Polymorphism may play an important role in speciation because new species could originate from the distinctive morphs observed in polymorphic populations. However, much remains to be understood about the process by which morphs found new species. To detail the steps of this mode of speciation, we studied the geographic variation and evolutionary history of a throat color polymorphism that distinguishes the "rock-paper-scissors" mating strategies of the sideblotched lizard, Uta stansburiana. We found that the polymorphism is geographically widespread and has been maintained for millions of years. However, there are many populations with reduced numbers of throat color morphs. Phylogenetic reconstruction showed that the polymorphism is ancestral, but it has been independently lost eight times, often giving rise to morphologically distinct subspecies/species. Changes to the polymorphism likely involved selection because the allele for one particular male strategy, the "sneaker" morph, has been lost in all cases. Polymorphism loss was associated with accelerated evolution of male size, female size, and sexual dimorphism, which suggests that polymorphism loss can promote rapid divergence among populations and aid species formation.lizard | morph | phylogeny | rock-paper-scissors | Uta stansburiana P olymorphic forms within a population could be the starting material for new species (1-4). Tests of how polymorphisms diversify into new lineages have been relatively rare (2, 5, 6), despite increasing recognition that polymorphisms, such as alternative mating strategies, are common within species (7-9). Here we investigate the processes by which a morph in a polymorphic population may diverge to found a new species.Competition within a population can generate morphs possessing alternative adaptations, which may become as phenotypically distinct as separate species and thus primed for speciation (2, 3). WestEberhard theorized how morphs may promote speciation (2, 10) and suggested that selection in new environments may favor a particular morph, destabilizing the dynamics maintaining the polymorphism. If a population loses a morph, this can lead to rapid phenotypic divergence in the remaining morph(s) (2). One cause of rapid divergence is that genetic evolution may be constrained in polymorphic populations, because alleles that increase the fitness of a single morph but decrease the fitness of other morphs may be unable to spread. Loss of a morph allows these alleles to spread, which can result in rapid phenotypic evolution in the direction of specialization on the remaining morph phenotypes, a process called character release (2). Another reason for rapid evolution coincident with morph loss is that colonization of a new ecological environment may select against a particular morph and also favor novel phenotypes in the remaining morphs. In addition, loss of a morph changes the competitive environment, because the fitness of morphs can depend on the frequency of other types in the population (8). Rapid evolution may occur ...

Population-level comparative analyses can link microevolutionary processes within populations to macroevolutionary patterns of diversification. We used the comparative method to study the evolution of sexual size dimorphism (SSD) among populations of side-blotched lizards (Uta stansburiana). Uta stansburiana is polymorphic for different male mating and female life-history strategies in some populations, but monomorphic in others. We tested whether intrasexual selection among males, fecundity selection on females, and the presence of polymorphic strategies affected levels of SSD. We first resolved a phylogeny for 41 populations across the range of the species and documented a substantial regional structure. Our intraspecific data had significant phylogenetic signal, and correcting for phylogeny using independent contrasts had large effects on our results. Polymorphic populations had male-biased SSD and changes in male body size, levels of tail breaks, and SSD consistent with the intrasexual selection hypothesis. Monomorphic populations had changes in female size, clutch size, and SSD consistent with the fecundity selection hypothesis. Fecundity selection is a likely cause of some monomorphic populations having no SSD or female-biased SSD.Our results suggest that changes in mating strategies are associated with phenotypic diversification and multiple evolutionary forces can shape SSD. Sexual selection arises from competition for mates and can lead to phenotypic diversification within and among species (Darwin 1871;West-Eberhard 1983;Andersson 1994). Most studies of sexual selection have focused either on selection within populations (Sinervo and Lively 1996; Krakauer 2005; Clutton-Brock et al.

Aim The salamander Ensatina eschscholtzii Gray is a classic example of a ring species, or a species that has expanded around a central barrier to form a secondary contact characterized by species-level divergence. In the original formulation of the ring species scenario, an explicit biogeographical model was proposed to account for the occurrence of intraspecific sympatry between two subspecies in southern California (the 'southern closure' model). Here we develop an alternative ring species model that is informed by the geomorphological development of the California Coast Ranges, and which situates the point of ring closure in the Monterey Bay region of central coastal California (the 'Monterey closure' model). Our study has two aims. The first is to use phylogenetic methods to evaluate the two competing biogeographical models. The second is to describe patterns of phylogeographical diversity throughout the range of the Ensatina complex, and to compare these patterns with previously published molecular systematic data.Location Western North America, with a focus on the state of California, USA.Methods We obtained mitochondrial DNA sequence data from 385 individuals from 224 populations. A phylogeny was inferred using Bayesian techniques, and the geographical distributions of haplotypes and clades were mapped. The two biogeographical ring species models were tested against our Bayesian topology, including the associated Bayesian 95% credible set of trees.Results High levels of phylogeographical diversity were revealed, especially in central coastal and northern California. Our Bayesian topology contradicts the Monterey closure model; however, 0.08% of the trees in our Bayesian 95% credible set are consistent with this model. In contrast, the classic ring species biogeographical model (the southern closure model) is consistent with our Bayesian topology, as were 99.92% of the trees in our 95% credible set.Main conclusions Our Bayesian phylogenetic analysis most strongly supports the classic ring species model, modified to accommodate an improved understanding of the complex geomorphological evolution of the California Coast Ranges. In addition, high levels of phylogeographical diversity in central and northern California were identified, which is consistent with the striking levels of allozymic differentiation reported previously from those regions.

Allozymes and mitochondrial DNA sequences were used to examine the phylogeographical history of the rough-skinned newt, Taricha granulosa, in western North America. Nineteen populations were analysed for allozyme variation at 45 loci, and 23 populations were analysed for cytochrome b sequence variation. Both data sets agree that populations in the southern part of the range are characterized by isolation by distance, whereas northern populations fit the expectations of a recent range expansion. However, the northern limit of isolation by distance (and the southern limit of range expansion) is located in Oregon State by the mtDNA data, and in Washington State by the allozyme data. Nevertheless, both data sets are consistent with the known Pleistocene history of western North America, with phylogenetically basal populations in central and northern California, and a recent range expansion in the north following the retreat of the Cordilleran ice sheet 10 000 years ago. Additionally, a population in Idaho, previously considered introduced from central California based on morphometric analyses, possesses a distinct mtDNA haplotype, suggesting it could be native. The relevance of these results for Pacific Northwest biogeography is discussed.