Contemporary Parallel Diversification, Antipredator Adaptations and Phenotypic Integration in an Aquatic Isopod

It is increasingly being recognized that predation can be a strong diversifying agent promoting ecological divergence. Adaptations against different predatory regimes can emerge over short periods of time and include many different traits. We studied antipredator adaptations in two ecotypes of an isopod (Asellus aquaticus) that have, diverged in parallel in two Swedish lakes over the last two decades. We quantified differences in escape speed, morphology and behavior for isopods from different ecotypes present in these lakes. Isopods from the source habitat (reed) coexist with mainly invertebrate predators. They are more stream-profiled and have higher escape speeds than isopods in the newly colonized stonewort habitat, which has higher density of fish predators. Stonewort isopods also show more cautious behaviors and had higher levels of phenotypic integration between coloration and morphological traits than the reed isopods. Colonization of a novel habitat with a different predation regime has thus strengthened the correlations between pigmentation and morphology and weakened escape performance. The strong signature of parallelism for these phenotypic traits indicates that divergence is likely to be adaptive and is likely to have been driven by differences in predatory regimes. Furthermore, our results indicate that physical performance, behavior and morphology can change rapidly and in concert as new habitats are colonized

Resolution of conflict between parental genomes in a hybrid species

AbstractThe development of reproductive barriers against parent species is crucial during hybrid speciation, and post-zygotic isolation can be important in this process. Genetic incompatibilities that normally isolate the parent species can become sorted in hybrids to form reproductive barriers towards either parent. However, the extent to which this sorting process is systematically biased and therefore predictable in which loci are involved and which alleles are favored is largely unknown. Theoretically, reduced fitness in hybrids due to the mixing of differentiated genomes can be resolved through rapid evolution towards allelic combinations ancestral to lineage-splitting of the parent species, as these alleles have successfully coexisted in the past. However, for each locus, this effect may be influenced by its chromosomal location, function, and interactions with other loci. We use the Italian sparrow, a homoploid hybrid species that has developed post-zygotic barriers against its parent species, to investigate this prediction. We show significant bias towards fixation of the ancestral allele among 57 nuclear intragenic SNPs, particularly those with a mitochondrial function whose ancestral allele came from the same parent species as the mitochondria. Consistent with increased pleiotropy leading to stronger fitness effects, genes with more protein-protein interactions were more biased in favor of the ancestral allele. Furthermore, the number of protein-protein interactions was especially low among candidate incompatibilities still segregating within Italian sparrows, suggesting that low pleiotropy allows steep intraspecific clines in allele frequencies to form. Finally, we report evidence for pervasive epistatic interactions within one Italian sparrow population, particularly involving loci isolating the two parent species but not hybrid and parent. However there was a lack of classic incompatibilities and no admixture linkage disequilibrium. This suggests that parental genome admixture can continue to constrain evolution and prevent genome stabilization long after incompatibilities have been purged.</jats:p

Phenotypic Plasticity in Response to the Social Environment: Effects of Density and Sex Ratio on Mating Behaviour Following Ecotype Divergence

The ability to express phenotypically plastic responses to environmental cues might be adaptive in changing environments. We studied phenotypic plasticity in mating behaviour as a response to population density and adult sex ratio in a freshwater isopod (Asellus aquaticus). A. aquaticus has recently diverged into two distinct ecotypes, inhabiting different lake habitats (reed Phragmites australis and stonewort Chara tomentosa, respectively). In field surveys, we found that these habitats differ markedly in isopod population densities and adult sex ratios. These spatially and temporally demographic differences are likely to affect mating behaviour. We performed behavioural experiments using animals from both the ancestral ecotype (“reed” isopods) and from the novel ecotype (“stonewort” isopods) population. We found that neither ecotype adjusted their behaviour in response to population density. However, the reed ecotype had a higher intrinsic mating propensity across densities. In contrast to the effects of density, we found ecotype differences in plasticity in response to sex ratio. The stonewort ecotype show pronounced phenotypic plasticity in mating propensity to adult sex ratio, whereas the reed ecotype showed a more canalised behaviour with respect to this demographic factor. We suggest that the lower overall mating propensity and the phenotypic plasticity in response to sex ratio have evolved in the novel stonewort ecotype following invasion of the novel habitat. Plasticity in mating behaviour may in turn have effects on the direction and intensity of sexual selection in the stonewort habitat, which may fuel further ecotype divergence

Just How Much is the G-matrix Actually Constraining Adaptation?

The genetic variance-covariance matrix (G) has long been considered to summarize the genetic constraints biasing evolution in its early stages, although in some instances, G can enhance divergence and facilitate adaptation. However, the effects of G on the response to selection might be of less importance than previously thought. In addition, it has been suggested that selection itself, under certain conditions, might rapidly alter the genetic covariance structure. If selection can indeed affect the stability of G to facilitate evolution, the overall structure of G might not be as important to consider as the past selective conditions that G was subject to. Thus, more empirical work is needed on the stability of G in the early stages of divergence before one can really assess to what extent G constrains evolution

The Interplay Between Selection And Constraints On Adaptive Divergence And Phenotypic Evolution

Under natural or sexual selection, individuals with advantageous traits or combinations of traits will be more successful than their peers at surviving and/or reproducing. Provided these traits are heritable, meaning that they have a genetic basis, the traits combinations which are selected for, will increase in frequency in the population. When selection is intense and persistent, adaptive traits may become ubiquitous in the population, and we may then say that this population has evolved and become adapted. However, this process might not always occur rapidly. This is because adaptive evolution occurs only when the individuals of a population are diverse in their trait combinations and when there is a significant amount of genetic variation for the trait(s) upon which selection is acting. However, if the adaptive optimum of a combination of traits is situated in a direction where there is little variation available, adaptation will be slowed down, and we will say it is constrained. This is precisely what I attempted to study in this thesis. The outcome of the interplay between selection and constraints might lead to evolution, to divergence between populations, and finally to the emergence of new species and biodiversity. By using different statistical techniques used in quantitative genetics or geometric morphometrics, combined with behavioral and breeding experiments, I tried to draw some conclusions on the role of constraints both in the early stages of adaptation and divergence (using isopod lake populations as a model system) and in the latter stages of divergence and speciation (using damselfly species as model organisms). My main conclusions are that in the context of strong divergent selection, constraints may be overcome and adaptation may proceed, provided that gene flow between populations is restrained

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

Effects of natural and sexual selection on adaptive population divergence and premating isolation in a damselfly

The relative strength of different types of directional selection has seldom been compared directly in natural populations. A recent meta-analysis of phenotypic selection studies in natural populations suggested that directional sexual selection may be stronger in magnitude than directional natural selection, although this pattern may have partly been confounded by the different time scales over which selection was estimated. Knowledge about the strength of different types of selection is of general interest for understanding how selective forces affect adaptive population divergence and how they may influence speciation. We studied divergent selection on morphology in parapatric, natural damselfly (Calopteryx splendens) populations. Sexual selection was stronger than natural selection measured on the same traits, irrespective of the time scale over which sexual selection was measured. Visualization of the fitness surfaces indicated that population divergence in overall morphology is more strongly influenced by divergent sexual selection rather than natural selection. Courtship success of experimental immigrant males was lower than that of resident males, indicating incipient sexual isolation between these populations. We conclude that current and strong sexual selection promotes adaptive population divergence in this species and that premating sexual isolation may have arisen as a correlated response to divergent sexual selection. Our results highlight the importance of sexual selection, rather than natural selection in the adaptive radiation of odonates, and supports previous suggestions that divergent sexual selection promotes speciation in this group

The role of different reproductive barriers during phenotypic divergence of isopod ecotypes.

The question of how diverging populations become separate species by restraining gene flow is a central issue in evolutionary biology. Assortative mating might emerge early during adaptive divergence, but the role of other types of reproductive barriers such as migration modification have recently received increased attention. We demonstrate that two recently diverged ecotypes of a freshwater isopod (Asellus aquaticus) have rapidly developed premating isolation, and this isolation barrier has emerged independently and in parallel in two south Swedish lakes. This is consistent with ecological speciation theory, which predicts that reproductive isolation arises as a byproduct of ecological divergence. We also find that in one of these lakes, habitat choice acts as the main barrier to gene flow. These observations and experimental results suggest that migration modification might be as important as assortative mating in the early stages of ecological speciation. Simulations suggest that the joint action of these two isolating barriers is likely to greatly facilitate adaptive divergence, compared to if each barrier was acting alone