Speciation by distance and the evolution of ring species

Author(s)

Ayana de Brito Martins

• Document type: Doctoral Thesis
• Press: São Paulo.
• Institution: Universidade de São Paulo (USP). Instituto de Biociências (IBIOC/SB)
• Defense date: 2014-08-15

Examining board members

Marcus Aloizio Martinez de Aguiar; José Garcia Vivas Miranda; Paulo Inácio de Knegt López de Prado; Tiago Bosisio Quental; Clarisse Palma da Silva

• Advisor: Marcus Aloizio Martinez de Aguiar

Abstract

Ring species are circular chains of gradually changing taxa which are formed when a population expands around a geographic barrier in such a way that the two expanding fronts, which meet after many generations, are reproductively isolated. These structures are considered a prime example of speciation by distance, a process in which the acquisition of reproductive isolation depends on the attenuation of gene flow with distance. Geographical barriers that limit dispersal have played a central role in the study of speciation, since their presence has the potential of facilitating divergence. In addition, dispersal is also limited by endogenous factors, so that the area accessible to each individual is less than total area of the species distribution. In speciation by distance, this limitation is the main mechanism promoting the reduction of gene flow. Using agent-based models, we simulated the expansion and divergence of a population around a central barrier with ongoing gene flow. Our results show that ring species are unstable to speciation or mixing, but can persist for extended times, even in the absence of environmental selection. This persistence is affected by landscape configuration and we suggest that the shape of the distribution area of the greenish warbler ring species, one of the best documented examples of this phenomenon, may be important for its existence. These results imply that agent-based models can aid the understanding of patterns of spatial distribution of genetic diversity and the effect of spatial attributes in particular cases. Given the demonstration of ring species in the simulations, we investigated the conditions for their formation in more detail. We conclude that, in the absence of environmental selection, these complexes are rare and form only under very restricted conditions which depend on landscape, population and individual features. However, population structuring leading to the acquisition of reproductive isolation is better explained by landscape configuration than by local mating. We suggest that speciation by distance can be particularly favored in the case of ring species due to population expansion, since under these circumstances genetic drift can enhance differential allele fixation. The conditions for ring species formation by speciation by distance are very limited, which by itself could explain their rarity. Since species are subject to environmental variation, gaps in their distributions may appear over time. Ring species may be especially susceptible to the emergence of gaps, since their formation is favored by particularly narrow areas. Furthermore, in the absence of environmental selection, these complexes are intrinsically unstable due to the presence of gene flow, and are expected to completely speciate or mix. This transience in time is another factor contributing to the rarity of ring species (AU)

• FAPESP's process: 10/10155-3 - Spatially explicit speciation models: formation of ring species and the role of sex chromossomes in reproductive isolation
• Grantee: Ayana de Brito Martins
• Support Opportunities: Scholarships in Brazil - Doctorate