Density-dependent selection, which promotes contrasting patterns of trait means at different population densities, has a long history in population genetics and ecology. The unifying principle from theory is that density-dependent selection operates on phenotypic traits whose values counter the effects of whatever ecological agent is limiting population growth, be it resource competition, predators, or pathogens. However, the complexity inherent in density dependence means that the same selective process can generate multiple outcomes, depending upon the details of how population density affects vital rates and the age or size structure of a population. Failure to appreciate the potential for multiple outcomes confounded many early studies of the process. Nonetheless, careful empirical work in laboratory studies, long-term field studies, and studies of sexual selection demonstrates the wide reach of density-dependent selection. The inconsistent outcomes observed in these studies call for renewed research into how the details of density dependence channel adaptive responses.
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Abstract The genetic basis of traits shapes and constrains how adaptation proceeds in nature; rapid adaptation can proceed using stores of polygenic standing genetic variation or hard selective sweeps, and increasing polygenicity fuels genetic redundancy, reducing gene re-use (genetic convergence). Guppy life history traits evolve rapidly and convergently among natural high- and low-predation environments in northern Trinidad. This system has been studied extensively at the phenotypic level, but little is known about the underlying genetic architecture. Here, we use four independent F2 QTL crosses to examine the genetic basis of seven (five female, two male) guppy life history phenotypes and discuss how these genetic architectures may facilitate or constrain rapid adaptation and convergence. We use RAD-sequencing data (16,539 SNPs) from 370 male and 267 female F2 individuals. We perform linkage mapping, estimates of genome-wide and per-chromosome heritability (multi-locus associations), and QTL mapping (single-locus associations). Our results are consistent with architectures of many loci of small-effect for male age and size at maturity and female interbrood period. Male trait associations are clustered on specific chromosomes, but female interbrood period exhibits a weak genome-wide signal suggesting a potentially highly polygenic component. Offspring weight and female size at maturity are also associated with a single significant QTL each. These results suggest rapid, repeatable phenotypic evolution of guppies may be facilitated by polygenic trait architectures, but subsequent genetic redundancy may limit gene re-use across populations, in agreement with an absence of strong signatures of genetic convergence from recent analyses of wild guppies.more » « less
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Abstract The placenta is a complex organ that shows high morphological diversity. Among fish, the first vertebrates that have evolved a placenta, the family Poeciliidae exhibits very diverse modes of maternal provisioning even among congeneric species. Here, we investigated the embryonic growth curve across seven recently-described species of the highly diverse genus Phalloceros (Eigenmann, 1907). We also investigated possible intraspecific differences and whether other female characteristics affected embryo mass. We found that embryo mass decreased until around stage 20 and then increased, resulting in a 1.5 to 3-fold mass gain from fertilization to birth. Embryo mass changed non-linearly with stage of development and was affected by species identity (or locality) and female somatic dry mass. This initial loss then gain of embryonic mass during development is unique among other Poeciliidae species and was conserved across populations and species, even though size at birth can vary. Other species instead either lose mass if they lack placentas or gain mass exponentially if they have placentas. The Phalloceros mode of maternal provisioning could thus represent a different form from that seen in other species of Poeciliidae.more » « less
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Teeling, Emma (Ed.)Abstract The evolutionary origin of complex organs challenges empirical study because most organs evolved hundreds of millions of years ago. The placenta of live-bearing fish in the family Poeciliidae represents a unique opportunity to study the evolutionary origin of complex organs, because in this family a placenta evolved at least nine times independently. It is currently unknown whether this repeated evolution is accompanied by similar, repeated, genomic changes in placental species. Here, we compare whole genomes of 26 poeciliid species representing six out of nine independent origins of placentation. Evolutionary rate analysis revealed that the evolution of the placenta coincides with convergent shifts in the evolutionary rate of 78 protein-coding genes, mainly observed in transporter- and vesicle-located genes. Furthermore, differences in sequence conservation showed that placental evolution coincided with similar changes in 76 noncoding regulatory elements, occurring primarily around genes that regulate development. The unexpected high occurrence of GATA simple repeats in the regulatory elements suggests an important function for GATA repeats in developmental gene regulation. The distinction in molecular evolution observed, with protein-coding parallel changes more often found in metabolic and structural pathways, compared with regulatory change more frequently found in developmental pathways, offers a compelling model for complex trait evolution in general: changing the regulation of otherwise highly conserved developmental genes may allow for the evolution of complex traits.more » « less
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Abstract A major question in ecology is how often competing species evolve to reduce competitive interactions and facilitate coexistence. One untested route for a reduction in competitive interactions is through ontogenetic changes in the trophic niche of one or more of the interacting species. In such cases, theory predicts that two species can coexist if the weaker competitor changes its resource niche to a greater degree with increased body size than the superior competitor.
We tested this prediction using stable isotopes that yield information about the trophic position (δ15N) and carbon source (δ13C) of two coexisting fish species: Trinidadian guppies
Poecilia reticulata and killifishRivulus hartii .We examined fish from locations representing three natural community types: (1) where killifish and guppies live with predators, (2) where killifish and guppies live without predators and (3) where killifish are the only fish species. We also examined killifish from communities in which we had introduced guppies, providing a temporal sequence of the community changes following the transition from a killifish only to a killifish–guppy community.
We found that killifish, which are the weaker competitor, had a much larger ontogenetic niche shift in trophic position than guppies in the community where competition is most intense (killifish–guppy only). This result is consistent with theory for size‐structured populations, which predicts that these results should lead to stable coexistence of the two species. Comparisons with other communities containing guppies, killifish and predators and ones where killifish live by themselves revealed that these results are caused primarily by a loss of ontogenetic niche changes in guppies, even though they are the stronger competitor. Comparisons of these natural communities with communities in which guppies were translocated into sites containing only killifish showed that the experimental communities were intermediate between the natural killifish–guppy community and the killifish–guppy–predator community, suggesting contemporary evolution in these ontogenetic trophic differences.
These results provide comparative evidence for ontogenetic niche shifts in contributing to species coexistence and comparative and experimental evidence for evolutionary or plastic changes in ontogenetic niche shifts following the formation of new communities.
