Rapid changes in genetic architecture of behavioural syndromes following colonization of a novel environment.

Behavioural syndromes, that is correlated behaviours, may be a result from adaptive correlational selection, but in a new environmental setting, the trait correlation might act as an evolutionary constraint. However, knowledge about the quantitative genetic basis of behavioural syndromes, and the stability and evolvability of genetic correlations under different ecological conditions, is limited. We investigated the quantitative genetic basis of correlated behaviours in the freshwater isopod Asellus aquaticus. In some Swedish lakes, A. aquaticus has recently colonized a novel habitat and diverged into two ecotypes, presumably due to habitat-specific selection from predation. Using a common garden approach and animal model analyses, we estimated quantitative genetic parameters for behavioural traits and compared the genetic architecture between the ecotypes. We report that the genetic covariance structure of the behavioural traits has been altered in the novel ecotype, demonstrating divergence in behavioural correlations. Thus, our study confirms that genetic correlations behind behaviours can change rapidly in response to novel selective environments.

Hybridization and speciation.

Hybridization has many and varied impacts on the process of speciation. Hybridization may slow or reverse differentiation by allowing gene flow and recombination. It may accelerate speciation via adaptive introgression or cause near-instantaneous speciation by allopolyploidization. It may have multiple effects at different stages and in different spatial contexts within a single speciation event. We offer a perspective on the context and evolutionary significance of hybridization during speciation, highlighting issues of current interest and debate. In secondary contact zones, it is uncertain if barriers to gene flow will be strengthened or broken down due to recombination and gene flow. Theory and empirical evidence suggest the latter is more likely, except within and around strongly selected genomic regions. Hybridization may contribute to speciation through the formation of new hybrid taxa, whereas introgression of a few loci may promote adaptive divergence and so facilitate speciation. Gene regulatory networks, epigenetic effects and the evolution of selfish genetic material in the genome suggest that the Dobzhansky-Muller model of hybrid incompatibilities requires a broader interpretation. Finally, although the incidence of reinforcement remains uncertain, this and other interactions in areas of sympatry may have knock-on effects on speciation both within and outside regions of hybridization.

Local adaptation within a hybrid species.

Ecological divergence among populations may be strongly influenced by their genetic background. For instance, genetic admixture through introgressive hybridization or hybrid speciation is likely to affect the genetic variation and evolvability of phenotypic traits. We studied geographic variation in two beak dimensions and three other phenotypic traits of the Italian sparrow (Passer italiae), a young hybrid species formed through interbreeding between house sparrows (P. domesticus) and Spanish sparrows (P. hispaniolensis). We found that beak morphology was strongly influenced by precipitation regimes and that it appeared to be the target of divergent selection within Italian sparrows. Interestingly, however, the degree of parental genetic contribution in the hybrid species had no effect on phenotypic beak variation. Moreover, beak height divergence may mediate genetic differentiation between populations, consistent with isolation-by-adaptation within this hybrid species. The study illustrates how hybrid species may be relatively unconstrained by their admixed genetic background, allowing them to adapt rapidly to environmental variation.

Evolution and stability of the G-matrix during the colonization of a novel environment.

Populations that undergo a process of rapid evolution present excellent opportunities to investigate the mechanisms driving or restraining adaptive divergence. The genetic variance-covariance matrix (G) is often considered to constrain adaptation but little is known about its potential to evolve during phenotypic divergence. We compared the G-matrices of ancestral and recently established ecotype populations of an aquatic isopod (Asellus aquaticus) that have diverged in parallel in two south Swedish lakes. Phenotypic changes after colonization involved a reduction in overall size, lost pigmentation and changes in shape. Comparisons between G-matrices reveal close similarity within the same ecotype from different lakes but some degree of differentiation among ecotypes. Phenotypic divergence has apparently not been much influenced by the orientation of G. Additive genetic variation in the newly colonized habitats has also decreased substantially. This suggests that a process of adaptation from standing genetic variation has occurred and has probably facilitated phenotypic divergence.

Changes in behavioural trait integration following rapid ecotype divergence in an aquatic isopod.

Colonization of new habitats can relax selection pressures, and traits or trait combinations no longer selected for might become reduced or lost. We investigated behavioural differentiation and behavioural trait integration in the freshwater isopod Asellus aquaticus. This isopod has recently colonized a novel habitat and diverged into two ecotypes which encounter different predator faunas. We investigated sex-specific behavioural differences and phenotypic integration in three behavioural assays: (i) time to emerge (TE) from a shelter, (ii) activity and (iii) escape behaviour. General activity and escape behaviour differed between ecotypes. Furthermore, general activity and TE differed between sexes. Behavioural traits were more frequently correlated in the ancestral habitat, and phenotypic integration tended to be higher in this habitat as well. Our study suggests that different predator types, but also other ecological factors such as habitat matrices and population densities, might explain the differences in behavioural integration in these ecotypes.

Parallel divergence in mate guarding behaviour following colonization of a novel habitat.

Ecological factors can have profound effects on mating system and mating behaviour. We investigated the effect of altered ecological conditions, following colonization of a novel habitat, on precopulatory mate guarding in a freshwater isopod (Asellus aquaticus). This isopod occurs in two different ecotypes, which coexist within several different lakes in Sweden but which utilize different habitats. These ecotypes have rapidly (ca. 40 generations) diverged in parallel among lakes in several phenotypic characters, presumably as a response to different predatory pressures. Here, we demonstrate that also mate guarding characteristics have diverged in parallel between the ecotypes in different lakes. This is one of the few studies reporting parallel evolution of mating behaviour. Furthermore, our results also indicate a potential sexual conflict, as the length of mate guarding appears to lower components of female fitness. We discuss how novel environments might have strong and rapid effects on mate guarding dynamics and mating behaviour.

Parallelism and historical contingency during rapid ecotype divergence in an isopod.

Recent studies on parallel evolution have focused on the relative role of selection and historical contingency during adaptive divergence. Here, we study geographically separate and genetically independent lake populations of a freshwater isopod (Asellus aquaticus) in southern Sweden. In two of these lakes, a novel habitat was rapidly colonized by isopods from a source habitat. Rapid phenotypic changes in pigmentation, size and sexual behaviour have occurred, presumably in response to different predatory regimes. We partitioned the phenotypic variation arising from habitat ('selection': 81­94%), lake ('history': 0.1­6%) and lake × habitat interaction ('unique diversification': 0.4­13%) for several traits. There was a limited role for historical contingency but a strong signature of selection. We also found higher phenotypic variation in the source populations. Phenotype sorting during colonization and strong divergent selection might have contributed to these rapid changes. Consequently, phenotypic divergence was only weakly influenced by historical contingency.

Phenotypic integration and conserved covariance structure in calopterygid damselflies.

By comparing the phenotypic (P) variance-covariance matrices between closely related taxa or conspecific populations, one can study the outcome of the interplay between selection and developmental constraints in phenotypic evolution. Shared patterns of phenotypic integration are also of interest and might result from similarities in either selection or developmental pathways. We compared P-matrices and phenotypic integration indices between populations and species of the damselfly genus Calopteryx. P(max)-comparisons between parapatric C. splendens populations revealed stronger conserved phenotypic covariance structure than P(max)-comparisons between species, suggesting that divergence in its early stages proceeds along phenotypic lines of least resistance. Within- and among-population correlations in C. splendens were highly concordant, in further support of initial divergence along P(max). Despite some similarities in overall phenotypic integration between C. splendens and C. virgo, these two species only had several P-matrix eigenvectors in common, indicating that after reproductive isolation, divergence has proceeded against P(max).