Negative Fitness Consequences Research Articles

Spatial variation in population genomic responses to over a century of anthropogenic change within a tidal marsh songbird.

Combating the current biodiversity crisis requires the accurate documentation of population responses to human-induced ecological change. However, our ability to pinpoint population responses to human activities is often limited to the analysis of populations studied well after the fact. Museum collections preserve a record of population responses to anthropogenic change that can provide critical baseline data on patterns of genetic diversity, connectivity, and population structure prior to the onset of human perturbation. Here, we leverage a spatially replicated time series of specimens to document population genomic responses to the destruction of nearly 90% of coastal habitats occupied by the Savannah sparrow (Passerculus sandwichensis) in California. We sequenced 219 sparrows collected from 1889 to 2017 across the state of California using an exome capture approach. Spatial-temporal analyses of genetic diversity found that the amount of habitat lost was not predictive of genetic diversity loss. Sparrow populations from southern California historically exhibited lower levels of genetic diversity and experienced the most significant temporal declines in genetic diversity. Despite experiencing the greatest levels of habitat loss, we found that genetic diversity in the San Francisco Bay area remained relatively high. This was potentially related to an observed increase in gene flow into the Bay Area from other populations. While gene flow may have minimized genetic diversity declines, we also found that immigration from inland freshwater-adapted populations into tidal marsh populations led to the erosion of divergence at loci associated with tidal marsh adaptation. Shifting patterns of gene flow through time in response to habitat loss may thus contribute to negative fitness consequences and outbreeding depression. Together, our results underscore the importance of tracing the genomic trajectories of multiple populations over time to address issues of fundamental conservation concern.

Consistent traffic noise impacts few fitness-related traits in a field cricket

BackgroundAnthropogenic habitat change is occurring rapidly, and organisms can respond through within-generation responses that improve the match between their phenotype and the novel conditions they encounter. But, plastic responses can be adaptive or maladaptive and are most likely to be adaptive only when contemporary conditions reasonably mimic something experienced historically to which a response has already evolved. Noise pollution is a ubiquitous anthropogenic stressor that accompanies expanding urbanization. We tested whether the amplitude of traffic noise influences a suite of fitness-related traits (e.g. survival, life history, reproductive investment, immunity) and whether that depends on the life stage at which the noise is experienced (juvenile or adult). Our treatments mimic the conditions experienced by animals living in urban roadside environments with variable vehicle types, but continuous movement of traffic. We used the Pacific field cricket, an acoustically communicating insect that was previously shown to experience some negative behavioral and life history responses to very loud, variable traffic noise, as a model system.ResultsAfter exposing crickets to one of four traffic noise levels (silence, 50dBA, 60dBA, and 70dBA which are commonly experienced in their natural environment) during development, at adulthood, or both, we measured a comprehensive suite of fifteen fitness-related traits. We found that survival to adulthood was lower under some noise treatments than under silence, and that the number of live offspring hatched depended on the interaction between a female’s juvenile and adult exposure to traffic noise. Both of these suggest that our noise treatments were indeed a stressor. However, we found no evidence of negative or positive fitness effects of noise on the other thirteen measured traits.ConclusionsOur results suggest that, in contrast to previous work with loud, variable traffic noise, when noise exposure is relatively constant, plasticity may be sufficient to buffer many negative fitness effects and/or animals may be able to habituate to these conditions, regardless of amplitude. Our work highlights the importance of understanding how the particular characteristics of noise experienced by animals influence their biological responses and provides insight into how commensal animals thrive in human-dominated habitats.

Fitness effects of assisted gene flow in an endangered salmonid population

AbstractAssisted gene flow is increasingly used to combat severe population declines. However, the associated risks, such as outbreeding depression, are often insufficiently assessed. Here, we studied the impact of assisted gene flow on the fitness of a highly endangered landlocked salmon population (Salmo salar m. sebago) from the lake Saimaa complex (Finland), using an anadromous Atlantic salmon population (S. salar) as donor. We released individuals of both parental populations and their hybrids into seminatural streams, monitoring their survival under predation risk from Northern pike (Esox lucius) and their growth on a natural diet. Before release, we exposed half of the salmon to the parasite Diplostomum pseudospathaceum to study whether assisted gene flow affects infection susceptibility, which could indirectly shape predation susceptibility and growth. The parental populations differed in both studied traits and the hybrids showed intermediate values. Relative to the target landlocked salmon population, hybrids experienced 21–26% lower survival, but 1.4–2.2% higher growth. They also carried 0.6–2.8 more parasites than the landlocked salmon, contributing to survival differences. These findings indicate that assisted gene flow can induce both negative and positive fitness effects. We propose that the overall net effects of this conservation tool need to be carefully evaluated before its implementation.

Multiple mechanisms for inbreeding avoidance used simultaneously in a wild ape.

Mating with close kin can have considerable negative fitness consequences, which are expected to result in selective pressure for inbreeding avoidance mechanisms, such as dispersal, mate choice and post-copulatory biases. Captive studies have suggested that inbreeding avoidance through mate choice is far less widespread than expected and may be absent where other mechanisms already limit inbreeding. However, few studies have examined multiple mechanisms of inbreeding avoidance simultaneously, particularly in the wild. We use 13 years of detailed dispersal, copulation and paternity data from mountain gorillas to examine inbreeding avoidance. We find that partial dispersal of both sexes results in high kinship in multimale groups, but that copulations between close kin occur 40% less than expected. We find strong kin discrimination in mate choice, with significant avoidance of maternal kin but more limited avoidance of paternal kin. We find no evidence for post-copulatory inbreeding avoidance. Our analyses support familiarity-based mechanisms of kin identification and age-based avoidance that limits mating between fathers and daughters in their natal group. Our findings demonstrate that multiple complementary mechanisms for inbreeding avoidance can evolve in a single species and suggest that inbreeding avoidance through mate choice may enable more flexible dispersal systems to evolve.

Improving the science and practice of using artificial roosts for bats.

Worldwide, artificial bat roosts (e.g., bat boxes, bark mimics, bat condos) are routinely deployed for conservation, mitigation, and community engagement. However, scant attention has been paid to developing best practices for the use of artificial roosts as conservation tools. Although bats readily occupy artificial roosts, occupancy and abundance data are misleading indicators of habitat quality. Lacking information on bat behavior, health, and fitness in artificial roosts, their conservation efficacy cannot be adequately validated. We considered the proximal and ultimate factors, such as evolutionarily reliable cues, that may prompt bats to preferentially use and show fidelity to suboptimal artificial roosts even when high-quality alternatives are available. Possible negative health and fitness consequences for artificial roost inhabitants include exposure to unstable and extreme microclimates in poorly designed roosts, and vulnerability to larger numbers of ectoparasites in longer lasting artificial roosts that house larger bat colonies than in natural roosts. Bats using artificial roosts may have lower survival rates if predators have easy access to roosts placed in conspicuous locations. Bats may be lured into occupying low-quality habitats if attractive artificial roosts are deployed on polluted urban and agricultural landscapes. To advance the science behind artificial bat roosts, we present testable research hypotheses and suggestions to improve the quality of artificial roosts for bats and decrease risks to occupants. Because continued loss of natural roosts may increase reliance on alternatives, such as artificial roosts, it is imperative that this conservation practice be improved.

Bacterial-fungal interactions promote parallel evolution of global transcriptional regulators in a widespread Staphylococcus species.

Experimental studies of microbial evolution have largely focused on monocultures of model organisms, but most microbes live in communities where interactions with other species may impact rates and modes of evolution. Using the cheese rind model microbial community, we determined how species interactions shape the evolution of the widespread food- and animal-associated bacterium Staphylococcus xylosus. We evolved S. xylosus for 450 generations alone or in co-culture with one of three microbes: the yeast Debaryomyces hansenii, the bacterium Brevibacterium aurantiacum, and the mold Penicillium solitum. We used the frequency of colony morphology mutants (pigment and colony texture phenotypes) and whole-genome sequencing of isolates to quantify phenotypic and genomic evolution. The yeast D. hansenii strongly promoted diversification of S. xylosus. By the end of the experiment, all populations co-cultured with the yeast were dominated by pigment and colony morphology mutant phenotypes. Populations of S. xylosus grown alone, with B. aurantiacum, or with P. solitum did not evolve novel phenotypic diversity. Whole-genome sequencing of individual mutant isolates across all four treatments identified numerous unique mutations in the operons for the SigB, Agr, and WalRK global regulators, but only in the D. hansenii treatment. Phenotyping and RNA-seq experiments highlighted altered pigment and biofilm production, spreading, stress tolerance, and metabolism ofS. xylosus mutants.Fitness experiments revealed antagonistic pleiotropy, where beneficial mutations that evolved in the presence of the yeast had strong negative fitness effects in other biotic environments. This work demonstrates that bacterial-fungal interactions can have long-term evolutionary consequences within multispecies microbiomes by facilitating the evolution of strain diversity.

Stop, then go! Rapid acceleration offsets the costs of intermittent locomotion when turning in Florida scrub lizards.

Intermittent locomotion is a common locomotor mode in small vertebrates. Pausing is thought to aid in locating a predator or prey, enhancing crypsis, lowering energy costs, and/or maneuvering around obstacles or toward a refuge. Many lizards flee predators by turning into potential refugia and subsequently pausing, presumably to conceal themselves. Intermittent locomotion may be associated with turning by allowing an animal time to assess its surroundings and/or decreasing the likelihood of losing its footing. In this study, we quantify locomotor performance and the use of intermittent locomotion in Florida scrub lizards (Sceloporus woodi) when navigating either a 45° or 90° turn. Lizards paused in 92.91% of all trials, and yet despite pausing, instantaneous speed was not different entering or exiting the turn. This result suggests that turning comes at minimal cost to forward speed for lizards under these conditions. Pausing during a turn, however, did slow speed in the turn. Interestingly, the speed in the turn did not differ in trials with a pause before the turn versus trials without a pause. The angle of the turn also had no effect on whether lizards paused. We found that lizards increase peak acceleration following pauses to compensate for lost speed during the pause, providing a mechanism that may minimize negative fitness effects associated with slow running speeds and allow intermittent locomotion to be such a common strategy in lizards.

Evolutionary barriers to horizontal gene transfer in macrophage-associated Salmonella.

Horizontal gene transfer (HGT) is a powerful evolutionary force facilitating bacterial adaptation and emergence of novel phenotypes. Several factors, including environmental ones, are predicted to restrict HGT, but we lack systematic and experimental data supporting these predictions. Here, we address this gap by measuring the relative fitness of 44 genes horizontally transferred from Escherichia coli to Salmonella enterica in infection-relevant environments. We estimated the distribution of fitness effects in each environment and identified that dosage-dependent effects across different environments are a significant barrier to HGT. The majority of genes were found to be deleterious. We also found longer genes had stronger negative fitness consequences than shorter ones, showing that gene length was negatively associated with HGT. Furthermore, fitness effects of transferred genes were found to be environmentally dependent. In summary, a substantial fraction of transferred genes had a significant fitness cost on the recipient, with both gene characteristics and the environment acting as evolutionary barriers to HGT.

Demographic history and genomic consequences of 10,000 generations of isolation in a wild mammal

Increased human activities caused the isolation of populations in many species-often associated with genetic depletion and negative fitness effects. The effects of isolation are predicted by theory, but long-term data from natural populations are scarce. We show, with full genome sequences, that common voles (Microtus arvalis) in the Orkney archipelago have remained genetically isolated from conspecifics in continental Europe since their introduction by humans over 5,000 years ago. Modern Orkney vole populations are genetically highly differentiated from continental conspecifics as a result of genetic drift processes. Colonization likely started on the biggest Orkney island and vole populations on smaller islands were gradually split off, without signs of secondary admixture. Despite having large modern population sizes, Orkney voles are genetically depauperate and successive introductions to smaller islands resulted in further reduction of genetic diversity. We detected high levels of fixation of predicted deleterious variation compared with continental populations, particularly on smaller islands, yet the fitness effects realized in nature are unknown. Simulations showed that predominantly mildly deleterious mutations were fixed in populations, while highly deleterious mutations were purged early in the history of the Orkney population. Relaxation of selection overall due to benign environmental conditions on the islands and the effects of soft selection may have contributed to the repeated, successful establishment of Orkney voles despite potential fitness loss. Furthermore, the specific life history of these small mammals, resulting in relatively large population sizes, has probably been important for their long-term persistence in full isolation.

Is now the time? Review of genetic rescue as a conservation tool for brook trout.

Brook trout populations have been declining throughout their native range in the east coast of the United States. Many populations are now distributed in small, isolated habitat patches where low genetic diversity and high rates of inbreeding reduce contemporary viability and long-term adaptive potential. Although human-assisted gene flow could theoretically improve conservation outcomes through genetic rescue, there is widespread hesitancy to use this tool to support brook trout conservation. Here, we review the major uncertainties that have limited genetic rescue from being considered as a viable conservation tool for isolated brook trout populations and compare the risks of genetic rescue with other management alternatives. Drawing on theoretical and empirical studies, we discuss methods for implementing genetic rescue in brook trout that could yield long-term evolutionary benefits while avoiding negative fitness effects associated with outbreeding depression and the spread of maladapted alleles. We also highlight the potential for future collaborative efforts to accelerate our understanding of genetic rescue as a viable tool for conservation. Ultimately, while we acknowledge that genetic rescue is not without risk, we emphasize the merits that this tool offers for protecting and propagating adaptive potential and improving species' resilience to rapid environmental change.

Unpredictable fasting transiently alters corticosterone and feeding behavior but not body mass or later HPA axis function in zebra finches (Taeniopygia guttata)

The hypothalamic-pituitary-adrenal (HPA) axis is commonly activated in response to unpredictable conditions, including unstable or inadequate food supply. Extended exposure to unpredictable food resources can alter HPA axis function, with the potential for negative fitness consequences. We addressed the interrelationships of unpredictable food resources, HPA axis activity, and food intake in adult and juvenile zebra finches (Taeniopygia guttata). Finches exposed to prolonged periods of unpredictable food elevate corticosterone (the primary avian glucocorticoid) when food is unavailable; however, whether they experience chronic elevation in baseline corticosterone during periods of unpredictability, even when food is available, is unclear. We subjected adults and juveniles to an extended period of temporally unpredictable food (consisting of a random daily fast) or predictable food supply. We investigated baseline corticosterone under fed conditions and in response to an acute fast (mimicking the daily fasting periods in the unpredictable treatment), and assessed differences in body mass, food intake, and corticosterone responses to restraint. Regardless of sex and age, individuals in both treatment groups elevated corticosterone when fasted, and baseline corticosterone under fed conditions was indistinguishable between groups. Thus, corticosterone levels were not persistently elevated in the unpredictably fed group. Treatment groups did not differ in body mass or corticosterone responses to restraint, but unpredictably fed birds consumed food more rapidly when food was available. Our findings suggest that the unpredictably fed birds experienced repeated, moderate elevations in corticosterone. Such elevations may aid birds in coping with unpredictable food sources, in part by activating compensatory changes in foraging behavior.

The terrestrial isopod symbiont ‘Candidatus Hepatincola porcellionum’ is a potential nutrient scavenger related to Holosporales symbionts of protists

The order Holosporales (Alphaproteobacteria) encompasses obligate intracellular bacterial symbionts of diverse Eukaryotes. These bacteria have highly streamlined genomes and can have negative fitness effects on the host. Herein, we present a comparative analysis of the first genome sequences of ‘Ca. Hepatincola porcellionum’, a facultative symbiont occurring extracellularly in the midgut glands of terrestrial isopods. Using a combination of long-read and short-read sequencing, we obtained the complete circular genomes of two Hepatincola strains and an additional metagenome-assembled draft genome. Phylogenomic analysis validated its phylogenetic position as an early-branching family-level clade relative to all other established Holosporales families associated with protists. A 16S rRNA gene survey revealed that this new family encompasses diverse bacteria associated with both marine and terrestrial host species, which expands the host range of Holosporales bacteria from protists to several phyla of the Ecdysozoa (Arthropoda and Priapulida). Hepatincola has a highly streamlined genome with reduced metabolic and biosynthetic capacities as well as a large repertoire of transmembrane transporters. This suggests that this symbiont is rather a nutrient scavenger than a nutrient provider for the host, likely benefitting from a nutrient-rich environment to import all necessary metabolites and precursors. Hepatincola further possesses a different set of bacterial secretion systems compared to protist-associated Holosporales, suggesting different host-symbiont interactions depending on the host organism.

Herbivory selects for tolerance and constitutive defence across stages of community succession.

Plants defend themselves from herbivory by either reducing damage (resistance) or minimizing its negative fitness effects with compensatory growth (tolerance). Herbivore pressure can fluctuate from year to year in an early secondary successional community, which can create temporal variation in selection for defence traits. We manipulated insect herbivory and successional age of the community as agents of natural selection in replicated common gardens with the perennial herb Solidago altissima. In these genotypic selection experiments, herbivory consistently selected for better defended plants in both successional communities. Herbivore suppression increased plant survival and the probability of flowering only in mid-succession. Despite these substantial differences in the effects of herbivory between early and mid-succession, the selection on defence traits did not change. Succession affected selection only on aboveground biomass, with positive selection in early but not mid-succession, suggesting an important role of competition in the selective environment. These results demonstrate that changes in the community that affect key life-history traits in an individual species can occur over very short timescales in a dynamic secondary successional environment. The resulting community context-driven variation in natural selection may be an important, yet overlooked, contributor to adaptive mosaics across populations.

High-density linkage maps and chromosome level genome assemblies unveil direction and frequency of extensive structural rearrangements in wood white butterflies (Leptidea spp.)

Karyotypes are generally conserved between closely related species and large chromosome rearrangements typically have negative fitness consequences in heterozygotes, potentially driving speciation. In the order Lepidoptera, most investigated species have the ancestral karyotype and gene synteny is often conserved across deep divergence, although examples of extensive genome reshuffling have recently been demonstrated. The genus Leptidea has an unusual level of chromosome variation and rearranged sex chromosomes, but the extent of restructuring across the rest of the genome is so far unknown. To explore the genomes of the wood white (Leptidea) species complex, we generated eight genome assemblies using a combination of 10X linked reads and HiC data, and improved them using linkage maps for two populations of the common wood white (L. sinapis) with distinct karyotypes. Synteny analysis revealed an extensive amount of rearrangements, both compared to the ancestral karyotype and between the Leptidea species, where only one of the three Z chromosomes was conserved across all comparisons. Most restructuring was explained by fissions and fusions, while translocations appear relatively rare. We further detected several examples of segregating rearrangement polymorphisms supporting a highly dynamic genome evolution in this clade. Fusion breakpoints were enriched for LINEs and LTR elements, which suggests that ectopic recombination might be an important driver in the formation of new chromosomes. Our results show that chromosome count alone may conceal the extent of genome restructuring and we propose that the amount of genome evolution in Lepidoptera might still be underestimated due to lack of taxonomic sampling.

Ontogenetic effects of brood parasitism by the Brown-headed Cowbird on host offspring.

Nest-sharer avian brood parasites do not evict or otherwise kill host chicks, but instead inflict a range of negative effects on their nestmates that are mediated by interactions between the parasite and host life history traits. Although many of the negative fitness effects of avian brood parasitism are well documented across diverse host species, there remains a paucity of studies that have examined the impacts of parasitism across the entirety of host ontogeny (i.e., from when an egg is laid until independence). More specifically, few studies have examined the impact of brood parasitism on the pre- and post-fledging development, physiology, behavior, and survival of host offspring. To help fill this knowledge gap, we assessed the effects of brood parasitism by Brown-headed Cowbirds (Molothrus ater) across the ontogeny (incubation, nestling, and post-fledging period) of nine sympatrically breeding host species in central Illinois, USA; due to sample sizes, impacts on the post-fledging period were only examined in two of the nine species. Specifically, we examined the impact of brood parasitism on ontogenetic markers including the embryonic heart rate, hatching rate, nestling period length, nest survival, and offspring growth and development. Additionally, in species in which we found negative impacts of cowbird parasitism on host nestmate ontogeny, we examined whether the difference in adult size between parasites and their hosts and their hatching asynchrony positively predicted variation in host costs across these focal taxa. We found that costs of cowbird parasitism were most severe during early nesting stages (reduction in the host clutch or brood size) and were predicted negatively by host size and positively by incubation length. In contrast, we only found limited costs of cowbird parasitism on other stages of host ontogeny; critically, post-fledging survival did not differ between host offspring that fledged alongside cowbirds and those that did not. Our findings (i) highlight the direct costs of cowbird parasitism on host fitness, (ii) provide evidence for when (the stage) those costs are manifested, and (iii) may help to explain why many anti-cowbird defenses of hosts have evolved for protection from parasitism during the laying and incubation stages.

Contrasting effects of an extended fall period and winter heatwaves on the overwintering fitness of diapausing disease vector, Aedes albopictus

Climate change is expected to dramatically alter autumnal and winter conditions in many temperate regions. However, limited data is available to accurately predict how these changes will impact species’ overwinter survival and post-winter fitness. Here, we determine how a longer, warmer fall period and winter heatwaves affect overwintering fitness and post-winter performance of the invasive mosquito vector, Aedes albopictus. We found that a longer, warmer fall period representative of early entry into diapause did not affect overwinter survival but did lead to reduced post-winter performance for multiple traits. Specifically, larvae that experienced longer, warmer fall conditions as diapause embryos exhibited reduced post-diapause larval starvation tolerance, increased post-diapause larval mortality, and longer post-diapause larval development compared to individuals from the short-fall treatments. These negative post-diapause fitness effects likely resulted from the greater energetic demands and/or damage incurred during the warmer, longer fall period. In contrast, exposure to winter heatwaves increased overwinter survival, possibly by allowing diapausing embryos to escape or repair cold injury. Finally, fall treatment and winter heatwaves had an interactive effect on male development time, while neither treatment impacted pupal mass in either sex. Overall, our results highlight that experiments that fail to measure post-diapause fitness are likely to substantially under-estimate the impacts of climate change on post-winter performance. Additionally, our results emphasize that it is crucial to consider the potentially conflicting effects of different aspects of climate change on a species’ overall overwintering success.

Heightened heart rate but similar flight responses to evolved versus recent predators in an Arctic seabird

Predator-prey dynamics in the Arctic are being altered with changing sea ice phenology. The increasing frequency of predation on colonial nesting seabirds and their eggs by the polar bear (<em>Ursus maritimus</em>) is a consequence of bears shifting to terrestrial food resources through a shortened seal-hunting season. We examined antipredator responses in a colony of nesting Common Eiders (<em>Somateria mollissima</em>) on East Bay Island, Nunavut, Canada, which is exposed to established nest predators, such as arctic fox (<em>Vulpes lagopus</em>), but also to recent increases in polar bear nest predation due to the bears’ lost on-ice hunting opportunities. Given eiders’ limited eco-evolutionary experience with bears, we aimed to experimentally contrast eider responses to the recent predation pressure by polar bears to those induced by their more traditional mammalian predator, the arctic fox. Our goal was to characterize whether this population of eiders was vulnerable to a changing predator regime. Using simulated approaches of visual stimuli of both predator types, we measured eider heart rate and flight initiation distance as physiological and behavioral metrics, respectively, to characterize the perceived risk of and subsequent response to imminent threat posed by these two predators that differ in historical encounter rates. Eider heart rates were more responsive to impending visual cues of arctic foxes compared to polar bears, but birds responded behaviorally to all simulated threats with similar flight initiation distances. Results suggest eiders may not perceive the full risk that bears pose as egg and adult predators, and are therefore expected to suffer negative fitness consequences from this ongoing and increasing interaction. Eiders may therefore require conservation intervention to aid in their management.

Fitness Costs of Female Competition Linked to Resource Defense and Relatedness of Competitors.

AbstractFemale reproductive success is often limited by access to resources, and this can lead to social competition both within and between kin groups. Theory predicts that both resource availability and relatedness should influence the fitness consequences of social competition. However, testing key predictions requires differentiating the effects of these two factors. Here, we achieve this experimentally by manipulating the social environment of house mice, a facultative communal breeding species with known kin discrimination ability. This allows us to investigate (1) the reproductive costs of defending a limited resource in response to cues of social competition and (2) whether such costs, or their potential mitigation via cooperative behavior, are influenced by the relatedness of competitors. Our results support the hypothesis that resource defense can be costly for females, potentially trading off against maternal investment. When the availability of protected nest sites was limited, subjects (1) were more active, (2) responded more strongly to simulated territory intrusions via competitive signaling, and (3) produced smaller weaned offspring. However, we found no evidence that the propensity for kin to cooperate was influenced by the relatedness of rivals. Communal breeding between sisters occurred independently of the relatedness of competitors and communally breeding sisters weaned fewer offspring when competing with unrelated females, despite our study being designed to prevent infanticide between kin groups. Our findings thus demonstrate that female competition has fitness costs and that associating with kin is beneficial to avoid negative fitness consequences of competing with nonkin, in addition to more widely recognized kin-selected benefits.

From high masked to high realized genetic load in inbred Scandinavian wolves.

When new mutations arise at functional sites they are more likely to impair than improve fitness. If not removed by purifying selection, such deleterious mutations will generate a genetic load that can have negative fitness effects in small populations and increase the risk of extinction. This is relevant for the highly inbred Scandinavian wolf (Canis lupus) population, founded by only three wolves in the 1980s and suffering from inbreeding depression. We used functional annotation and evolutionary conservation scores to study deleterious variation in a total of 209 genomes from both the Scandinavian and neighbouring wolf populations in northern Europe. The masked load (deleterious mutations in heterozygote state) was highest in Russia and Finland with deleterious alleles segregating at lower frequency than neutral variation. Genetic drift in the Scandinavian population led to the loss of ancestral alleles, fixation of deleterious variants and a significant increase in the per-individual realized load (deleterious mutations in homozygote state; an increase by 45% in protein-coding genes) over five generations of inbreeding. Arrival of immigrants gave a temporary genetic rescue effect with ancestral alleles re-entering the population and thereby shifting deleterious alleles from homozygous into heterozygote genotypes. However, in the absence of permanent connectivity to Finnish and Russian populations, inbreeding has then again led to the exposure of deleterious mutations. These observations provide genome-wide insight into the magnitude of genetic load and genetic rescue at the molecular level, and in relation to population history. They emphasize the importance of securing gene flow in the management of endangered populations.

Among-years rain variation is associated with flower size, but not with signal patch size in Iris petrana.

Among-years rain variation is associated with flower size, but not with signal patch size in Iris petrana.