The effect of gene flow on the coalescent time in the human-chimpanzee ancestral population

The coalescent process in the human-chimpanzee ancestral population is investigated using a model, which incorporates a certain time period of gene flow during the speciation process. a is a parameter to represent the degree and time of gene flow, and the model is identical to the null model with an instantaneous species split when a=infinity. A maximum likelihood (ML) method is developed to estimate a, and its power and reliability is investigated by coalescent simulations. The ML method is applied to nucleotide divergence data between human and chimpanzee. It is found that the null model with an instantaneous species split explains the data best, and no strong evidence for gene flow is detected. The result is discussed in the view of the mode of speciation. Another ML method is developed to estimate the male-female ratio (alpha) of mutation rate, in which the coalescent process in the ancestral population is taken into account.     

叶绿体DNA标记在谱系地理学中的应用研究进展

谱系地理学研究旨在探究历史上发生的影响目前遗传谱系系统发育和空间分布格局关系的生态与进化过程。叶绿体DNA具有单亲遗传、低突变率、单倍体等特征, 其分子标记不同程度地保留着植物长期进化的历史遗传痕迹, 有助于深度解析谱系地理变异的形成机制。本文探讨了上述特征是怎样影响分子标记的选择、扩大或缩小群体遗传结构分化、延长或缩短空间基因溯祖时间、促进或阻碍种间基因渐渗及谱系分选(复系、并系和单系形成)进程, 重点阐述了这些影响过程的理论基础, 并结合实际例子阐述谱系地理研究进展。由于位点间在突变率、选择强度及它们与漂变互作等方面存在异质性, 今后一个研究重点就是基于叶绿体全基因组序列分析谱系地理变化格局, 包括分析DNA位点间的基因渐渗或基因流动程度差异分布及沿着叶绿体DNA序列上谱系分选差异分布。     

Phylogeographic breaks in low-dispersal species: the emergence of concordance across gene trees

Computer simulations were used to investigate population conditions under which phylogeographic breaks in gene genealogies can be interpreted with confidence to infer the existence and location of historical barriers to gene flow in continuously distributed, low-dispersal species. We generated collections of haplotypic gene trees under a variety of demographic scenarios and analyzed them with regard to salient genealogical breaks in their spatial patterns. In the first part of the analysis, we estimated the frequency in which the spatial location of the deepest phylogeographic break between successive pairs of populations along a linear habitat coincided with a spatial physical barrier to dispersal. Results confirm previous reports that individual gene trees can show ‘haphazard’ phylogeographic discontinuities even in the absence of historical barriers to gene flow. In the second part of the analysis, we assessed the probability that pairs of gene genealogies from a set of population samples agree upon the location of a geographical barrier. Our findings extend earlier reports by demonstrating that spatially concordant phylogeographic breaks across independent neutral loci normally emerge only in the presence of longstanding historical barriers to gene flow. Genealogical concordance across multiple loci thus becomes a deciding criterion by which to distinguish between stochastic and deterministic causation in accounting for phylogeographic discontinuities in continuously distributed species.     

Go east: phylogeographies of Mauremys caspica and M. rivulata- discordance of morphology, mitochondrial and nuclear genomic markers and rare hybridization

In recent years many cases of hybridization and introgression became known for chelonians, requiring a better understanding of their speciation mechanisms. Phylogeographic investigations offer basic data for this challenge. We use the sister species Mauremys caspica and M. rivulata, the most abundant terrapins in the Near and Middle East and South-east Europe, as model. Their phylogeographies provide evidence that speciation of chelonians fits the allopatric speciation model, with both species being in the parapatric phase of speciation, and that intrinsic isolation mechanisms are developed during speciation. Hybridization between M. caspica and M. rivulata is very rare, suggesting that the increasing numbers of hybrids in other species are caused by human impact on environment (breakdown of ecological isolation). Genetic differentiation within M. caspica and M. rivulata resembles the paradigm of southern genetic richness and northern purity of European biota. However, in west Asia this pattern is likely to reflect dispersal and vicariance events older than the Holocene. For M. caspica three distinct Pleistocene refuges are postulated (Central Anatolia, south coast of Caspian Sea, Gulf of Persia). Morphologically defined subspecies within M. caspica are not supported by genetic data. This is one of the few studies available about the phylogeography of west and central Asian species.     

Refugial isolation versus ecological gradients

Hypotheses for divergence and speciation in rainforests generally fall into two categories: those emphasizing the role of geographic isolation and those emphasizing the role of divergent selection along gradients. While a majority of studies have attempted to infer mechanisms based on the pattern of species richness and congruence of geographic boundaries, relatively few have tried to simultaneously test alternative hypotheses for diversification. Here we discuss four examples, taken from our work on diversification of tropical rainforest vertebrates, in which we examine patterns of genetic and morphological variation within and between biogeographic regions to address two alternative hypotheses. By estimating morphological divergence between geographically contiguous and isolated populations under similar and different ecological conditions, we attempt to evaluate the relative roles of geographic isolation and natural selection in population divergence. Results suggest that natural selection, even in the presence of appreciable gene flow, can result in morphological divergence that is greater than that found between populations isolated for millions of years and, in some cases, even greater than that found between congeneric, but distinct, species. The relatively small phenotypic divergence that occurs among long-term geographic isolates in similar habitats suggests that morphological divergence via drift may be negligible and/or that selection is acting to produce similar phenotypes in populations occupying similar habitats. Our results demonstrate that significant phenotypic divergence: (1) is not necessarily coupled with divergence in neutral molecular markers; and (2) can occur without geographic isolation in the presence of gene flow.     

The anatomy of a ‘suture zone’ in Amazonian butterflies: a coalescent‐based test for vicariant geographic divergence and speciation

Attempts by biogeographers to understand biotic diversification in the Amazon have often employed contemporary species distribution patterns to support particular theories, such as Pleistocene rainforest refugia, rather than to test among alternative hypotheses. Suture zones, narrow regions where multiple contact zones and hybrid zones between taxa cluster, have been seen as evidence for past expansion of whole biotas that have undergone allopatric divergence in vicariant refuges. We used coalescent analysis of mutilocus sequence data to examine population split times in 22 pairs of geminate taxa in ithomiine and heliconiine butterflies. We test a hypothesis of simultaneous divergence across a suture zone in NE Peru. Our results reveal a scattered time course of diversification in this suture zone, rather than a tight cluster of split times. Additionally, we find rapid diversification within some lineages such as Melinaea contrasting with older divergence within lineages such as the Oleriina (Hyposcada and Oleria). These results strongly reject simple vicariance as a cause of the suture zone. At the same time, observed lineage effects are incompatible with a series of geographically coincident vicariant events which should affect all lineages similarly. Our results suggest that Pleistocene climatic forcing cannot readily explain this Peruvian suture zone. Lineage‐specific biological traits, such as characteristic distances of gene flow or varying rates of parapatric divergence, may be of greater importance.     

A parapatric propensity for breeding precludes the completion of speciation in common teal (Anas crecca, sensu lato)

Speciation is a process in which genetic drift and selection cause divergence over time. However, there is no rule dictating the time required for speciation, and even low levels of gene flow hinder divergence, so that taxa may be poised at the threshold of speciation for long periods of evolutionary time. We sequenced mitochondrial DNA (mtDNA) and eight nuclear introns (nuDNA) to estimate genomic levels of differentiation and gene flow between the Eurasian common teal (Anas crecca crecca) and the North American green‐winged teal (Anas crecca carolinensis). These ducks come into contact in Beringia (north‐eastern Asia and north‐western North America) and have probably done so, perhaps cyclically, since the Pliocene–Pleistocene transition, ～2.6 Ma, when they apparently began diverging. They have diagnosable differences in male plumage and are 6.9% divergent in the mtDNA control region, with only 1 of 58 crecca and 2 of 86 carolinensis having haplotypes grouping with the other. Two nuclear loci were likewise strongly structured between these teal (Φ_st ≥ 0.35), but six loci were undifferentiated or only weakly structured (Φ_st = 0.0–0.06). Gene flow between crecca and carolinensis was ～1 individual per generation in both directions in mtDNA, but was asymmetrical in nuDNA, with ～1 and ～20 individuals per generation immigrating into crecca and carolinensis, respectively. This study illustrates that species delimitation using a single marker oversimplifies the complexity of the speciation process, and it suggests that even with divergent selection, moderate levels of gene flow may stall the speciation process short of completion.     

Endemic bird species and conservation in the Cerrado Region, South America

A general overview of the largest South American savanna region, the Cerrado Region, is presented. Using simple approaches, I analyse the distribution of the bird species endemic to this region in order to test the hypothesis that areas harbouring local aggregates of restricted-range endemics have remained stable during the successive Quaternary climatic-vegetational changes. This hypothesis seems not to be generally valid for birds of the Cerrado Region, as restricted-range endemic species exist both in areas that remained stable (Central Goia´s Plateau) and areas that remain unstable (Araguaya and Parana~ River valleys, Espinhac¸o Plateau) during the Late Pleistocene. I also discuss some aspects of the conservation in the Cerrado Region, and suggest that the dry forests of the River Parana~ Valley, Goia´s, must be considered the area with the highest priority for the creation of new reserves in the Cerrado Region.