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Raia et al. propose that the evolution of the shape and complexity of animal ornaments (e.g., deer antlers) can be explained by interspecific variation in body size and is not influenced by sexual selection. They claim to show that ornament complexity is related to body size by an 0.25-power law and argue that this finding precludes a role for sexual selection in the evolution of ornament complexity. However, their study does not test alternative hypotheses and mismeasures antler shape allometry by omitting much of the published data. We show that an index of sexual selection (sexual size dimorphism) is positively correlated with size-corrected antler complexity and that the allometric slope of complexity is substantially greater than 0.25, contra Raia et al. We conclude that sexual selection and physical constraints both affect the evolution of antler shape. [ABSTRACT FROM AUTHOR]

Genetic rescue, an increase in population growth owing to the infusion of new alleles, can aid the persistence of small populations. Its use as a management tool is limited by a lack of empirical data geared toward predicting effects of gene flow on local adaptation and demography. Experimental translocations provide an ideal opportunity to monitor the demographic consequences of gene flow. In this study we take advantage of two experimental introductions of Trinidadian guppies to test the effects of gene flow on downstream native populations. We individually marked guppies from the native populations to monitor population dynamics for 3 months before and 26 months after gene flow. We genotyped all individuals caught during the first 17 months at microsatellite loci to classify individuals by their genetic ancestry: native, immigrant, F1 hybrid, F2 hybrid, or backcross. Our study documents a combination of demographic and genetic rescue over multiple generations under fully natural conditions. Within both recipient populations, we found substantial and long-term increases in population size that could be attributed to high survival and recruitment caused by immigration and gene flow from the introduction sites. Our results suggest that low levels of gene flow, even from a divergent ecotype, can provide a substantial demographic boost to small populations, which may allow them to withstand environmental stochasticity. [ABSTRACT FROM AUTHOR]

Aim: The theory of island biogeography provides a predictive framework relating species richness to island size and distance from the mainland. However, the theory as originally formulated does not necessarily scale to large islands and continental landmasses that are capable of generating species through in situ speciation (rather than entirely by colonization), nor does it necessarily account for how human introduction of species alters traditional biogeographical patterns. Here, we examine the ecological (colonization and extinction), evolutionary (in situ speciation) and human-mediated (deliberate introductions) determinants of species richness in a taxonomic group that has undergone a radiation on Caribbean islands: live-bearing fishes of the family Poeciliidae. Location: The Caribbean. Methods: We created a database of both native and introduced poeciliid species occurrence on Caribbean islands through literature review, and estimated the number of colonizations versus speciation events on each island using a molecular phylogeny. Linear regression and other statistical tests were used to explore species-area and species-isolation relationships. Results: Species richness on small islands results entirely from colonization and does not significantly increase with island area, whereas on larger islands species richness increases dramatically as a function of area due primarily to in situ speciation. Poeciliid fishes have been introduced widely, both as a byproduct of their popularity in the aquarium hobby and as a means of mosquito control. We show that such establishments have occurred disproportionately on islands depauperate in native species, and that introduced species richness is positively correlated with economic interconnectedness (shipping traffic) and human population size. Main conclusions: On large Caribbean islands in situ speciation has elevated the number of poeciliid species beyond that predicted from ecological processes alone. Introduced species significantly alter biogeographical patterns. [ABSTRACT FROM AUTHOR]

The livebearing Trinidadian guppy ( Poecilia reticulata) produces bigger offspring in populations exposed to low predation and produces smaller, more numerous offspring in populations subject to high predation (HP)., Like most fishes, guppies respond to predator attacks with a fast-start escape response. From the scaling of teleost fast-start performance, we predict that larger guppy neonates should exhibit faster, more effective escape responses than smaller neonates. Increasing performance with increasing size could be due simply to size, or to both size and the acquisition of more mature body forms, as is seen in larval-stage fishes., We find no difference in external body proportions among guppy offspring varying in size from 5·1 mm to 7·1 mm at birth, suggesting offspring are born morphologically mature. However, based on the degree of skeletal ossification, as a proxy for internal maturity, we find that guppies are still maturing rapidly around the time of birth. The smallest neonates from the very HP Caroni confluence lack ossification of key skeletal elements that are present in their larger low predation counterparts., In guppies from the Aripo/Caroni drainage, we show that neonatal escape performance covaries with responsiveness and increases with size along a gradient of HP offspring, but is lower in the largest low predation offspring. The scaling of escape performance in the HP populations performing maximally (scaling with length as L2·76) exceeds predictions from size alone (scaling exponent of L1·0 indicates performance increases linearly with body size). This suggests that guppy neonates, like rapidly developing larval fishes, vary substantially in morphological maturity among populations., The finding that neonatal guppy offspring covary in both size and maturity at birth means that being smaller also means being less mature, which amplifies the negative escape performance effects of being born small. Despite the negative consequences of being born both small and immature, HP environments select heavily for high fecundity, and thus small offspring size. We find selection favours female life-history traits over offspring escape performance. [ABSTRACT FROM AUTHOR]

Mate choice is a common feature of sexually reproducing species. In sessile or sedentary external fertilizers, however, direct interactions between reproductive partners are minimal, and instead mate recognition and choice must occur at the level of gametes. It is common for some sperm and egg combinations to have higher fertilization success than others, but it remains unclear whether differences in fertilization reflect gamete-level mate choice (GMC) for paternal quality or parental compatibility. Here, we examine the mechanisms underlying GMC in an externally fertilizing ascidian. A manipulative mate-choice assay confirmed that offspring viability was greater in clutches where we allowed GMC than in clutches where we precluded GMC. A complementary quantitative genetic experiment then revealed that paternal quality effects were generally weaker than parental compatibility effects, particularly for the trait combination underlying the benefits of GMC. Overall, our data suggest that gametes that are more compatible at fertilization produce more viable offspring than gametes that are less compatible at fertilization. Therefore, although the regalia we typically associate with sexual selection are absent in external fertilizers, mechanisms that allow females to bias fertilization in favor of some males over others produce significant fitness benefits in organisms reproducing via the ancestral strategy. [ABSTRACT FROM AUTHOR]

Environments that vary within a generation of an organism provide opportunities for adaptation if the level of variation is severe and predictable. We describe a model of evolution in such environments with genotypes that show trade-offs in viability and fecundity. One genotype develops rapidly and has superior viability but reduced fertility relative to the alternative genotype. Conditions that allow the evolution of the rapidly developing genotypes are explored. We show how the evolution of ovoviviparity and resource specialization in Drosophila sechellia shares many important features of this model. We suggest that our model may capture many of the evolutionary forces responsible for the evolution of niche specialization and ovoviviparity seen in D. sechellia. [ABSTRACT FROM AUTHOR]

Selection on naturally occurring hybrid individuals is a key component of speciation theory, but few studies examine the functional basis of hybrid performance. We examine the functional consequences of hybridization in nature, using the freshwater sunfishes (Centrarchidae), where natural hybrids have been studied for more than a century and a half. We examined bluegill (Lepomis macrochirus), green sunfish (Lepomis cyanellus), and their naturally occurring hybrid, using prey-capture kinematics and morphology to parameterize suction-feeding simulations on divergent parental resources. Hybrid individuals exhibited kinematics intermediate between those of the two parental species. However, performance assays indicated that hybrids display performance most similar to the worse-performing species for a given parental resource. Our results show that intermediate hybrid phenotypes can be impaired by a lessthan- intermediate performance and hence suffer a larger loss in fitness than could be inferred from morphology alone. [ABSTRACT FROM AUTHOR]

Phenotypic plasticity is the capacity for an individual genotype to produce different phenotypes in response to environmental variation. Most traits are plastic, but the degree to which plasticity is adaptive or non-adaptive depends on whether environmentally induced phenotypes are closer or further away from the local optimum. Existing theories make conflicting predictions about whether plasticity constrains or facilitates adaptive evolution. Debate persists because few empirical studies have tested the relationship between initial plasticity and subsequent adaptive evolution in natural populations. Here we show that the direction of plasticity in gene expression is generally opposite to the direction of adaptive evolution. We experimentally transplanted Trinidadian guppies (Poecilia reticulata) adapted to living with cichlid predators to cichlid-free streams, and tested for evolutionary divergence in brain gene expression patterns after three to four generations. We find 135 transcripts that evolved parallel changes in expression within the replicated introduction populations. These changes are in the same direction exhibited in a native cichlid-free population, suggesting rapid adaptive evolution. We find 89% of these transcripts exhibited non-adaptive plastic changes in expression when the source population was reared in the absence of predators, as they are in the opposite direction to the evolved changes. By contrast, the remaining transcripts exhibiting adaptive plasticity show reduced population divergence. Furthermore, the most plastic transcripts in the source population evolved reduced plasticity in the introduction populations, suggesting strong selection against non-adaptive plasticity. These results support models predicting that adaptive plasticity constrains evolution, whereas non-adaptive plasticity potentiates evolution by increasing the strength of directional selection. The role of non-adaptive plasticity in evolution has received relatively little attention; however, our results suggest that it may be an important mechanism that predicts evolutionary responses to new environments. [ABSTRACT FROM AUTHOR]

Luxuriant, bushy antlers, bizarre crests, and huge, twisting horns and tusks are conventionally understood as products of sexual selection. This view stems from both direct observation and from the empirical finding that the size of these structures grows faster than body size (i.e., ornament size shows positive allometry).We contend that the familiar evolutionary increase in the complexity of ornaments over time in many animal clades is decoupled from ornament size evolution. Increased body size comes with extended growth. Since growth scales to the quarter power of body size, we predicted that ornament complexity should scale according to the quarter power law as well, irrespective of the role of sexual selection in the evolution and function of the ornament. To test this hypothesis, we selected three clades (ammonites, deer, and ceratopsian dinosaurs) whose species bore ornaments that differ in terms of the importance of sexual selection to their evolution.We found that the exponent of the regression of ornament complexity to body size is the same for the three groups and is statistically indistinguishable from0.25.We suggest that the evolution of ornament complexity is a by-product of Cope's rule. We argue that although sexual selection may control size in most ornaments, it does not influence their shape. [ABSTRACT FROM AUTHOR]

Decades of theory and recent empirical results have shown that evolutionary, population, community and ecosystem properties are the result of feedbacks between ecological and evolutionary processes. The vast majority of theory and empirical research on these eco-evolutionary feedbacks has focused on interactions among population size and mean traits of populations., However, numbers and mean traits represent only a fraction of the possible feedback dimensions. Populations of many organisms consist of different size classes that differ in their impact on the environment and each other. Moreover, rarely do we know the map of ecological pathways through which changes in numbers or size structure cause evolutionary change. The goal of this study was to test the role of size structure in eco-evolutionary feedbacks of Trinidadian guppies and to begin to build an eco-evolutionary map along this unexplored dimension., We used a factorial experiment in mesocosms wherein we crossed high- and low-predation guppy phenotypes with population size structure. We tested the ability of changes in size structure to generate selection on the demographic rates of guppies using an integral projection model ( IPM). To understand how fitness differences among high- and low-predation phenotypes may be generated, we measured the response of the biomass of lower trophic levels and nutrient cycling to the different phenotype and size structure treatments., We found a significant interaction between guppy phenotype and the size structure treatments for absolute fitness. Size structure had a very large effect on invertebrate biomass in the mesocosms, but there was little or no effect of the phenotype. The effect of size structure on algal biomass depended on guppy phenotype, with no difference in algal biomass in populations with more, smaller guppies, but a large decrease in algal biomass in mesocosms with phenotypes adapted to low-predation risk., These results indicate an important role for size structure partially driving eco-evolutionary feedbacks in guppies. The changes in the ecosystem suggest that the absence of a steep decline in guppy fitness of the low-predation risk populations is likely due to higher consumption of algae when invertebrates are comparatively rare. Overall, these results demonstrate size structure as a possible dimension through which eco-evolutionary feedbacks may occur in natural populations. [ABSTRACT FROM AUTHOR]

Two ways in which organisms adapt to variable environments are phenotypic plasticity and bet-hedging. Theory suggests that bet-hedging is expected to evolve in unpredictable environments for which reliable cues indicative of future conditions (or season length) are lacking. Alternatively, if reliable cues exist indicating future conditions, organisms will be under selection to produce the most appropriate phenotype -that is, adaptive phenotypic plasticity. Here, we experimentally test which of these modes of adaptation are at play in killifish that have evolved an annual life cycle. These fish persist in ephemeral pools that completely dry each season through the production of eggs that can remain in developmental arrest, or diapause, buried in the soil, until the following rainy season. Consistent with diversified bet-hedging (a risk spreading strategy), we demonstrate that the eggs of the annual killifish Nothobranchius furzeri exhibit variation at multiple levels-whether or not different stages of diapause are entered, for how long diapause is entered, and the timing of hatching-and this variation persists after controlling for both genetic and environmental sources of variation. However, we show that phenotypic plasticity is also present in that the proportion of eggs that enter diapause is influenced by environmental factors (temperature and light level) that vary seasonally. In nature there is typically a large parameter zone where environmental cues are somewhat correlated with seasonality, but not perfectly so, such that it may be advantageous to have a combination of both bet-hedging and plasticity. [ABSTRACT FROM AUTHOR]

The evolution of convergent phenotypes in lineages subject to similar selective pressures is a common feature of adaptive radiation. In geographically replicated radiations, repeated convergence occurs between clades occupying distinct regions or islands. Alternatively, a clade may repeatedly reach the same adaptive peaks in broadscale sympatry, resulting in extensive convergence within a region. Rockfish (Sebastes sp.) have radiated in both the northeast and northwest Pacific, allowing tests of the extent and geographic pattern of convergence in a marine environment. We used a suite of phylogenetically informed methods to test for morphological convergence in rockfish. We examined patterns of faunal similarity using nearest neighbor distances in morphospace and the frequency of morphologically similar yet distantly related species pairs. The extent of convergence both between regions and within the northeast Pacific exceeds the expectation under a Brownian motion null model, although constraints on trait space could account for the similarity. We then used a recently developed method (SURFACE) to identify adaptive peak shifts in Sebastes evolutionary history. We found that the majority of convergent peak shifts occur within the northeast Pacific rather than between regions and that the signal of peak shifts is strongest for traits related to trophic morphology. Pacific rockfish thus demonstrate a tendency toward morphological convergence within one of the two broad geographic regions in which they have diversified. [ABSTRACT FROM AUTHOR]

Most of what we know about placentas comes from mammals, yet little can be learned from them about the adaptive significance of the placental mode of reproduction because they all derived their placenta from a single common ancestor that lived over 100 million years ago. We can make inferences about the adaptive significance of placentation from fish in the family Poeciliidae because there have been multiple, recent origins of placentation, affording an opportunity to compare close relatives with and without placentas and to seek properties that are common to each origin of placentation., Here, we used field collections and a common garden study to quantify the degree of placentation and related it to aspects of the life history in two clades of live-bearing fish from the genus Poeciliopsis that each contains an independent origin of placentation. Doing so enables us to test the 'life history facilitation hypothesis', or the proposal that the placenta evolved to facilitate the evolution of some other feature of the life history., We found that the evolution of placentation in each clade is tightly correlated with the evolution of other components of the life history, but that the nature of the association is radically different across the two clades. In the Northern Clade the magnitude of post-fertilization maternal provisioning is negatively correlated with age at maturity, mass at maturity, offspring dry mass and interlitter interval. In contrast, degree of matrotrophy in the Southern Clade is positively correlated with age at maturity, mass at maturity, offspring dry mass and inter-litter interval., There is thus no consistent relationship between the evolution of placentas and other features of the life history, which negates those proposals that the placenta evolved to facilitate the evolution of other features of the life history. However, there is a negative correlation between degree of placentation and ovary dry mass and reproductive allocation common to both clades, suggesting that placentation may be an adaptation that facilitates a reduction in reproductive allocation. [ABSTRACT FROM AUTHOR]

Traits that increase mating success in males may come at a cost, such as an increased risk of predation. However, predatormediated selection is challenging to document in natural populations, hampering our understanding of the trade-offs between sexual selection and predation. Here we report on a study of predator- mediated natural selection on wing traits in the damselfly Calopteryx splendens, the males of which possess conspicuous wing patches. Wagtails (genus Motacilla) are important avian predators of C. splendens, capturing them in flight and removing the wings prior to consumption. Using geometric morphometric techniques, we quantified the strength and mode of selection on wing traits by comparing wings from depredated individuals with the standing variation present in the population. Our results reveal that predator- mediated selection is stronger on secondary sexual characters than on size and shape, suggesting that traits related to flight performance are closer to their adaptive peaks. This could be a consequence of the long-term evolutionary association with avian predators, whereas stronger selection on conspicuous secondary sexual traits may reflect trade-offs between sexual and natural selection. Finally, even though C. splendens possesses nearly identical fore- and hindwings, we found evidence for divergent selection between them. [ABSTRACT FROM AUTHOR]

Sexual selection can increase morphological diversity within and among species. Little is known regarding how interspecific variation produced through sexual selection affects other functional systems. Here, we examine how morphological diversity resulting from sexual selection impacts aerobic locomotor performance. Using Xiphophorus (swordtail fish) and their close relatives ( N = 19 species), we examined whether the evolution of a longer sexually selected sword affects critical swimming speed. We also examined the effect of other suborganismal, physiological, and morphological traits on critical swimming speed, as well as their relationship with sword length. In correlation analyses, we found no significant relationship between sword length and critical swimming speed. Unexpectedly, we found that critical swimming speed was higher in species with longer swords, after controlling for body size in multiple regression analyses. We also found several suborganismal and morphological predictors of critical swimming speed, as well as a significant negative relationship between sword length and heart and gill mass. Our results suggest that interspecific variation in sword length is not costly for this aspect of swimming performance, but further studies should examine potential costs for other types of locomotion and other components of Darwinian fitness (e.g., survivorship, life span). [ABSTRACT FROM AUTHOR]

Novel environments often impose directional selection for a new phenotypic optimum. Novel environments, however, can also change the distribution of phenotypes exposed to selection by inducing phenotypic plasticity. Plasticity can produce phenotypes that either align with or oppose the direction of selection. When plasticity and selection are parallel, plasticity is considered adaptive because it provides a better pairing between the phenotype and the environment. If the plastic response is incomplete and falls short of producing the optimum phenotype, synergistic selection can lead to genetic divergence and bring the phenotype closer to the optimum. In contrast, non-adaptive plasticity should increase the strength of selection, because phenotypes will be further from the local optimum, requiring antagonistic selection to overcome the phenotype–environment mismatch and facilitate adaptive divergence. We test these ideas by documenting predator-induced plasticity for resting metabolic rate and growth rate in populations of the Trinidadian guppy (Poecilia reticulata) adapted to high and low predation. We find reduced metabolic rates and growth rates when cues from a predator are present during development, a pattern suggestive of adaptive and non-adaptive plasticity, respectively. When we compared populations recently transplanted from a high-predation environment into four streams lacking predators, we found evidence for rapid adaptive evolution both in metabolism and growth rate. We discuss the implications for predicting how traits will respond to selection, depending on the type of plasticity they exhibit. [ABSTRACT FROM AUTHOR]