on plant species diversity and identify abiotic factors capable of preventing their dispersal or persistence at a site. As predicted, shallow, drier soils and steeper terrain limited earthworm occurrence and abundance in our highly heterogeneous study area, and earthworm presence appeared to reduce plant species richness. Our results indicated that HCVCs endemic to shallow‐soil (

Immigration into small recipient populations is expected to alleviate inbreeding and increase genetic variation, and hence facilitate population persistence through genetic and/or evolutionary rescue. Such expectations depend on three standard assumptions: that immigrants are outbred, unrelated to existing natives at arrival, and unrelated to each other. These assumptions are rarely explicitly verified, including in key field systems in evolutionary ecology. Yet, they could be violated due to non-random or repeated immigration from adjacent small populations. We combined molecular genetic marker data for 150-160 microsatellite loci with comprehensive pedigree data to test the three assumptions for a song sparrow (Melospiza melodia) population that is a model system for quantifying effects of inbreeding and immigration in the wild. Immigrants were less homozygous than existing natives on average, with mean homozygosity that closely resembled outbred natives. Immigrants can therefore be considered outbred on the focal population scale. Comparisons of homozygosity of real or hypothetical offspring of immigrant-native, native-native and immigrant-immigrant pairings implied that immigrants were typically unrelated to existing natives and to each other. Indeed, immigrants’ offspring would be even less homozygous than outbred individuals on the focal population scale. The three standard assumptions of population genetic and evolutionary theory were consequently largely validated. Yet, our analyses revealed some deviations that should be accounted for in future analyses of heterosis and inbreeding depression, implying that the three assumptions should be verified in other systems to probe patterns of non-random or repeated dispersal and facilitate precise and unbiased estimation of key evolutionary parameters.

Identifying causes and consequences of variation in species life history has the potential to improve predictions about how climate and land-use change may affect the demography and distribution of species in future. Sooty Fox Sparrows (Passerella unalaschcensis (J.F. Gmelin, 1789); or commonly grouped within the Fox Sparrow, Passerella iliaca (Merrem, 1786)) were migrants that rarely bred in the Georgia Basin of British Columbia prior to ∼1950 but have since established resident populations. Data on 270 color-banded birds and 54 nests on Mandarte Island, British Columbia, allowed us to estimate demographic vital rates and population growth in one recently established population. Annual fecundity (F), estimated as the product of the number of broods initiated (1.5 ± 0.01; mean ± SD), clutch size (2.82 ± 0.44), and probability of survival to fledging (0.68 ± 0.02), exceeded values reported for migrants, supporting the hypothesis that residents invest more in reproduction, on average, than migrants within species. Estimating juvenile and adult overwinter survival (Sj = 0.32 ± 0.06 and Sa = 0.69 ± 0.05) next allowed us to simulate an expected distribution of population growth rates as λexp = Sa + (Sj × F), given parameter error. Our estimate of λexp (1.61 ± 0.57) implies expeditious population growth, consistent with the species’ recent colonization of the region.

Factors that affect the survival of nestling birds can drive the dynamics of populations and the evolution of life histories. Such factors may include a variety of intrinsic, biotic, or abiotic factors that vary in magnitude or effect over ecological time scales. Consequently, estimates of the influence of such factors on nest success may differ when estimated from short- vs. long-term studies. We tested this possibility by estimating the influence of 2 intrinsic factors (female age and inbreeding coefficient), 2 abiotic factors (rainfall and temperature), and 3 biotic factors (breeding density, parasitism rate, and brood parasitism) previously found to predict success (≥1 fledged young) in a 39 yr study of a Song Sparrow (Melospiza melodia) population on Mandarte Island, British Columbia, Canada. We then compared those estimates to others obtained from parallel models based on the the first 13 yr, middle 13 yr, and last 13 yr of the study. Over the entire study, nest success increased in females...

Species invasions and range shifts can lead to novel competitive interactions between historically resident and colonizing species, but the demographic consequences of such interactions remain controversial. We present results from field experiments and 45 years of demographic monitoring to test the hypothesis that the colonization of Mandarte Is., BC, Canada, by fox sparrows (Passerella iliaca) caused the long-term decline of the resident population of song sparrows (Melospiza melodia). Several lines of evidence indicate that competition with fox sparrows for winter food reduced over-winter survival in juvenile song sparrows by 48% from 1960 to 2015, enforcing population decline despite an increase in annual reproductive rate in song sparrows over the same period. Preference for locally abundant seeds presented at experimental arenas suggested complete overlap in diet in song and fox sparrows, and observations at arenas baited with commercial seed showed that fox sparrows displaced song sparrows in 91-100% of interactions in two periods during winter. In contrast, we found no evidence of interspecific competition for resources during the breeding season. Our results indicate that in the absence of marked shifts in niche dimension, range expansions by dominant competitors have the potential to cause the extirpation of historically resident species when competitive interactions between them are strong and resources not equitably partitioned.

Individual animal fitness can be strongly influenced by the ability to recognise habitat features which may be beneficial. Many studies focus on the effects of habitat on annual reproductive rate, even though adult survival is typically a greater influence on fitness and population growth in vertebrate species with intermediate to long lifespans. Understanding the effects of preferred habitat on individuals over the annual cycle is therefore necessary to predict its influences on individual fitness. This is particularly true in species that are resident and territorial year-round in the temperate zone, which may face potential trade-offs between habitat that maximises reproduction and that which maximises non-breeding season (‘over-winter’) survival. We used a 37-year study of Song Sparrows Melospiza melodia residing territorially year-round on a small island to examine what habitat features influenced adult over-winter survival, how site-specific variation in adult survival versus annual reproductive rate influenced long-term habitat preference, and if preferred sites conferred on average higher individual fitness. Habitat features such as area of shrub cover and exposure to intertidal coastline predicted adult over-winter survival independent of individual age or sex, population size, or winter weather. Long-term habitat preference (measured as occupation rate) was better predicted by site-specific annual reproductive rate than by expected over-winter survival, but preferred sites maximised fitness on average over the entire annual cycle,. Although adult over-winter survival had a greater influence on population growth (λ) than reproductive rate, the influence of reproductive rate on λ increased in preferred sites because site-specific variation in reproductive rate was higher than variation in expected over-winter survival. Because preferred habitats tended to have higher mean site-specific reproductive and adult survival rates, territorial birds in this population do not appear to experience seasonal trade-offs in preferred habitat, but are predicted to incur substantial fitness costs of settling in less-preferred sites. This article is protected by copyright. All rights reserved.

Theory predicts that reproduction will change as individuals near the end of their lives by either increasing reproductive allocation (terminal allocation hypothesis) or decreasing allocation (senescence hypothesis) toward the end of life. Although senescence has received more support, few studies examine how both age and years to death influence late-life reproduction. We used a 37-yr study of Song Sparrows (Melospiza melodia) to ask how age and years to death influenced reproductive allocation late in life. We observed both senescence and terminal allocation, and that age and years to death interacted to influence individual variation in allocation tactics. In particular, we observed a decline in allocation in older individuals (senescence), but only when comparing individuals with the same number of years to death. Likewise, we observed terminal allocation, but only in young females. Reproductive tactics were most variable in young females, with many exerting high effort but living 1-2 yr, and others exerting low effort annually and living longer. Our results suggest that late-life reproductive tactics are influenced by both chronological age and years to death, and that short- and long-lived females with differing reproductive tactics exist within this population of Song Sparrows.

Many studies report mixed results on the influence of invasive plants on native animals, partly due to uncertainties about habitat preference and reproductive performance in native animals before and after plant invasion. We used vegetation surveys 20 years apart and 18 years of breeding data from Song Sparrows (Melospiza melodia (A. Wilson, 1810)) to compare nest-site preference and reproductive performance during the colonization of Mandarte Island, British Columbia, by one shrub species native to the region but historically absent from the island (red elderberry, Sambucus racemosa L.) and another species that is exotic to North America (Himalayan blackberry, Rubus armeniacus Focke = Rubus bifrons Vest). Nest-site preference declined where red elderberry increased but was unrelated to change in the cover of Himalayan blackberry. Song Sparrows nested in trailing blackberry (Rubus ursinus Cham. and Schltdl.) and its exotic congener Himalayan blackberry in preference to two common shrubs native to Mandarte Island (Nootka rose, Rosa nutkana C. Presl; snowberry, Symphoricarpos albus (L.) S.F. Blake) and built just 1 of 1051 nests in red elderberry. In contrast, reproductive performance was similar in all shrub species used regularly as nest substrates. Our results show that Song Sparrow nest-site preference and reproductive performance were independent of plant species origin.

Inclusive fitness theory predicts that parental care will vary with relatedness between potentially caring parents and offspring, potentially shaping mating system evolution. Systems with extra-pair paternity (EPP), and hence variable parent–brood relatedness, provide valuable opportunities to test this prediction. However, existing theoretical and empirical studies assume that a focal male is either an offspring's father with no inbreeding, or is completely unrelated. We highlight that this simple dichotomy does not hold given reproductive interactions among relatives, complicating the effect of EPP on parent–brood relatedness yet providing new opportunities to test inclusive fitness theory. Accordingly, we tested hierarchical hypotheses relating parental feeding rate to parent–brood relatedness, parent kinship and inbreeding, using song sparrows ( Melospiza melodia ) experiencing natural variation in relatedness. As predicted, male and female feeding rates increased with relatedness to a dependent brood, even controlling for brood size. Male feeding rate tended to decrease as paternity loss increased, and increased with increasing kinship and hence inbreeding between socially paired mates. We thereby demonstrate that variation in a key component of parental care concurs with subtle predictions from inclusive fitness theory. We additionally highlight that such effects can depend on the underlying social mating system, potentially generating status-specific costs of extra-pair reproduction.

Spatial variation in the occurrence of browsing ungulates can drive local adaptation in plant traits but also lead to trade-offs among traits potentially enhancing competitive ability versus resistance or tolerance to browsing. Plectritis congesta populations co-occurring on islands with and without ungulates offer striking examples of population-level variation in traits, such as plant height and fruit morphology, which may also affect fitness. We monitored split-plot common gardens exposed to and protected from browsing ungulates for 5 yr to test for local adaptation (local vs. foreign comparison) in P. congesta by comparing the survival and fecundity of 4,392 sown fruits from six island populations where ungulates were present ("historically exposed") and six where they were absent ("historically naive"). Our results indicate that local adaptation to browsing in P. congesta favored rosette formation, delayed flowering, reduced height, and the production of wingless fruits, all of which appeared to enhance survival, fecundity, and population growth in plants from populations historically exposed to ungulate browsers, as compared to plants from historically naive populations. In contrast, plants from historically naive populations displayed higher relative fitness in the absence of ungulates, increased in height, flowered earlier, and produced fewer but larger, winged fruits, often in large terminal inflorescences. Our results support the hypothesis that variation in the occurrence of ungulate browsers has led to (1) spatial heterogeneity in natural selection and rapid adaptation in P. congesta populations on islands, and (2) context-dependent trade-offs in the fitness value of traits linked to the resistance or tolerance of browsing versus success in competition for light, pollinators, or other resources. Because patterns of selection in plant communities will vary with the introduction or extirpation of top predators or browsers, we suggest historical context, local adaptation, and the capacity for rapid adaptation should be a focal concern of those aiming to maximize or predict population persistence under environmental change in conservation plans.

Individual variation in nest defense behaviour is common in altricial birds, but despite clear predictions about why such variation exists, there is no consensus on its causes. We tested for an influence of five predictors of individual variation in nest defense behaviour, including time of season, offspring age, parental age and sex, and clutch size in a well-studied Song Sparrow (Melospiza melodia (A. Wilson, 1810)) population. We recorded parental responses to a standardized human approach and used model selection to assess support for each predictor. Parents tended to approach observers less closely and alarm-call less as the breeding season progressed, indicating a modest seasonal decline in parental nest defense, which is consistent with the hypothesis that the reproductive value of offspring influenced parental defense behaviour. Clutch size effect estimates were insignificant, but it was weakly supported as a predictor of nest defense, which is expected if parental investment in defense and current reproductive effort are positively related. Offspring age, as well as parental age and sex, received little or no support as affecting parent nest defense. Our results offer limited support for the hypothesis that the reproductive value of offspring affects parental nest defense.

Female extra-pair reproduction in socially monogamous systems is predicted to cause cuckolded socially-paired males to conditionally reduce paternal care, causing selection against extra-pair reproduction and underlying polyandry. However, existing models and empirical studies have not explicitly considered that cuckolded males might be related to their socially-paired female and/or to her extra-pair mate, and therefore be related to extra-pair offspring that they did not sire but could rear. Selection against paternal care, and hence against extra-pair reproduction, might then be weakened. We derive metrics that quantify allele-sharing between within-pair and extra-pair offspring and their mother and her socially-paired male in terms of coefficients of kinship and inbreeding. We use song sparrow (Melospiza melodia) paternity and pedigree data to quantify these metrics, and thereby quantify the joint effects of extra-pair reproduction and inbreeding on a brood's total allelic value to its socially-paired parents. Cuckolded male song sparrows were almost always detectably related to extra-pair offspring they reared. Consequently, although brood allelic value decreased substantially following female extra-pair reproduction, this decrease was reduced by within-pair and extra-pair reproduction among relatives. Such complex variation in kinship within nuclear families should be incorporated into models considering coevolutionary dynamics of extra-pair reproduction, parental care, and inbreeding.