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The shared functions of the skull are thought to result in common evolutionary patterns in mammalian cranial shape. Craniofacial evolutionary allometry (CREA) is a particularly prominent pattern where larger species display proportionally elongate facial skeletons and smaller braincases. It was recently proposed that CREA arises from biomechanical effects of cranial scaling when diets are similar. Thus, deviations from CREA should occur with changes in cranial biomechanics, for example due to dietary change. Here, we test this using 3D geometric morphometric analysis in a dataset of Australian murine crania, which are highly allometric. We contrast allometric and non‐allometric variation in the cranium by comparing evolutionary mode, allometry, ordinations, as well as allometry, integration, and modularity in functional modules. We found evidence of stabilising selection in allometry‐containing and size‐free shape, and substantial non‐allometric variation aligned with dietary specialisation in parallel with CREA. Integration among cranial modules was higher, and modularity lower, with size included, but integration between rostrum and cranial vault, which are involved in the CREA pattern, dropped dramatically after size removal. Our results thus support the hypothesis that CREA is a composite arising from selection on cranial function, with substantial non‐allometric shape variation occurring alongside CREA where dietary specialisation impacts selection on gnawing function. This emphasises the need to research mammalian cranial evolution in the context of allometric and non‐allometric selection on biomechanical function. [ABSTRACT FROM AUTHOR]

As temperatures rise, timing of reproduction is changing at different rates across trophic levels, potentially resulting in asynchrony between consumers and their resources. The match–mismatch hypothesis (MMH) suggests that trophic asynchrony will have negative impacts on average productivity of consumers. It is also thought to lead to selection on timing of breeding, as the most asynchronous individuals will show the greatest reductions in fitness.Using a 30‐year individual‐level dataset of breeding phenology and success from a population of European shags on the Isle of May, Scotland, we tested a series of predictions consistent with the hypothesis that fitness impacts of trophic asynchrony are increasing.These predictions quantified changes in average annual breeding success and strength of selection on timing of breeding, over time and in relation to rising sea surface temperature (SST) and diet composition.Annual average (population) breeding success was negatively correlated with average lay date yet showed no trend over time, or in relation to increasing SST or the proportion of principal prey in the diet, as would be expected if trophic mismatch was increasing. At the individual level, we found evidence for stabilising selection and directional selection for earlier breeding, although the earliest birds were not the most productive. However, selection for earlier laying did not strengthen over time, or in relation to SST or slope of the seasonal shift in diet from principal to secondary prey. We found that the optimum lay date advanced by almost 4 weeks during the study, and that the population mean lay date tracked this shift.Our results indicate that average performance correlates with absolute timing of breeding of the population, and there is selection for earlier laying at the individual level. However, we found no fitness signatures of a change in the impact of climate‐induced trophic mismatch, and evidence that shags are tracking long‐term shifts in optimum timing. This suggests that if asynchrony is present in this system, breeding success is not impacted. Our approach highlights the advantages of examining variation at both population and individual levels when assessing evidence for fitness impacts of trophic asynchrony. [ABSTRACT FROM AUTHOR]

The capacity of species to tolerate physical stressors is critical in a world of increasing environmental instability, however, past selective environments should dramatically impact on future stress tolerance, particularly in isolated populations. Through stabilising selection, long-term environmental stasis may reduce physiological tolerance, creating an evolutionary legacy where populations are less fit if environments change. Few empirical studies have investigated this evolutionary legacy of past selection, and of particular interest whether stabilising selection in a benign environment reduces stress tolerance in natural systems. Here we use multiple populations of salt-lake invertebrates (Coxiella striata, Austrochiltonia subtenuis) with either stable or fluctuating environmental histories to investigate the relationship between stabilising selection and environmental stress resistance. Tolerance to both salinity and temperature stress were examined in invertebrate populations from lakes with long-term (decadal) stable environments and compared with populations from lakes with extreme salinity variations. Individuals from stable environments demonstrated significantly lower survival under both increasing salinity and temperature stresses when compared with environmentally unstable populations. Our results support the hypothesis that the evolutionary legacy from stabilising selection in constant environments leads to reduced stress tolerance. This finding demonstrates that under an increasingly variable climate, the evolutionary legacies of populations will be critical for future survival and adaptation. [ABSTRACT FROM AUTHOR]

Research linking functional traits to competitive ability of coexisting species has largely relied on rectilinear correlations, yielding inconsistent results. Based on concepts borrowed from natural selection theory, we propose that trait–competition relationships can generally correspond to three univariate selection modes: directional (a rectilinear relationship), stabilising (an n‐shaped relationship), and disruptive (a u‐shaped relationship). Moreover, correlational selection occurs when two traits interact in determining competitive ability and lead to an optimum trait combination (i.e., a bivariate nonlinear selection mode).We tested our ideas using two independent datasets, each one characterising a group of species according to (a) their competitive effect on a target species in a pot experiment and (b) species‐level values of well‐known functional traits extracted from existing databases. The first dataset comprised 10 annual plant species frequent in a summer‐rainfall desert in Argentina, while the second consisted of 37 herbaceous species from cool temperate vegetation types in Canada. Both experiments had a replacement design where the identity of neighbours was manipulated holding total plant density in pots constant. We modelled the competitive ability of neighbours (i.e., the log inverse of target plant biomass) as a function of traits using normal multiple regression.Leaf dry matter content (LDMC) was consistently subjected to negative directional selection in both experiments as well as to stabilising selection among temperate species. Leaf size was subjected to stabilising selection among desert species while among temperate species, leaf size underwent correlational selection in combination with specific leaf area (SLA): selection on SLA was negative directional for large‐leaved species, while it was slightly positive for small‐leaved species.Synthesis. Multiple quadratic regression adds functional flexibility to trait‐based community ecology while providing a standardised basis for comparison among traits and environments. Our analyses of two datasets from contrasting environmental conditions indicate (a) that leaf dry matter content can capture an important component of plant competitive ability not accounted for by widely used competitive traits, such as specific leaf area, leaf size, and plant height and (b) that optimum relationships (either univariate or bivariate) between competitive ability and plant traits may be more common than previously realised. [ABSTRACT FROM AUTHOR]

Illnesses caused by a variety of micro- and macro- organisms can negatively affect individuals’ fitness, leading to the expectation that immunity is under positive selection. However, immune responses are costly and individuals must trade-off their immune response with other fitness components (e.g. survival or reproductive success) meaning that individuals with intermediate response may have the greatest overall fitness. Such a process might be particularly acute in species with strong sexual selection because the condition-dependence of male secondary sexual-traits might lead to striking phenotypic differences amongst males of different immune response levels. We tested whether there is selection on immune response by survival and reproduction in yearling and adult male black grouse (Lyrurus tetrix) following an immune challenge with a novel antigen and tested the hypothesis that sexual signals and body mass are honest signals of the immune response. We show that yearling males with highest immune response to these challenges had higher survival, but the reverse was true for adults. Adults with higher responses had highest mass loss and adult males with intermediate immune response had highest mating success. Tail length was related to baseline response in adults and more weakly in yearlings. Our findings reveal the complex fitness consequences of mounting an immune response across age classes. Such major differences in the direction and magnitude of selection in multiple fitness components is an alternative route underpinning the stabilising selection of immune responses with an intermediate immune response being optimal. [ABSTRACT FROM AUTHOR]

Phylogenetic studies of lineages growing in extreme environments have frequently recovered evidence not only of high level of homoplasy but also of discordance of morphological disparity and species diversity. It has been suggested that this divergence may be caused by developmental constraints and/or natural selection. Here we explored these hypotheses by inferring the phenotypic evolution of the derived liverwort genus Cololejeunea. These liverworts occur preferentially on the surface of leaves or other aerial parts of vascular plants growing in wet forests. The evolution of 12 morphological characters was studied using a phylogenetic framework comprising 70 species of Cololejeunea. The phylogeny was reconstructed using DNA sequences of one nuclear and two plastid regions and enabled the inference of the evolution of the studied morphological characters by determining the frequency of homoplasy. Mantel tests were used to test for correlations of morphological disparity × species diversity and morphological disparity × epiphytism. The phylogenetic informativeness of each binary character was estimated by the D metric of the Fritz and Purvis test, and the relationship between each character and epiphytism was inferred by Pearson's coefficient. We evaluated the morphospace occupation using principal coordinate analyses. Our results not only recovered high levels of homoplasy but also weak correlations of morphological disparity and species diversity. Morphological disparity was not linked to epiphytism, although positive or negative relationships between some characters and epiphytism were found. The Brownian model of character evolution was not rejected for the studied morphological disparity in Cololejeunea with the exception of asexual propagules. The observations support the prediction that iterative evolution in a well-defined morphospace may result in rampant homoplasy and the observed divergence of disparity and diversity. [ABSTRACT FROM AUTHOR]

We analysed 76 accessions of the medicinal and nutritive rice ( Oryza sativa) landrace Njavara ( Shashtika in Sanskrit) and 67 traditional cultivars of Kerala state using phenotypic traits and microsatellite markers. Multivariate analyses of 13 quantitative phenotypic traits revealed two distinct clusters, the Njavara accessions and the traditional cultivars ( Qst = 0.4753). Njavara accessions belonging to the same morphotype were clustered together, although no specific pattern could be deciphered from the clustering of traditional cultivars. A total of 222 alleles were generated at the 24 microsatellite loci, with a mean number of 4.42 and 6.623 alleles per locus and a mean gene diversity ( He) of 0.479 and 0.596 in Njavara and traditional cultivars, respectively. Different diversity analyses clearly separated Njavara and traditional cultivars from each other. As with the phenotypic analysis, Njavara accessions clustered according to their morphotypes, but the topology of the two dendrograms were different. However, the clustering pattern of traditional cultivars in genotypic dendrogram was inconsistent with that of phenotypic dendrogram. The scented rice cultivars formed a distinct cluster while the remaining traditional cultivars were clustered according to their photosensitivity. Significant ( P < 0.0000) partitioning of molecular diversity was recorded between Njavara and traditional cultivars ( Fst = 0.4293), within Njavara types ( Fst = 0.5616) and traditional cultivars ( Fst = 0.2546). The results indicate that Njavara is a cryptic variant within the traditional rice gene pool in Kerala. The study provided valuable information on the genetic structure and population differentiation of traditional rice cultivars in Kerala, which are relevant in breeding and conservation. [ABSTRACT FROM AUTHOR]

A tradeoff between immune response and life history traits, in particular growth rate, has been documented in various bird species. Ostriches are fast-growing birds and a typical feature of cohorts is that offspring often differ greatly in size. We investigated the relationship between hatching date and growth rate of chicks and both cell-mediated (measured using a phytohaemagglutinin (PHA) injection) and humoral immune responses in ostrich chicks maintained on a research farm. Chicks with higher growth rates had intermediate responses to both diphtheria and tetanus toxoids. By contrast, no relation between growth rates and responses to PHA injection were found. We conclude that chick growth rate variation may be explained beyond a certain threshold by a tradeoff between the humoral response and growth. Both responses to PHA injection and humoral responses in chicks were found to decrease with chick hatching date. Within the context of ostrich farming, these results could partially explain size variations observed in cohorts of chicks, as well as high mortality rates during their first 3 months of age. [ABSTRACT FROM AUTHOR]

Deleterious impacts of major mutations can be ameliorated by stabilising selection acting on modifier genes. We hypothesise that a new hyperactive circling mouse (counterspin: Cr) arises when modifier genes inadvertently selected to ameliorate the negative impacts of a growth hormone transgenic insertion segregate into the normal genetic background that lacks the transgene. We hypothesise that such modifiers generate a phenotype “mirror image” to the transgenics on the otherwise normal background. We highlight this by testing a priori hypotheses that counterspin and transgenic growth hormone mice deviate oppositely from normal mice across a broad spectrum of characteristics. Results spanning growth, sensorimotor performance, cognition and striatal neurotransmitters provide strong circumstantial evidence for the hypothesis. In a more direct test for selection in the transgenic mice, we found that those examined in 2008 slept ~3 h/d less than they did 14 years ago ( P < 0.0005). This is a profound change strongly supporting the reality of modifier selection in these mice. Our results highlight that modifiers may act powerfully on genetically engineered constructs given a genetically variable background. Furthermore, we suggest that modifier selection might provide a novel method for deriving genetic models, and specifically, models phenotypically opposite to engineered constructs or natural mutations. [ABSTRACT FROM AUTHOR]

Abstract: We investigate a genetic model of a large population of sexual organisms in a changing environment. The organisms are subject to stabilising selection on a quantitative trait, with environmental change causing the fitness optimum to move. When the fitness optimum moves slowly, adaptation to the changing environment occurs by means of reasonably well-separated substitutions at the loci controlling the trait. In this way, the trait generally tracks the moving optimum, but in such a case, the population may exhibit periods of time where the mean trait value overshoots the moving optimal trait value, thereby exhibiting an apparent anticipation of selection. The mechanism underlying this phenomenon is determined from consideration of a simpler model that correctly captures the observed dynamical behaviour. We note that very slow rates of changes of traits are seen in the fossil record and the present work may be relevant to this topic. [Copyright &y& Elsevier]

Abstract: Fisher''s geometrical model amounts to a description of mutation and selection for individuals characterised by a number of quantitative traits. In the present work the fitness landscape is not assumed to be spherically symmetric, hence different points, i.e. phenotypes, on a surface of constant fitness generally have different curvatures. We investigate two different approximations of Fisher''s geometrical model that have appeared in the literature. One approximation uses the average curvature of the fitness surface at the parental phenotype. The other approach is based on a normal approximation of a distribution associated with new mutations. Analytical results and simulations are used to compare the accuracy of the two approximations. [Copyright &y& Elsevier]

Abstract: We consider a large population of asexual organisms characterised by a number of quantitative traits that are subject to stabilising selection. Mutation is taken to act pleiotropically, with every mutation generally changing all of the traits under selection. We focus on the equilibrium distribution of the population, where mutation and selection are in balance. It has been previously established that the equilibrium distribution of genotypic effects may be anomalous, as it may contain a singular spike—a Dirac delta function—corresponding to a non-zero proportion of the population having exactly optimal genotypic values. In the present work, we present exact results for the case where three traits are under selection. These results give the equilibrium genetic variance of the population, and the proportion of the population that have the optimal genotype. This is achieved for two different spherically symmetric distributions of mutant effects. Additionally, a simple and robust numerical approach is also presented that allows the treatment of some other mutation distributions, where there are an arbitrary number of selected traits. [Copyright &y& Elsevier]

Timing of arrival at a resource often determines an individual’s reproductive success. Tree-killing bark beetles can reproduce in healthy trees by attacking in adequate numbers to overcome host defences that could otherwise be lethal. This process is mediated by aggregation and antiaggregation pheromones. Beetles that arrive early in such a “mass attack” must contend with undiminished tree defences, and produce enough pheromones to attract more beetles, but have a head start on gallery construction and egg-laying. Beetles that arrive late may be impeded by competition and diminishing availability of phloem, but should experience fewer costs associated with pheromone production and battling tree defences. We investigated relationships between timing of arrival, body size, pheromone production and fitness in the southern pine beetle, Dendroctonus frontalis. In field experiments, we captured beetles that arrived early (pioneers) and late on slash pine trees, Pinus elliottii, and measured pheromone amounts in their hindguts. We marked gallery entrances of beetles as they landed on a tree and measured their reproductive success after the attack terminated. We found no difference in body size or pheromone amounts between early and late arrivers. Most beetles arrived at the middle of the attack sequence, and excavated longer galleries per day than early arrivers. The number of offspring produced per day by beetles that established galleries midway through mass attack was higher than those that arrived early or very late in the sequence. Our results suggest that beetles do not exhibit adaptive phenotypic plasticity in pre-landing pheromone production, depending on the extent of previous colonisation of a host. Rather, it appears that stabilising selection favours beetles that attack in the middle of the sequence, and contributes to attack synchrony. Synchronous attack on trees is essential before population booms characteristic of tree-killing bark beetles can occur in nature. [ABSTRACT FROM AUTHOR]

We consider the implications of mutationally non-equivalent loci for large populations of randomly mating diploid organisms under mutation–selection balance. Variation, across loci, of parameters such as the allelic mutational variance and the mutation rate, is shown to reduce the equilibrium genetic variance. This is proved to follow from the genetic variance contributed by a single locus having an underlying convexity. We give approximate results indicating the way small deviations of the mutational parameters, from their mean values, reduce the genetic variance. Numerical estimates of the size of the effect are given for more general variations of the parameters. Variation in the mutation rates has a significantly smaller effect than variation in the mutational variances. Under accepted parameter values, the reduction in genetic variance can be substantial. [Copyright &y& Elsevier]

The theory of pleiotropic mutation and selection is investigated and developed for a large population of asexual organisms. Members of the population are subject to stabilising selection on Ω phenotypic characters, which each independently affect fitness. Pleiotropy is incorporated into the model by allowing each mutation to simultaneously affect all characters. To expose differences with continuous-allele models, the characters are taken to originate from discrete-effect alleles and thus have discrete genotypic effects. Each character can take the values n×Δ where n=0,±1,±2,…, and the splitting in character effects, Δ, is a parameter of the model. When the distribution of mutant effects is normally distributed around the parental value, and Δ is large, a “stepwise” model of mutation arises, where only adjacent trait effects are accessible from a single mutation. The present work is primarily concerned with the opposite limit, where Δ is small and many different trait effects are accessible from a single mutation.In contrast to what has been established for continuous-effect models, discrete-effect models do not yield a singular equilibrium distribution of genotypic effects for any value of Ω. Instead, for different values of Ω, the equilibrium frequencies of trait values have very different dependencies on Δ. For Ω=1 and 2, decreasing Δ broadens the width of the frequency distribution and hence increases the equilibrium level of polymorphism. For all sufficiently large values of Ω, however, decreasing Δ decreases the width of the frequency distribution and the equilibrium level of polymorphism. The connection with continuous trait models follows when the limit Δ→0 is considered, and a singular probability density of trait values is obtained for all sufficiently large Ω. [Copyright &y& Elsevier]

Sokolovska et al. (2002) used 57 weighted correlation coefficients derived from 33 published studies of 20 species to produce the first meta-analysis of body size and fitness in odonates. They concluded that 'there is a general fitness benefit to large size of odonates'. Given the inherent biases of their analyses, such a conclusion is hardly surprising. In this paper, their study was used to illustrate major problems that arise from (1) ignoring the possibility of stabilising selection on body size, (2) using statistics from the literature indiscriminately, when the original studies are inappropriate for comparative analysis, and (3) relying heavily on transformed statistics derived from small samples. Suggestions are made as to how data can be presented and analysed in ways conducive to future comparative analyses on size relations to fitness in odonates and other insects. [ABSTRACT FROM AUTHOR]

It is the differences between sperm and eggs that fundamentally underpin the differences between the sexes within reproduction. For males, it is theorized that widespread sperm competition leads to selection for investment in sperm numbers, achieved by minimizing sperm size within limited resources for spermatogenesis in the testis. Here, we empirically examine how sperm competition shapes sperm size, after more than 77 generations of experimental selection of replicate lines under either high or low sperm competition intensities in the promiscuous flour beetle Tribolium castaneum . After this experimental evolution, populations had diverged significantly in their sperm competitiveness, with sperm in ejaculates from males evolving under high sperm competition intensities gaining 20% greater paternity than sperm in ejaculates from males that had evolved under low sperm competition intensity. Males did not change their relative investment into sperm production following this experimental evolution, showing no difference in testis sizes between high and low intensity regimes. However, the more competitive males from high sperm competition intensity regimes had evolved significantly longer sperm and, across six independently selected lines, there was a significant association between the degree of divergence in sperm length and average sperm competitiveness. To determine whether such sperm elongation is costly, we used dietary restriction experiments, and revealed that protein-restricted males produced significantly shorter sperm. Our findings therefore demonstrate that sperm competition intensity can exert positive directional selection on sperm size, despite this being a costly reproductive trait.