The fitness effects of mutations on a given genotype are rarely constant across environments to which this genotype is more or less adapted, that is, between more or less stressful conditions. This can have important implications, especially on the evolution of ecological specialization. Stress is thought to increase the variance of mutations' fitness effects, their average, or the number of expressed mutations. Although empirical evidence is available for these three mechanisms, their relative magnitude is poorly understood. In this paper, we propose a simple approach to discriminate between these mechanisms, using a survey of empirical measures of mutation effects in contrasted environments. This survey, across various species and environments, shows that stress mainly increases the variance of mutations' effects on fitness, with a much more limited impact on their average effect or on the number of expressed mutations. This pattern is consistent with a simple model in which fitness is a Gaussian function of phenotypes around an environmentally determined optimum. These results suggest that a simple, mathematically tractable landscape model may not be quantitatively as unrealistic as previously suggested. They also suggest that mutation parameter estimates may be strongly biased when measured in stressful environments.

Inbreeding is known to reduce heterozygosity of neutral genetic markers, but its impact on quantitative genetic variation is debated. Theory predicts a linear decline in additive genetic variance (V_A) with increasing inbreeding coefficient (F) when loci underlying the trait act additively, but a nonlinear hump-shaped relationship when dominance and epistasis are important. Predictions for heritability (h²) are similar, although the exact shape depends on the value of h² in the absence of inbreeding. We located 22 published studies in which the level of genetic variation in experimentally inbred populations (measured by V_A or h²) was compared with that in outbred control populations. For life-history traits, the data strongly supported a nonlinear change in genetic variation with increasing F. V_A and h² were, respectively, 244% and 50% higher at F = 0.4 than in outbred populations, and dominance plus epistatic variance together exceeded additive variance by a factor of four. For nonfitness traits the decline was linear and estimates of nonadditive variance were small. These results confirm that population bottlenecks frequently increase V_A in some traits, and imply that life-history traits are underlain by substantial dominance or epistasis. However, the importance of drift-induced genetic variation in conservation or evolutionary biology is questionable, in part because inbreeding depression usually accompanies inbreeding.

Comparative phylogeographic studies often reveal disparate levels of sequence divergence between lineages spanning a common geographic barrier, leading to the conclusion that isolation was nonsynchronous. However, only rarely do researchers account for the expected variance associated with ancestral coalescence and among-taxon variation in demographic history. We introduce a flexible approximate Bayesian computational (ABC) framework that can test for simultaneous divergence (TSD) using a hierarchical model that incorporates idiosyncratic differences in demographic history across taxon pairs. The method is tested across a range of conditions and is shown to be accurate even with single-locus mitochondrial DNA (mtDNA) data. We apply this method to a landmark dataset of putative simultaneous vicariance, eight geminate echinoid taxon pairs thought to have been split by the Isthmus of Panama 3.1 million years ago. The ABC posterior estimates are not consistent with a history of simultaneous vicariance given these data. Subsequent ABC estimates under a constrained model that assumes two divergence times across the eight taxon pairs suggests simultaneous divergence 3.1 million years ago in seven of the taxon pairs and a more recent divergence in the remaining taxon pair. These ABC estimates on the simultaneous divergence of the seven taxon pairs correspond to a DNA substitution rate of approximately 1.59% per lineage per million years at the mtDNA cytochrome oxidase I gene. This ABC framework can easily be modified to analyze single taxon-pair datasets and/or be expanded to include multiple loci, migration, recombination, and other idiosyncratic demographic histories. The flexible aspect of ABC and its built-in evaluation of estimator bias and statistical power has the potential to greatly enhance statistical rigor in phylogeographic studies.

In flowering plants, the onset and duration of female receptivity vary among species. In several species the receptive structures wilt upon pollination. Here we explore the hypothesis that postpollination wilting may be influenced by pollen and serve as a general means to secure paternity of the pollen donor at the expense of female fitness. Taking a game-theoretical approach, we examine the potential for the evolution of a pollen-borne wilting substance, and for the coevolution of a defense strategy by the recipient plant. The model without defense predicts an evolutionarily stable strategy (ESS) for the production of wilting substance. The ESS value is highest when pollinator visiting rates are intermediate and when the probability that pollen from several donors arrives at the same time is low. This finding has general implications in that it shows that male traits to secure paternity also can evolve in species, such as plants, where mating is not strictly sequential. We further model coevolution of the wilting substance with the timing of stigma receptivity. We assume that pollen-receiving plants can reduce the costs induced by toxic pollen by delaying the onset of stigmatic receptivity. The model predicts a joint ESS, but no female counter-adaptation when the wilting substance is highly toxic. This indicates that toxicity affects the probability that a male manipulative trait stays beneficial (i.e., not countered by female defense) over evolutionary time. We discuss parallels to male induced changes in female receptivity known to occur in animals and the role of harm for the evolution of male manipulative adaptations.

The timing of when to initiate reproduction is an important transition in any organism's life cycle. There is much variation in flowering time among populations, but we do not know to what degree this variation contributes to local adaptation. Here we use a reciprocal transplant experiment to examine the presence of divergent natural selection for flowering time and local adaptation between two distinct populations of Mimulus guttatus. We plant both parents and hybrids (to tease apart differences in suites of associated parental traits) between these two populations into each of the two native environments and measure floral, vegetative, life-history, and fitness characters to assess which traits are under selection at each site. Analysis of fitness components indicates that each of these plant populations is locally adapted. We obtain striking evidence for divergent natural selection on date of first flower production at these two sites. Early flowering is favored at the montane site, which is inhabited by annual plants and characterized by dry soils in midsummer, whereas intermediate (though later) flowering dates are selectively favored at the temperate coastal site, which is inhabited by perennial plants and is almost continually moist. Divergent selection on flowering time contributes to local adaptation between these two populations of M. guttatus, suggesting that genetic differentiation in the timing of reproduction may also serve as a partial reproductive isolating barrier to gene flow among populations.

Drought strongly influences plant productivity, suggesting that water limitation has shaped the evolution of many plant physiological traits. One functional strategy that plants employ to cope with decreasing water availability is drought escape. For drought-escaping species, high metabolic activity (gas exchange) and rapid growth are hypothesized to confer a fitness advantage, because this enables a plant to complete its life cycle before the most intense period of drought. By growing an annual grass species (Avena barbata) under well-watered or water-limited conditions in a greenhouse, we directly tested whether high photosynthesis, increased stomatal opening, and early flowering are adaptive under drought. We measured phenotypic selection on instantaneous gas exchange and flowering time as well as the underlying biochemical traits that regulate photosynthesis. We found strong selection for earlier flowering in the dry environment, but no evidence that increased photosynthesis was adaptive under drought. Photosynthetic rate (A) and stomatal conductance (g_s) were both adaptively neutral in the dry environment. Increased photosynthetic capacity (A_max) was maladaptive in the dry environment, perhaps because of the respiratory cost associated with maintaining excess enzyme and substrate capacity. There was no correlational selection on the combination of physiology and flowering time in the dry environment, suggesting that accelerated development and high gas exchange may not need to be tightly linked to promote drought escape. In contrast, there was selection for both high photosynthetic function (A_max and A) and early flowering in the well-watered environment. These combinations of traits may have been favored because they maximize both energy and time available for reproduction. Our results suggest that the benefit of increased photosynthesis for plant fitness may be strongest in the absence of drought stress.

Variation in ontogenetic development among individuals may be a major contributor to morphological variation within species. Evolution of different growth trajectories might, for example, evolve as a response to varying ecological contexts of individuals living in different environments, or by life-stage or gender differences. The intertidal periwinkle Littorina saxatilis is strongly polymorphic in shell shape. We compared ontogenetic trajectories between life stages, local populations, and sexes to understand how different morphological end points are reached during ontogeny and what might cause these differences. Applying landmark-based geometric morphometrics, we captured shell shape variation for four Swedish populations of this species. We also derived a method to visualize ontogenetic trajectories described by the relationship of size to the multivariate shape space. We found that growth trajectories differed between individuals living in different habitats, as well as between sexes and maturity stages. Males living on rocky cliffs grew isometrically throughout life, whereas females from the same habitat switched from isometric growth as juveniles to allometric growth as adults. In contrast, males and females living on boulders grew allometrically as juveniles but changed to isometric growth at sexual maturity. Thus, in this species, ontogenetic growth seems influenced by habitat-associated selection as well as by gender and age-specific selection. These differing selection regimes result in ontogenetic shifts in allometry in three of the four groups examined.

The view that marine species with planktonic dispersal have highly connected, demographically open populations is giving way to recognition that populations may often be largely self-recruiting, or demographically closed. This raises the question of what local conditions might favor isolation of populations. To test the importance of islands for local isolation in species with planktonic larvae, we examined allozyme variation among 35 populations of the intertidal snail Austrocochlea constricta in the Houtman Abrolhos Islands, Western Australia, spanning 60 km. Heterogeneity of allozyme frequencies among populations was high, with average F_ST of 0.237, indicating highly localized populations. Increased subdivision was associated with islands at different scales: between island groups, separated by deep water gaps, and between disconnected sets of islands within groups. At short distances, up to two km, subdivision increased fivefold between islands compared with that between populations on the same island. Along 11 km of continuous, sheltered shore, there was isolation by distance but among a linear series of islands over similar distance, there was greater subdivision at short distances but no association with distance. These patterns had been seen previously in the direct-developing snail Bembicium vittatum, but its finding in A. constricta confirms for a planktonic disperser the importance of this complex archipelago for both retention of locally produced larvae and disruption of patterns of connectivity. Taken together, these results indicate that islands can increase both the “open” and the “closed” components of recruitment and that applicable models of genetic connectivity depend substantially on local conditions.

The Alpheidae—possibly the most diverse family of recent decapod crustaceans—offers attractive opportunities to study the evolution of many intriguing phenomena, including key morphological innovations like spectacular snapping claws, highly specialized body forms, facultative and obligate symbioses with many animal groups, and sophisticated behaviors like eusociality. However, studies of these remarkable adaptations remain hampered by insufficient phylogenetic information. We present the first phylogenetic hypothesis of relationships among 36 extant genera of alpheid shrimps, based on a cladistic analysis of 122 morphological characters from 56 species, and we use this hypothesis to explore evolutionary trends in morphology and species diversity. Our results strongly supported a monophyletic Alpheidae that included two hitherto difficult-to-place genera (Yagerocaris and Pterocaris). Of 35 nodes among genera, all were supported by at least one morphological character (24 were supported by two or more) and 17 received greater than 50% jackknife support. Unfortunately, many basal nodes were only weakly supported. Six genera appeared nonmonophyletic, including the dominant genus Alpheus (paraphyletic due to inclusion of one clade with three minor genera). Evolutionary trends in alpheid claw form shed some revealing light on how key innovations evolve. First, several functionally significant features of the cheliped (claw bearing leg) evolved independently multiple times, including: asymmetry, folding, inverted orientation, sexual dimorphism, adhesive plaques that enhance claw cocking, and tooth-cavity systems on opposing claw fingers, a preadaptation for snapping. Many conspicuous features of alpheid claw form therefore appear prone to parallel evolution. Second, although tooth-cavity systems evolved multiple times, a functional snapping claw, which likely facilitated an explosive radiation of over 550 species, evolved only once (in Synalpheus [Alpheus satellite genera]). Third, adhesive plaques (claw cocking aids) also evolved multiple times, and within snapping alpheids are associated with the most diverse clade (Alpheus derivative genera). This pattern of parallel preadaptation—multiple independent evolutionary origins of precursors (preadaptations) to what ultimately became a key innovation (adaptation)—suggests alpheid shrimp claws are predisposed to develop features like tooth-cavity and adhesive plaque systems for functional or developmental reasons. Such functional/developmental predisposition may facilitate the origin of key innovations. Finally, moderate orbital hoods—anterior projections of the carapace partly or completely covering the eyes—occur in many higher Alpheidae and likely evolved before snapping claws. They are unique among decapod crustaceans, and their elaboration in snapping alpheids suggests they may protect the eyes from the stress of explosive snaps. Thus one key innovation (orbital hoods) may have facilitated evolution of a second (snapping claws).

The molecular chaperone protein Hsp90 has been widely discussed as a candidate gene for developmental buffering. We used the methods of geometric morphometrics to analyze its effects on the variation among individuals and fluctuating asymmetry of wing shape in Drosophila melanogaster. Three different experimental approaches were used to reduce Hsp90 activity. In the first experiment, developing larvae were reared in food containing a specific inhibitor of Hsp90, geldanamycin, but neither individual variation nor fluctuating asymmetry was altered. Two further experiments generated lines of genetically identical flies carrying mutations of Hsp83, the gene encoding the Hsp90 protein, in heterozygous condition in nine different genetic backgrounds. The first of these, introducing entire chromosomes carrying either of two Hsp83 mutations, did not increase shape variation or asymmetry over a wild-type control in any of the nine genetic backgrounds. In contrast, the third experiment, in which one of these Hsp83 alleles was introgressed into the wild-type background that served as the control, induced an increase in both individual variation and fluctuating asymmetry within each of the nine genetic backgrounds. No effect of Hsp90 on the difference among lines was detected, providing no evidence for cryptic genetic variation of wing shape. Overall, these results suggest that Hsp90 contributes to, but is not controlling, the buffering of phenotypic variation in wing shape.

The evolution of biological materials is a critical, yet poorly understood, component in the generation of biodiversity. For example, the diversification of spiders is correlated with evolutionary changes in the way they use silk, and the material properties of these fibers, such as strength, toughness, extensibility, and stiffness, have profound effects on ecological function. Here, we examine the evolution of the material properties of dragline silk across a phylogenetically diverse sample of species in the Araneomorphae (true spiders). The silks we studied are generally stronger than other biological materials and tougher than most biological or man-made fibers, but their material properties are highly variable; for example, strength and toughness vary more than fourfold among the 21 species we investigated. Furthermore, associations between different properties are complex. Some traits, such as strength and extensibility, seem to evolve independently and show no evidence of correlation or trade-off across species, even though trade-offs between these properties are observed within species. Material properties retain different levels of phylogenetic signal, suggesting that traits such as extensibility and toughness may be subject to different types or intensities of selection in several spider lineages. The picture that emerges is complex, with a mosaic pattern of trait evolution producing a diverse set of materials across spider species. These results show that the properties of biological materials are the target of selection, and that these changes can produce evolutionarily and ecologically important diversity.

When social interactions occur, the phenotype of an individual is influenced directly by its own genes (direct genetic effects) but also indirectly by genes expressed in social partners (indirect genetic effects). Social insect colonies are characterized by extensive behavioral interactions among workers, brood, and queens so that indirect genetic effects are particularly relevant. I used a series of experimental manipulations to disentangle the contribution of direct effects, maternal (queen) effects, and sibsocial (worker) effects to variation for worker, gyne, and male mass; caste ratio; and sex ratio in the ant Temnothorax curvispinosus. The results indicate genetic variance for direct, maternal, and sibsocial effects for all traits, except for male mass there was no significant maternal variance, and for sex ratio the variance for direct effects was not separable from maternal variance for the primary sex ratio. Estimates of genetic correlations between direct, maternal, and sibsocial effects were generally negative, indicating that these effects may not evolve independently. These results have broad implications for social insect evolution. For example, the genetic architecture underlying social insect traits may constrain the realization of evolutionary conflicts between social partners.

Our understanding of the evolution of genes of the major histocompatibility complex (MHC) is rapidly increasing, but there are still enigmatic questions remaining, particularly regarding the maintenance of high levels of MHC polymorphisms in small, isolated populations. Here, we analyze the genetic variation at eight microsatellite loci and sequence variation at exon 2 of the MHC class IIB (DAB) genes in two wild populations of the Trinidadian guppy, Poecilia reticulata. We compare the genetic variation of a small (N_e ≈ 100) and relatively isolated upland population to that of its much larger (N_e ≈ 2400) downstream counterpart. As predicted, microsatellite diversity in the upland population is significantly lower and highly differentiated from the population further downstream. Surprisingly, however, these guppy populations are not differentiated by MHC genetic variation and show very similar levels of allelic richness. Computer simulations indicate that the observed level of genetic variation can be maintained with overdominant selection acting at three DAB loci. The selection coefficients differ dramatically between the upland (s ≥ 0.2) and lowland (s ≤ 0.01) populations. Parasitological analysis on wild-caught fish shows that parasite load is significantly higher on upland than on lowland fish, which suggests that large differences in selection intensity may indeed exist between populations. Based on the infection intensity, a substantial proportion of the upland fish would have suffered direct or indirect fitness consequences as a result of their high parasite loads. Selection by parasites plays a particularly important role in the evolution of guppies in the upland habitat, which has resulted in high levels of MHC diversity being maintained in this population despite considerable genetic drift.

Morphological diversity is routinely used to infer ecological variation among species because differences in form underlie variation in functional performance of ecological tasks like capturing prey, avoiding predators, or defending territories. However, many functions have complex morphological bases that can weaken associations between morphological and functional diversification. We investigate the link between these levels of diversity in a mechanically explicit model of fish suction-feeding performance, where the map of head morphology to feeding mechanics is many-to-one: multiple, alternative forms can produce the same mechanical property. We show that many-to-one mapping leads to discordance between morphological and mechanical diversity in the freshwater fish family, the Centrarchidae, despite close associations between morphological changes and their mechanical effects. We find that each of the model's five morphological variables underlies evolution of suction capacity. Yet, the major centrarchid clades exhibit an order of magnitude range in diversity of suction mechanics in the absence of any clear difference in diversity of the morphological variables. This cryptic pattern of mechanical diversity suggests an evolutionary history for suction performance that is unlike the one inferred from comparisons of morphological diversity. Because many-to-one mapping is likely to be common in functional systems, this property of design may lead to widespread discordance between functional and morphological diversity. Although we focus on the interaction between morphology and mechanics, many-to-one mapping can decouple diversity between levels of organization in any hierarchical system.

The history of life has been marked by several spectacular radiations, in which many lineages arise over a short period of time. A possible consequence of such rapid splitting in the recent past is that the intrinsic barriers that prevent gene flow between many species may have too little time to develop fully, leading to extensive hybridization among recently evolved lineages. The salamander genus Plethodon in eastern North America has been proposed as a possible example of this scenario, but without explicit statistical tests. In this paper, we present a nearly comprehensive phylogeny for the 45 extant species of eastern Plethodon, based on DNA sequences of mitochondrial (two genes, 1335 base pairs) and nuclear genes (two genes, up to 3481 base pairs). We then use this phylogeny to examine rates and patterns of diversification and hybridization. We find significantly rapid diversification within the glutinosus species group. Examining patterns of natural hybridization in light of the phylogeny shows considerable hybridization within this clade, including introgression between species that are morphologically distinct and distantly related. Reproductive isolation increases over time and may be very weak among the most recently diverged species. These results suggest that the origin of species and the evolution of intrinsic reproductive isolating mechanisms, rather than being synonymous, may be decoupled in some cases (i.e., rapid origin of lineages outstrips the “speciation clock”). In contrast to the conclusions of a recent review of adaptive radiation and hybridization, we suggest that extensive hybridization sometimes may be a consequence, rather than a cause, of rapid diversification.

Recent speciation research has generally focused on how lineages that originate in allopatry evolve intrinsic reproductive isolation, or how ecological divergence promotes nonallopatric speciation. However, the ecological basis of allopatric isolation, which underlies the most common geographic mode of speciation, remains poorly understood and largely unstudied. Here, we explore the ecological and evolutionary factors that promote speciation in Desmognathus and Plethodon salamanders from temperate eastern North America. Based on published molecular phylogenetic estimates and the degree of geographic range overlap among extant species, we find strong evidence for a role for geographic isolation in speciation. We then examine the relationship between climatic variation and speciation in 16 sister-taxon pairs using geographic information system maps of climatic variables, new methods for modeling species' potential geographic distributions, and data on geographic patterns of genetic variation. In contrast to recent studies in tropical montane regions, we found no evidence for parapatric speciation along climatic gradients. Instead, many montane sister taxa in the Appalachian Highlands inhabit similar climatic niches and seemingly are allopatric because they are unable to tolerate the climatic conditions in the intervening lowlands. This temporal and spatial-ecological pattern suggests that niche conservatism, rather than niche divergence, plays the primary role in promoting allopatric speciation and montane endemism in this species-rich group of vertebrates. Our results demonstrate that even the relatively subtle climatic differences between montane and lowland habitats in eastern North America may play a key role in the origin of new species.

Interpopulation variation in morphology, such as that among small island populations, plays a key role in speciation and diversification. There are two approaches to investigating evolution of morphological characters: comparing patterns of trait variances and covariances within and among populations, and testing particular adaptive scenarios. Here, we combine both approaches to infer the role of natural selection in shaping morphological variation in body size, head color pattern, and body shape among 10 populations of a day gecko, Phelsuma ornata, and its close relative, P. inexpectata, in the Mascarene Islands. We find that local populations are morphologically distinct, and that natural selection has likely influenced phenotypic diversification in the group. Lizards on small outer islands tend to be larger than lizards on the mainland of Mauritius. For body shape and head color pattern, comparisons of variation within and among populations reveal that differences among populations for some variables are too great to be explained by neutral processes alone, although we cannot identify the causal agents for this selection. These results reveal that the forces shaping different sets of organismal traits may be distinct, such that a variety of statistical approaches are needed to investigate selection in natural populations.

Sensory exploitation occurs when signals trigger behavioral reactions that diminish the receiver's fitness. Research in this area focuses on the match between the signal's form and the receiver's sensitivity, but the effect of habitat on interspecific sensory exploitation is rarely addressed. Myioborus redstarts use conspicuous wing and tail displays of contrasting black-and-white plumage patches to flush dipteran insects, which are then pursued and captured in flight. Previous studies have shown that by increasing the distance at which insects perform an escape response, conspicuous visual displays improve the birds' foraging performance. We tested the hypothesis that selection for a visual signal that maximizes prey escape distance under local habitat conditions can lead to the evolution of geographic variation in plumage pattern among Myioborus redstarts. Using models of foraging birds, we recorded the escape responses of Dipterous insects to a range of plumage patterns and background tones (from light to dark) to determine whether the plumage pattern that maximizes prey flushing is dependent upon that habitat (background) against which birds are viewed by their prey. Our results indicate that the effectiveness of a particular plumage pattern in flushing dipteran prey depends strongly on the background against which that plumage pattern is displayed, and darker habitat (background) conditions generally favor plumages with more extensive patches of white in the tail. However, the addition of white wing patches that imitate the plumage of the painted redstart (Myioborus pictus) generally increases insect escape responses but reduces the effect that tail pattern variation and background tone have on escape behavior. These experiments support the hypothesis that habitat-specific natural selection to enhance sensory exploitation of prey escape responses could produce geographic variation in plumage patterns of flush-pursuers.

Why do all mammals, except for sloths and manatees, have exactly seven cervical vertebrae? In other vertebrates and other regions, the vertebral number varies considerably. We investigated whether natural selection constrains the number of cervical vertebrae in humans. To this end, we determined the incidence of cervical ribs and other homeotic vertebral changes in radiographs of deceased human fetuses and infants, and analyzed several existing datasets on the incidence in infants and adults. Our data show that homeotic transformations that change the number of cervical vertebrae are extremely common in humans, but are strongly selected against: almost all individuals die before reproduction. Selection is most probably indirect, caused by a strong coupling of such changes with major congenital abnormalities. Changes in the number of thoracic vertebrae appear to be subject to weaker selection, in good correspondence with the weaker evolutionary constraint on these numbers. Our analysis highlights the role of prenatal selection in the conservation of our common body plan.

The pattern and extent of pleiotropic gene action can contribute substantially to the internal structure and shape of the additive genetic variance-covariance matrix (G)—a key determinant of evolutionary trajectories. We use data from our study (Estes et al. 2004) on the univariate effects of mutation in a mismatch-repair-defective strain, msh-2, of Caenorhabditis elegans to address the impact of increasing levels of selection on the magnitude and pattern of genetic covariance due to new mutations. Mutational covariances between three life-history traits are shown to exhibit a weak pattern of decline with increasing population size (increasing selection), while the orientation of mutational matrices remains reasonably constant. This suggests that mutations with smaller effects on fitness may tend to be slightly more confined in their influence than large-effect mutations (i.e., small-effect mutations reduce the magnitude of covariation between characters), but do not change the direction of this covariation.