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Focuses on the issues of terminology, gradient dynamics and the ease of sympatric speciation in adaptive dynamics. Importance of terminology in adaptive dynamics; Justification of the claims often made within the Adaptive Dynamics; Criticism of the initial conditions used in the allele frequencies.

Adaptive Dynamics is an approach to studying evolutionary change when fitness is density or frequency dependent. Modern papers identifying themselves as using this approach first appeared in the 1990s, and have greatly increased up to the present. However, because of the rather technical nature of many of the papers, the approach is not widely known or understood by evolutionary biologists. In this review we aim to remedy this situation by outlining the methodology and then examining its strengths and weaknesses. We carry this out by posing and answering 20 key questions on Adaptive Dynamics. We conclude that Adaptive Dynamics provides a set of useful approximations for studying various evolutionary questions. However, as with any approximate method, conclusions based on Adaptive Dynamics are valid only under some restrictions that we discuss. [ABSTRACT FROM AUTHOR]

Testing whether some species traits have a significant effect on diversification rates is central in the assessment of macroevolutionary theories. However, we still lack a powerful method to tackle this objective. I present a new method for the statistical analysis of diversification with species traits. The required data are observations of the traits on recent species, the phylogenetic tree of these species, and reconstructions of ancestral values of the traits. Several traits, either continuous or discrete, and in some cases their interactions, can be analyzed simultaneously. The parameters are estimated by the method of maximum likelihood. The statistical significance of the effects in a model can be tested with likelihood ratio tests. A simulation study showed that past random extinction events do not affect the Type I error rate of the tests, whereas statistical power is decreased, though some power is still kept if the effect of the simulated trait on speciation is strong. The use of the method is illustrated by the analysis of published data on primates. The analysis of these data showed that the apparent overall positive relationship between body mass and species diversity is actually an artifact due to a clade-specific effect. Within each clade the effect of body mass on speciation rate was in fact negative. The present method allows to take both effects (clade and body mass) into account simultaneously. [ABSTRACT FROM AUTHOR]

Sexual conflict, where male and female reproductive interests differ, is probably widespread and often mediated by male or sperm proteins and female or egg proteins that bind to each other during mating or fertilization. One potential consequence is maintenance of polymorphism in these proteins, which might result in reproductive isolation between sympatric subpopulations. I investigate the conditions for polymorphism maintenance in a series of mathematical models of sexual conflict over mating or fertilization frequency. The models represent a male or sperm ligand and a female or egg receptor, and they differ in whether expression of either protein is haploid or diploid. For diploid expression, the conditions imply that patterns of dominance, which involve neither overdominance nor underdominance, can determine whether polymorphism is maintained. For example, suppose ligand expression is diploid, and consider ligand alleles L1 and L2 in interactions with a given receptor genotype; if L1/L1 males are fitter than L2/L2 males in these interactions, then polymorphism is more likely to be maintained when L1/L2 males more closely resemble L1/L1 males in these interactions. Such fitter-allele dominance might be typical of a ligand or its receptor due to their biochemistry, in which case polymorphism might be typical of the pair. [ABSTRACT FROM AUTHOR]

Although reinforcement is ostensibly driven by selection against hybrids, there are often other components in empirical cases and theoretical models of reinforcement that may contribute to premating isolation. One of these components is local adaptation of a trait used in mate choice. I use several different comparisons to assess the roles that local adaptation and selection against hybrids may play in reinforcement models. Both numerical simulations of exact recursion equations and analytical weak selection approximations are employed. I find that selection against hybrids may play a small role in driving preference evolution in a reinforcement model where the mating cue is separate from loci causing hybrid incompatibilities. When females have preferences directly for purebreds of their own population, however, selection against hybrids can play a large role in premating isolation evolution. I present some situations in which this type of selection is likely to exist. This work also illustrates shortfalls of using a weak selection approach to address questions about reinforcement. [ABSTRACT FROM AUTHOR]

While it now appears likely that sympatric speciation is possible, its generality remains contentious. If it really is rare, then most natural populations must not fit the assumptions of sympatric speciation theory. A better understanding of these assumptions may help identify when sympatric speciation is or is not likely. This paper investigates two such assumptions: that genetic variation for stringent assortative mating is not limiting and that females are not penalized for mating assortatively. Simulations demonstrate that the speed of sympatric speciation is very sensitive to the population's capacity for stringent assortative mating and is potentially extremely slow. The rapid divergence often thought to be a hallmark of sympatric speciation may only occur in a restricted area of parameter space. [ABSTRACT FROM AUTHOR]

The tradition of classifying cases of speciation into discrete geographic categories (allopatric, parapatric and sympatric) fuelled decades of fruitful research and debate. Not surprisingly, as the science has become more sophisticated, this simplistic taxonomy has become increasingly obsolete. Geographic patterns are now reasonably well understood. Sister species are rarely sympatric, implying that sympatric speciation, it its most general sense, is rare. However, sympatric speciation, even in its most restricted population genetic sense, is possible. Several case studies have demonstrated that divergence has occurred in nature without geographic barriers to gene flow. Obviously, different sets of criteria for sympatric speciation will lead to different numbers of qualifying cases. But changing the rules of nomenclature to make ‘sympatric speciation’ more or less common does not constitute scientific progress. Advances in the study of speciation have come from studies of the processes that constrain or promote divergence, and how they are affected by geography. [ABSTRACT FROM AUTHOR]

Sympatric speciation has always fascinated evolutionary biologists, and for good reason; it pits diversifying selection directly against the tendency of sexual reproduction to homogenize populations. However, different investigators have used different definitions of sympatric speciation and different criteria for diagnosing cases of sympatric speciation. Here, we explore some of the definitions that have been used in empirical and theoretical studies. Definitions based on biogeography do not always produce the same conclusions as definitions based on population genetics. The most precise definitions make sympatric speciation an infinitesimal end point of a continuum. Because it is virtually impossible to demonstrate the occurrence of such a theoretical extreme, we argue that testing whether a case fits a particular definition is less informative than evaluating the biological processes affecting divergence. We do not deny the importance of geographical context for understanding divergence. Rather, we believe this context can be better understood by modelling and measuring quantities, such as gene flow and selection, rather than assigning cases to discrete categories like sympatric and allopatric speciation. [ABSTRACT FROM AUTHOR]

Maternally inherited endosymbionts have been implicated as significant drivers of sexual conflict within their hosts, typically through sex-ratio manipulation. Empirical studies show that some of these endosymbionts have the potential to influence sexual conflict not by sex-ratio distortion, but by altering reproductive traits within their hosts. Research has already shown that reproductive traits involved in mating/fertilization process are integral ‘players’ in sexual conflict, thus suggesting the novel hypothesis that endosymbiont-induced changes in reproductive phenotypes can impact the dynamics of sexual conflict. Here, we use a standard quantitative genetic approach to model the effects of endosymbiont-induced changes in a female reproductive trait on the dynamics of sexual conflict over mating/fertilization rate. Our model shows that an endosymbiont-induced alteration of a host female reproductive trait that affects mating rate can maintain the endosymbiont infection within the host population, and does so in the absence of sex-ratio distortion and cytoplasmic incompatibility. [ABSTRACT FROM AUTHOR]

We present here a stochastic two-locus, two-habitat model for the evolution of migration with local adaptation and kin selection. One locus determines the migration rate while the other causes local adaptation. We show that the opposing forces of kin competition and local adaptation can lead to the existence of one or two convergence stable migration rates, notably depending on the recombination rate between the two loci. We show that linkage between migration and local adaptation loci has two antagonist effects: when linkage is tight, cost of local adaptation increases, leading to smaller equilibrium migration rates. However, when linkage is tighter, the population structure at the migration locus tends to be very high because of the indirect selection, and thus equilibrium migration rates increases. This result, qualitatively different from results obtained with other models of migration evolution, indicates that ignoring drift or the detail of the genetic architecture may lead to incorrect conclusions. [ABSTRACT FROM AUTHOR]

We study the population genetics of adaptation in nonequilibrium haploid asexual populations. We find that the accumulation of deleterious mutations, due to the operation of Muller's ratchet, can considerably reduce the rate of fixation of advantageous alleles. Such reduction can be approximated reasonably well by a reduction in the effective population size. In the absence of Muller's ratchet, a beneficial mutation can only become fixed if it creates the best possible genotype; if Muller's ratchet operates, however, mutations initially arising in a nonoptimal genotype can also become fixed in the population, since the loss of the least-loaded class implies that an initially nonoptimal background can become optimal. We show that, while the rate at which adaptive mutations become fixed is reduced, the rate of fixation of deleterious mutations due to the ratchet is not changed by the presence of beneficial mutations as long as the rate of their occurrence is low and the deleterious effects of mutations (sd) are higher than the beneficial effects (sa). When sa > sd, the advantage of a beneficial mutation can outweigh the deleterious effects of associated mutations. Under these conditions, a beneficial allele can drag to fixation deleterious mutations initially associated with it at a higher rate than in the absence of advantageous alleles. We propose analytical approximations for the rates of accumulation of deleterious and beneficial mutations. Furthermore, when allowing for the possible occurrence of interference between beneficial alleles, we find that the presence of deleterious mutations of either very weak or very strong effect can marginally increase the rate of accumulation of beneficial mutations over that observed in the absence of such deleterious mutations. [ABSTRACT FROM AUTHOR]

Twenty years ago, Bulmer and Bull suggested that disruptive selection, produced by environmental fluctuations, can result in an evolutionary transition from environmental sex determination (ESD) to genetic sex determination (GSD). We investigated the feasibility of such a process, using mutation-limited adaptive dynamics and individual-based computer simulations. Our model describes the evolution of a reaction norm for sex determination in a metapopulation setting with partial migration and variation in an environmental variable both within and between local patches. The reaction norm represents the probability of becoming a female as a function of environmental state and was modeled as a sigmoid function with two parameters, one giving the location (i.e., the value of the environmental variable for which an individual has equal chance of becoming either sex) and the other giving the slope of the reaction norm for that environment. The slope can be interpreted as being set by the level of developmental noise in morph determination, with less noise giving a steeper slope and a more switchlike reaction norm. We found convergence stable reaction norms with intermediate to large amounts of developmental noise for conditions characterized by low migration rates, small differential competitive advantages between the sexes over environments, and little variation between individual environments within patches compared to variation between patches. We also considered reaction norms with the slope parameter constrained to a high value, corresponding to little developmental noise. For these we found evolutionary branching in the location parameter and a transition from ESD toward GSD, analogous to the original analysis by Bulmer and Bull. Further evolutionary change, including dominance evolution, produced a polymorphism acting as a GSD system with heterogamety. Our results point to the role of developmental noise in the evolution of sex determination. [ABSTRACT FROM AUTHOR]

Formation of partially reproductively isolated ecotypes in the rough periwinkle, Littorina saxatilis, may be a case of incipient nonallopatric ecological speciation. To better understand the dynamics of ecotype formation, its timescale, driving forces and evolutionary consequences, we developed a spatially explicit, individual-based model incorporating relevant ecological, spatial and mate selection data for Swedish L. saxatilis. We explore the impact of bounded hybrid superiority, ecological scenarios and mate selection systems on ecotype formation, gene flow and the evolution of prezygotic isolation. Our model shows that ecotypes are expected to form rapidly in parapatry under conditions applicable to Swedish L. saxatilis and may proceed to speciation. However, evolution of nonrandom mating had complex behaviour. Ecotype evolution was inhibited by pre-existing mating preferences, but facilitated by the evolution of novel preferences. While in many scenarios positive assortative mating reduced gene flow between ecotypes, in others negative assortative mating arose, preferences were lost after ecotype formation, preferences were confined to one ecotype or the ancestral ecotype became extinct through sexual selection. Bounded hybrid superiority (as observed in nature) enhanced ecotype formation but increased gene flow. Our results highlight that ecotype formation and speciation are distinct processes: factors that contribute to ecotype formation can be detrimental to speciation and vice versa. The complex interactions observed between local adaptation and nonrandom mating imply that generalization from data is unreliable without quantitative theory for speciation. [ABSTRACT FROM AUTHOR]

Within the Diptera, two different selfish genetic elements are known to cause the production of female-biased sex ratios: maternally inherited bacteria that kill male zygotes (male-killers), and X chromosomes causing the degeneration of Y-bearing sperm in males (meiotic drive). We here develop a mathematical model for the dynamics of these two sex-ratio distorters where they co-occur. We show that X chromosome meiotic drive elements can be expected to substantially lower the equilibrium frequency of male-killers and can even lead to their extinction. Conversely, male-killers can also decrease the equilibrium frequency of X drivers and cause their extinction. Thus, we predict that there will be some complementarity in the incidence of X chromosome meiotic drive and male-killing in natural populations, with a lower than expected number of species bearing both elements. [ABSTRACT FROM AUTHOR]

Learning processes potentially play a role in speciation but are often ignored in speciation models. Learning may, for instance, play a role when a new niche is being colonized, because the learning of niche features may cause niche-specific assortative mating and a tendency to produce young in this niche. Several animal species learn about their environmental features that may be important in finding or attracting mates. We use a gene-culture coevolutionary model to look into the effect of such learning on the colonization of new niches and on the genetic divergence between groups using different niches, which are steps necessary in achieving speciation. We assume that density is regulated separately in each of the two niches and that the viability of an individual depends on its genotype as well as on which niche it exploits. Our results show that genetic adaptation to the new niche is enhanced by a high female fecundity and a low viability selection against heterozygotes. Furthermore, when initial colonization (without genetic adaptation) fails, genetic divergence is more difficult when the mating preference is stronger. In contrast, when colonization without genetic adaptation is successful, a stronger mating preference makes genetic divergence easier. An increase in the number of egg-laying mistakes by females can have a positive or negative effect on the success of genetic adaptation depending on other parameters. We show that genetic divergence can be prevented by a niche shift, which can occur only if viabilities in the two niches are asymmetrical. [ABSTRACT FROM AUTHOR]