In recent years, strikingly consistent patterns of biodiversity have been identified over space, time, organism type and geographical region. A neutral theory (assuming no environmental selection or organismal interactions) has been shown to predict many patterns of ecological biodiversity. This theory is based on a mechanism by which new species arise similarly to point mutations in a population without sexual reproduction. Here we report the simulation of populations with sexual reproduction, mutation and dispersal. We found simulated time dependence of speciation rates, species-area relationships and species abundance distributions consistent with the behaviours found in nature. From our results, we predict steady speciation rates, more species in one-dimensional environments than two-dimensional environments, three scaling regimes of species-area relationships and lognormal distributions of species abundance with an excess of rare species and a tail that may be approximated by Fisher's logarithmic series. These are consistent with dependences reported for, among others, global birds and flowering plants, marine invertebrate fossils, ray-finned fishes, British birds and moths, North American songbirds, mammal fossils from Kansas and Panamanian shrubs. Quantitative comparisons of specific cases are remarkably successful. Our biodiversity results provide additional evidence that species diversity arises without specific physical barriers. This is similar to heavy traffic flows, where traffic jams can form even without accidents or barriers.
The branching of new species from an ancestral population requires the evolution of reproductive isolation between groups of individuals. Geographic separation of sub-populations by natural barriers, if sustained for sufficiently long times, may lead to the accumulation of independent genetic changes in each group and to mating incompatibilities (Mayr, 2001; Fitzpatrick et al., 2009). A similar phenomenon may occur in the absence of barriers via isolation by distance if the population is distributed over large areas (de Aguiar et al., 2009; Etienne and Haegeman, 2011; Gavrilets et al., 2000). The first demonstration of this process was based on computer simulations employing agent-based models. Recently, analytical results were derived combining network theory, to model the spatial structure of the population, and an ansatz that accounts for the effect of forbidding mating between individuals that are too different genetically (de Aguiar and Bar-Yam, 2011). The main result obtained with this approach is an expression that indicates when speciation is possible as a function of the parameters describing the population. The aim of this work is to test this analytical result by comparing it with numerical simulations for a hermaphroditic population (de Aguiar et al., 2009) and for a population whose individuals are explicitly separated into males and females (Baptestini et al., 2013). We show that the analytical formula is indeed a very good overall description of the simulations and that the exponents describing dependence of the critical threshold of speciation with the parameters are in good agreement with the simulations.
Many quantitative genetic and adaptive dynamic models suggest that disruptive selection can maintain genetic polymorphism and be the driving force causing evolutionary divergence. These models also suggest that disruptive selection arises from frequency-dependent intraspecific competition. For convenience or historical precedence, these models assume that carrying capacity and competition functions follow a Gaussian distribution. Here, we propose a new analytical framework that relaxes the assumption of Gaussian competition and carrying capacity functions, and investigate how alternative shapes affect the likelihood of disruptive selection. We found that the shape of both carrying capacity and competition kernels interact to determine the likelihood of disruptive selection. For certain regions of the parametric space disruptive selection is facilitated, whereas for others it becomes more difficult. Our results suggest that the relationship between the degree of frequency dependence and the likelihood of disruptive selection is more complex than previously thought, depending on how resources are distributed and competition interference takes place. It is now important to describe the empirical patterns of resource distribution and competition in nature as a way to determine the likelihood of disruptive selection in natural populations.
Neutral speciation mechanisms based on isolation by distance and sexual selection, termed topopatric, have recently been shown to describe the observed patterns of abundance distributions and species-area relationships. Previous works have considered this type of process only in the context of hermaphrodic populations. In this work we extend a hermaphroditic model of topopatric speciation to populations where individuals are explicitly separated into males and females. We show that for a particular carrying capacity speciation occurs under similar conditions, but the number of species generated decreases as compared to the hermaphroditic case. Evolution results in fewer species having more abundant populations.Comment: 18 pages + 8 figure
Neutral models of speciation based on isolation by distance and assortative mating, termed topopatric, have shown to be successful in describing abundance distributions and species-area relationships. Previous works have considered this type of process in the context of haploid genomes. Here we discuss the implementation of two schemes of dominance to analyze the effects of diploidy: a complete dominance model in which one allele dominates over the other and a perfect codominant model in which heterozygous genotypes give rise to a third phenotype. In the case of complete dominance, we observe that speciation requires stronger spatial inbreeding in comparison to the haploid model. For perfect codominance, instead, speciation demands stronger genetic assortativeness. Nevertheless, once speciation is established, the three models predict the same abundance distributions even at the quantitative level, revealing the robustness of the original mechanism to describe biodiversity features.
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