Determinants and consequences of age of primiparity in bighorn ewes

Because variation in age of first reproduction can have major effects on individual fitness and population dynamics, it is important to understand what maintains that variability. Although early primiparity is assumed to be costly, it is sometimes associated with high lifetime reproductive success. We used a long‐term study on bighorn sheep Ovis canadensis to determine what variables affect age at first reproduction, investigate the impact of primiparity on body resources and quantify the reproductive performance of primiparous ewes. We then examined the consequences of delayed primiparity on adult body mass, longevity and lifetime reproductive success. Environmental conditions during early development, body mass as a yearling, genotype and maternal effects affected age of primiparity. Primiparous ewes lost more mass in winter and gained less mass in summer than multiparous ewes. Small yearling ewes that postponed reproduction attained similar adult mass than heavy yearling ewes who reproduced at a younger age. Early primiparity did not reduce longevity and was positively associated with lifetime reproductive success. Starting to reproduce as soon as possible appears to maximize fitness of females. When early life conditions are unfavorable, however, delayed primiparity allows greater body growth and likely maximizes survival. The combination of a conservative reproductive strategy and maternal effects on age of primiparity may partly delay population recovery following density‐dependent declines.     

A negative association between horn length and survival in a weakly dimorphic ungulate

While all models of sexual selection assume that the development and expression of enlarged secondary sexual traits are costly, males with larger ornaments or weapons generally show greater survival or longevity. These studies have mostly been performed in species with high sexual size dimorphism, subject to intense sexual selection. Here, we examined the relationships between horn growth and several survival metrics in the weakly dimorphic Pyrenean chamois (Rupicapra pyrenaica). In this unhunted population living at high density, males and females were able to grow long horns without any apparent costs in terms of longevity. However, we found a negative relationship between horn growth and survival during prime age in males. This association reduces the potential evolutionary consequences of trophy hunting in male chamois. We also found that females with long horns tended to have lower survival at old ages. Our results illustrate the contrasting conclusions that may be drawn when different survival metrics are used in analyses. The ability to detect trade‐off between the expression of male secondary sexual traits and survival may depend more on environmental conditions experienced by the population than on the strength of sexual selection.     

Spatio-temporal shifts in gradients of habitat quality for an opportunistic avian predator

We used the conceptual framework of the theory of natural selection to study breeding habitat preferences by an opportunistic avian predator, the black kite Milvus migrans. In Europe, black kite populations are mostly found near large networks of aquatic habitats, usually considered optimal for foraging and breeding. We hypothesized that proximity to wetlands could vary among individuals and affect their fitness, and thus be subject to natural selection. We tested the hypothesis first on a population on Lake Lugano (Italian pre‐Alps) which has been monitored for nine years, and then on seven other populations, each studied for four–five years, located along a continuum of habitat from large water bodies to scarce aquatic habitat of any kind. In the Lake Lugano population, black kite abundance was negatively related to distance to the lake in all the nine years of study, consistent with long‐term natural selection. There was evidence of ongoing directional selection on strategic nest location in three of the years, and evidence of stabilizing selection in two years. In eight of the nine years the trend was for a linear increase in fitness with increasing proximity to the lake. At the population level, results were consistent with adaptive habitat choice in relation to the previous year's spatial variation in fitness: higher associations between fitness and distance to the lake (i.e. higher selection gradients) resulted in higher density variations in the following year, in turn related to the availability of fish, the main local prey. The progressive decline of inland pairs and increase in the density of lakeshore pairs caused a directional long‐term trend of declining mean distance to the lake. Breeding near aquatic habitats was associated with higher foraging success, and higher frequency and biomass of prey deliveries to offspring. There was weak evidence of selection in other populations. The inland‐wetland gradient of habitat quality may have been affected by predation risk, as estimated by density of a major predator of adults and nestlings, the eagle owl Bubo bubo. Behavioral decisions at the level of the individual probably translated into population effects on density and distribution at various spatial scales. Populations in optimal habitats showed higher density and produced six times as many young per unit space as those in sub‐optimal habitats.     

Traffic noise exposure affects telomere length in nestling house sparrows

In a consistently urbanizing world, anthropogenic noise has become almost omnipresent, and there are increasing evidence that high noise levels can have major impacts on wildlife. While the effects of anthropogenic noise exposure on adult animals have been widely studied, surprisingly, there has been little consideration of the effects of noise pollution on developing organisms. Yet, environmental conditions experienced in early life can have dramatic lifelong consequences for fitness. Here, we experimentally manipulated the acoustic environment of free-living house sparrows (Passer domesticus) breeding in nest boxes. We focused on the impact of such disturbance on nestlings’ telomere length and fledging success, as telomeres (the protective ends of chromosomes) appear to be a promising predictor of longevity. We showed that despite the absence of any obvious immediate consequences (growth and fledging success), nestlings reared under traffic noise exposure exhibited reduced telomere lengths compared with their unexposed neighbours. Although the mechanisms responsible for this effect remain to be determined, our results provide the first experimental evidence that noise alone can affect a wild vertebrate''s early-life telomere length. This suggests that noise exposure may entail important costs for developing organisms.     

Probing the adaptive landscape using experimental islands: density-dependent natural selection on lizard body size

Anolis lizards in the Greater Antilles are thought to have diversified through natural selection on body size and shape, presumably due to interspecific competition and variation in locomotor performance. Here we measure natural selection on body size over three years and across seven replicate populations of the brown anole, A. sagrei. We experimentally manipulated an important component of the environment (population density) on several small islands to test the role of density in driving natural selection. Results indicate that the strength of natural selection was proportional to population density (r2 = 0.81), and favored larger body sizes at higher density, presumably owing to the enhanced competitive ability afforded by large size. Changes in the distribution of body size by selective releases of lizards to islands show that this effect did not arise by pure density dependence, since smaller individuals were disproportionately selected against at higher densities. We measured significant broad sense heritability for body size in the laboratory (h2 = 0.55) indicating that selection in the wild could have an evolutionary response. Our results suggest an important effect of population density on natural selection in Anolis lizards.     

Genetic perspectives on northern population cycles: bridging the gap between theory and empirical studies

Many key species in northern ecosystems are characterised by high‐amplitude cyclic population demography. In 1924, Charles Elton described the ecology and evolution of cyclic populations in a classic paper and, since then, a major focus has been the underlying causes of population cycles. Elton hypothesised that fluctuations reduced population genetic variation and influenced the direction of selection pressures. In concordance with Elton, present theories concern the direct consequences of population cycles for genetic structure due to the processes of genetic drift and selection, but also include feedback models of genetic composition on population dynamics. Most of these theories gained mathematical support during the 1970s and onwards, but due to methodological drawbacks, difficulties in long‐term sampling and a complex interplay between microevolutionary processes, clear empirical data allowing the testing of these predictions are still scarce. Current genetic tools allow for estimates of genetic variation and identification of adaptive genomic regions, making this an ideal time to revisit this subject. Herein, we attempt to contribute towards a consensus regarding the enigma described by Elton almost 90 years ago. We present nine predictions covering the direct and genetic feedback consequences of population cycles on genetic variation and population structure, and review the empirical evidence. Generally, empirical support for the predictions was low and scattered, with obvious gaps in the understanding of basic population processes. We conclude that genetic variation in northern cyclic populations generally is high and that the geographic distribution and amount of diversity are usually suggested to be determined by various forms of context‐ and density‐dependent dispersal exceeding the impact of genetic drift. Furthermore, we found few clear signatures of selection determining genetic composition in cyclic populations. Dispersal is assumed to have a strong impact on genetic structuring and we suggest that the signatures of other microevolutionary processes such as genetic drift and selection are weaker and have been over‐shadowed by density‐dependent dispersal. We emphasise that basic biological and demographical questions still need to be answered and stress the importance of extensive sampling, appropriate choice of tools and the value of standardised protocols.     

Selection on incremental variation of eye size in a wild population of Daphnia

Several studies of eye morphology have analysed macroevolutionary patterns in the diversity of eyes, and although these studies are often linked to environment or behaviour, they provide only indirect evidence of selection. Specific data to show the microevolutionary potential for adaptation by natural selection in eye morphology have been lacking. We document directional selection on eye size, an important determinant of visual capabilities, in a wild population of the freshwater microcrustacean Daphnia. We show that even slight changes in eye size may have major consequences for fitness. An increase in eye diameter of 19.9 μm – slightly more than one standard deviation – is associated with an increase in clutch size of one egg, or an increase of nearly 20% of the mean clutch size. Furthermore, relative eye size is genetically variable and thus could evolve in response to the observed selective pressure. We conclude that selection on incremental variation in eye size may have led to differences observed on broader taxonomic scales.     

Selection on laying date is connected to breeding density in the pied flycatcher

Timing of reproduction and clutch size are important determinants of breeding success, especially in seasonal environments. Several recent bird population studies have shown changes in breeding time and in natural selection on it. These changes have often been linked with climate change, but few studies have investigated how the traits or natural selection are actually connected with climatic factors. Furthermore, the effect of population density on selection has been rarely considered, despite the potential importance of density in demographic processes. We studied variation in natural selection on laying date and on clutch size in relation to measures of spring phenology and population density in a long-term study of pied flycatchers in SW Finland. The phenological stage of the environment at mean egg-laying did not affect the direction of selection on either laying date or on clutch size. There was, however, stronger selection for earlier laying date when the breeding density of the population was high, suggesting that early breeding is not necessarily beneficial as such, but that its importance is emphasized when high population density increases competition. In addition, early breeding was favoured when the pre-breeding period was cool, which may indicate an increased advantage for the fittest individuals in harsher conditions. In the middle of the twentieth century, there was selection for large clutch size, which subsequently ceased, along with an overall decrease in recruit production. Our results indicate that attention should be paid to demographic factors such as breeding density when studying natural selection and temporal changes in it.     

Genomic divergence across ecological gradients in the Central African rainforest songbird (Andropadus virens)

The little greenbul, a common rainforest passerine from sub‐Saharan Africa, has been the subject of long‐term evolutionary studies to understand the mechanisms leading to rainforest speciation. Previous research found morphological and behavioural divergence across rainforest–savannah transition zones (ecotones), and a pattern of divergence with gene flow suggesting divergent natural selection has contributed to adaptive divergence and ecotones could be important areas for rainforests speciation. Recent advances in genomics and environmental modelling make it possible to examine patterns of genetic divergence in a more comprehensive fashion. To assess the extent to which natural selection may drive patterns of differentiation, here we investigate patterns of genomic differentiation among populations across environmental gradients and regions. We find compelling evidence that individuals form discrete genetic clusters corresponding to distinctive environmental characteristics and habitat types. Pairwise F_ST between populations in different habitats is significantly higher than within habitats, and this differentiation is greater than what is expected from geographic distance alone. Moreover, we identified 140 SNPs that showed extreme differentiation among populations through a genomewide selection scan. These outliers were significantly enriched in exonic and coding regions, suggesting their functional importance. Environmental association analysis of SNP variation indicates that several environmental variables, including temperature and elevation, play important roles in driving the pattern of genomic diversification. Results lend important new genomic evidence for environmental gradients being important in population differentiation.     

Interactions between parental traits,environmental harshness and growth rate in determining telomere length in wild juvenile salmon

A larger body size confers many benefits, such as increased reproductive success, ability to evade predators and increased competitive ability and social status. However, individuals rarely maximize their growth rates, suggesting that this carries costs. One such cost could be faster attrition of the telomeres that cap the ends of eukaryotic chromosomes and play an important role in chromosome protection. A relatively short telomere length is indicative of poor biological state, including poorer tissue and organ performance, reduced potential longevity and increased disease susceptibility. Telomere loss during growth may also be accelerated by environmental factors, but these have rarely been subjected to experimental manipulation in the natural environment. Using a wild system involving experimental manipulations of juvenile Atlantic salmon Salmo salar in Scottish streams, we found that telomere length in juvenile fish was influenced by parental traits and by direct environmental effects. We found that faster‐growing fish had shorter telomeres and there was a greater cost (in terms of reduced telomere length) if the growth occurred in a harsher environment. We also found a positive association between offspring telomere length and the growth history of their fathers (but not mothers), represented by the number of years fathers had spent at sea. This suggests that there may be long‐term consequences of growth conditions and parental life history for individual longevity.     

Cohort effects and population dynamics

Cohort effects originate from environmental conditions, and can have long‐term consequences for the cohort's performance. It has been proposed that cohort effects tend to increase population fluctuations. However, differences among individuals, which cohort effects introduce into a population, usually have stabilizing effects. There are thus two different predictions regarding the impact of cohort effects on population fluctuations. We argue that it is important to distinguish between environmental variability and its long‐term effects on individual quality, and approach the question with a population model that can include or exclude such effects. We show that the influence of cohort effects depends on the inherent dynamics: cohort effects can have stabilizing effects if dynamics are inherently unstable. However, the most common outcome is destabilization whenever cohort effects act on top of inherently stable dynamics. Intriguingly, both alternatives are due to individual differences affecting the structure of density dependence in a similar way.     

Natural Selection Constrains Neutral Diversity across A Wide Range of Species

The neutral theory of molecular evolution predicts that the amount of neutral polymorphisms within a species will increase proportionally with the census population size (Nc). However, this prediction has not been borne out in practice: while the range of Nc spans many orders of magnitude, levels of genetic diversity within species fall in a comparatively narrow range. Although theoretical arguments have invoked the increased efficacy of natural selection in larger populations to explain this discrepancy, few direct empirical tests of this hypothesis have been conducted. In this work, we provide a direct test of this hypothesis using population genomic data from a wide range of taxonomically diverse species. To do this, we relied on the fact that the impact of natural selection on linked neutral diversity depends on the local recombinational environment. In regions of relatively low recombination, selected variants affect more neutral sites through linkage, and the resulting correlation between recombination and polymorphism allows a quantitative assessment of the magnitude of the impact of selection on linked neutral diversity. By comparing whole genome polymorphism data and genetic maps using a coalescent modeling framework, we estimate the degree to which natural selection reduces linked neutral diversity for 40 species of obligately sexual eukaryotes. We then show that the magnitude of the impact of natural selection is positively correlated with Nc, based on body size and species range as proxies for census population size. These results demonstrate that natural selection removes more variation at linked neutral sites in species with large Nc than those with small Nc and provides direct empirical evidence that natural selection constrains levels of neutral genetic diversity across many species. This implies that natural selection may provide an explanation for this longstanding paradox of population genetics.     

Intra‐specific relatedness,spatial clustering and reduced demographic performance in tropical rainforest trees

Intra‐specific negative density dependence promotes species coexistence by regulating population sizes. Patterns consistent with such density dependence are frequently reported in diverse tropical tree communities. Empirical evidence demonstrating whether intra‐specific variation is related to these patterns, however, is lacking. The present study addresses this important knowledge gap by genotyping all individuals of a tropical tree in a long‐term forest dynamics plot in tropical China. We show that related individuals are often spatially clustered, but having closely related neighbours reduces the growth performance of focal trees. We infer from the evidence that dispersal limitation and negative density dependence are operating simultaneously to impact the spatial distributions of genotypes in a natural population. Furthermore, dispersal limitation decreases local intra‐specific genetic diversity and increases negative density dependence thereby promoting niche differences and species coexistence as predicted by theory.     

SEXUAL SELECTION COUNTERACTS EXTINCTION OF SMALL POPULATIONS OF THE BULB MITES

Genetic drift in small populations can increase frequency of deleterious recessives and consequently lead to inbreeding depression and population extinction. On the other hand, as homozygosity at deleterious recessives increases, they should be purged from populations more effectively by selection. Sexual selection has been postulated to strengthen selection against deleterious mutations, and should thus decrease extinction rate and intensify purging of inbreeding depression. We tested these predictions in the bulb mite Rhizoglyphus robini. We created 100 replicate lines of small populations (five males and five females) and in half of them experimentally removed sexual selection by enforcing monogamy. The lines were propagated for eight generations and then assayed for purging of inbreeding depression. We found that proportion of lines which went extinct was lower with sexual selection than without. We also found evidence for purging of inbreeding depression in the lines with sexual selection, but not in lines without sexual selection. Our results suggest that purging of inbreeding depression was more effective against mutations with relatively large deleterious effects. Thus, although our data clearly indicate a positive impact of sexual selection on short‐term survival of bottlenecked populations, long‐term consequences are less clear as they may be negatively impacted by accumulation of deleterious mutations of small effect.     

Crickets increase sexual signalling and sperm protection but live shorter in the presence of rivals

The sociosexual environment animals experience through their life can shape the evolution of key life history traits, including longevity. Male–male competition, for instance, may influence the resources allocated to traits involved in male reproductive success. Here, I test whether lifelong exposure to a competitor male influences male investment in pre‐ and post‐copulatory sexual traits (calling effort and sperm quality) and how this affected the oxidative status and longevity of male field crickets (Gryllus bimaculatus). As expected, the visual exposure to a mating competitor promoted a higher calling effort in the male crickets but resulted in a decline in the haemolymph antioxidant content. The haemolymph antioxidant content negatively covaried with the antioxidant content of the sperm but interestingly, only when the males were exposed to a competitor. This suggests that when mating competition is high, males may prioritize sperm antioxidant protection against somatic maintenance. Finally, I provide evidence that male–male competition imposes additional costs to reproduction, as males exposed to a mating competitor lost a significant proportion of body antioxidant defences and body mass over time and in long term, had a reduced lifespan. Overall, this study strongly suggests that intrasexual competition may impose additional oxidative costs during reproduction for males, with negative consequences on lifespan. Moreover, the results highlight the role of oxidative stress as a physiological mechanism underlying the trade‐off between reproduction‐longevity and through which sexual selection may contribute to the evolution of senescence patterns.     

EXPERIMENTAL MANIPULATION OF PUTATIVE SELECTIVE AGENTS PROVIDES EVIDENCE FOR THE ROLE OF NATURAL ENEMIES IN THE EVOLUTION OF PLANT DEFENSE

Although biologists have long assumed that plant resistance characters evolved under selection exerted by such natural enemies as herbivores and pathogens, experimental evidence for this assumption is sparse. We present evidence that natural enemies exert selection on particular plant resistance characters. Specifically, we demonstrate that elimination of natural enemies from an experimental field population of Arabidopsis thaliana alters the pattern of selection on genetic variation in two characters that have been shown to reduce herbivore damage in the field: total glucosinolate concentration and trichome density. The change in pattern of selection reveals that natural enemies imposed selection favoring increased glucosinolate concentration and increased trichome density, and thus, supports one of the major assumptions of the coevolution hypothesis. We also demonstrate that a pattern of stabilizing selection on glucosinolate concentration results from a balance between the costs and benefits associated with increasing levels of this resistance character. This result provides direct confirmation of the appropriateness of cost-benefit models for characterizing the evolution of plant defenses.     

The evolutionary history of Drosophila buzzatii. XVI. Fitness component analysis in an original natural population from Argentina

The adaptive significance of the chromosomal polymorphism of Drosophila buzzati has been studied by means of fitness component analysis in an original population from Argentina. The results show evidence of selection acting through pupal viability, longevity (adult viability) and fecundity on the second chromosome polymorphism, and through pupal viability and virility on the fourth chromosome polymorphism. Changes in chromosomal inversion frequencies throughout the life-cycle suggested an endocyclic pattern of directional selection, which at first seems to be the only detectable mechanism responsible for the maintenance of the polymorphism. However, slow, long-term frequency changes cannot be ruled out. The way in which endocyclic selection acts on this population is different from that in a colonized population previously studied; that is, different fitness components are involved in the maintenance of chromosomal polymorphism. The possible factors that may explain these differences are discussed.     

Recent evolutionary history predicts population but not ecosystem‐level patterns

In the face of rapid anthropogenic environmental change, it is increasingly important to understand how ecological and evolutionary interactions affect the persistence of natural populations. Augmented gene flow has emerged as a potentially effective management strategy to counteract negative consequences of genetic drift and inbreeding depression in small and isolated populations. However, questions remain about the long‐term impacts of augmented gene flow and whether changes in individual and population fitness are reflected in ecosystem structure, potentiating eco‐evolutionary feedbacks. In this study, we used Trinidadian guppies (Poecilia reticulata) in experimental outdoor mesocosms to assess how populations with different recent evolutionary histories responded to a scenario of severe population size reduction followed by expansion in a high‐quality environment. We also investigated how variation in evolutionary history of the focal species affected ecosystem dynamics. We found that evolutionary history (i.e., gene flow vs. no gene flow) consistently predicted variation in individual growth. In addition, gene flow led to faster population growth in populations from one of the two drainages, but did not have measurable impacts on the ecosystem variables we measured: zooplankton density, algal growth, and decomposition rates. Our results suggest that benefits of gene flow may be long‐term and environment‐dependent. Although small in replication and duration, our study highlights the importance of eco‐evolutionary interactions in determining population persistence and sets the stage for future work in this area.     

ANCIENT URBANIZATION PREDICTS GENETIC RESISTANCE TO TUBERCULOSIS

A link between urban living and disease is seen in recent and historical records, but the presence of this association in prehistory has been difficult to assess. If the transition to urban living does result in an increase in disease‐based mortality, we might expect to see evidence of increased disease resistance in longer‐term urbanized populations, as the result of natural selection. To test this, we determined the frequency of an allele (SLC11A1 1729 + 55del4) associated with natural resistance to intracellular pathogens such as tuberculosis and leprosy. We found a highly significantly correlation with duration of urban settlement—populations with a long history of living in towns are better adapted to resisting these infections. This correlation remains strong when we correct for autocorrelation in allele frequencies due to shared population history. Our results therefore support the interpretation that infectious disease loads became an increasingly important cause of human mortality after the advent of urbanization, highlighting the importance of population density in determining human health and the genetic structure of human populations.     

Pathogen frequency in an age-structured population of<Emphasis Type="Italic"> Plantago lanceolata</Emphasis>

Life-history traits can play important roles in determining the course of ecological species interactions. We explored the consequences of host age on a host-pathogen interaction by quantifying pathogen frequency in an age-structured host population. Our project was motivated by an interest in whether the demographic structure of a host population has consequences for species interactions. In 2 successive years, we planted large cohorts of the perennial herb Plantago lanceolata in its natural environment and observed infection by Fusarium moniliforme, a non-lethal floral fungal pathogen, over 3 years. We documented substantial variation of pathogen frequency across years and between cohorts. Logistic regression revealed that pathogen frequency increased with the number of inflorescences produced and with evidence of prior pathogen presence, whereas it decreased with increasing plant longevity. In addition, interannual variation and an age-year interaction contributed to the observed pathogen frequencies. There was a significant positive effect of age on pathogen frequency overall, but this was not consistent over all ages. Pathogen frequency was higher in 2-year-old plants than in 1-year-olds, suggesting that age-structure can influence the host-pathogen interaction. This pattern did not continue into 3-year-old plants. A possible explanation for this is that selective mortality allows only generally robust plants, and consequently the most resistant plants, to survive to the oldest ages.