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The increasing aridity during the Last Glacial Maximum (LGM) has been proposed as a major factor affecting Neotropical species. The character and intensity of this change, however, remains the subject of ongoing debate. This review proposes an approach to test contrasting paleoecological hypotheses by way of their expected demographic and genetic effects on Neotropical cloud forest species. We reviewed 48 paleoecological records encompassing the LGM in the Neotropics. The records show contrasting evidence regarding the changes in precipitation during this period. Some regions remained fairly moist and others had a significantly reduced precipitation. Many paleoecological records within the same region show apparently conflicting evidence on precipitation and forest stability. From these data, we propose and outline two demographic/genetic scenarios for cloud forests species based on opposite precipitation regimes: the dry refugia and the moist forests hypotheses. We searched for studies dealing with the population genetic structure of cloud forest and other montane taxa and compared their results with the proposed models. To date, the few available molecular studies show insufficient genetic evidence on the predominance of glacial aridity in the Neotropics. In order to disentangle the climatic history of the Neotropics, the present study calls for a general multi-disciplinary approach to conduct future phylogeographic studies. Given the contradictory paleoecological information, population genetic data on Neotropical cloud forest species should be used to explicitly test the genetic consequences of competing paleoecological models.

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BackgroundEcological adaptation to host taxa is thought to result in mistletoe speciation via race formation. However, historical and ecological factors could also contribute to explain genetic structuring particularly when mistletoe host races are distributed allopatrically. Using sequence data from nuclear (ITS) and chloroplast (trnL-F) DNA, we investigate the genetic differentiation of 31 Psittacanthus schiedeanus (Loranthaceae) populations across the Mesoamerican species range. We conducted phylogenetic, population and spatial genetic analyses on 274 individuals of P. schiedeanus to gain insight of the evolutionary history of these populations. Species distribution modeling, isolation with migration and Bayesian inference methods were used to infer the evolutionary transition of mistletoe invasion, in which evolutionary scenarios were compared through posterior probabilities.ResultsOur analyses revealed shallow levels of population structure with three genetic groups present across the sample area. Nine haplotypes were identified after sequencing the trnL-F intergenic spacer. These haplotypes showed phylogeographic structure, with three groups with restricted gene flow corresponding to the distribution of individuals/populations separated by habitat (cloud forest localities from San Luis Potosí to northwestern Oaxaca and Chiapas, localities with xeric vegetation in central Oaxaca, and localities with tropical deciduous forests in Chiapas), with post-glacial population expansions and potentially corresponding to post-glacial invasion types. Similarly, 44 ITS ribotypes suggest phylogeographic structure, despite the fact that most frequent ribotypes are widespread indicating effective nuclear gene flow via pollen. Gene flow estimates, a significant genetic signal of demographic expansion, and range shifts under past climatic conditions predicted by species distribution modeling suggest post-glacial invasion of P. schiedeanus mistletoes to cloud forests. However, Approximate Bayesian Computation (ABC) analyses strongly supported a scenario of simultaneous divergence among the three groups isolated recently.ConclusionsOur results provide support for the predominant role of isolation and environmental factors in driving genetic differentiation of Mesoamerican parrot-flower mistletoes. The ABC results are consistent with a scenario of post-glacial mistletoe invasion, independent of host identity, and that habitat types recently isolated P. schiedeanus populations, accumulating slight phenotypic differences among genetic groups due to recent migration across habitats. Under this scenario, climatic fluctuations throughout the Pleistocene would have altered the distribution of suitable habitat for mistletoes throughout Mesoamerica leading to variation in population continuity and isolation. Our findings add to an understanding of the role of recent isolation and colonization in shaping cloud forest communities in the region.Electronic supplementary materialThe online version of this article (doi:10.1186/s12862-016-0648...

ResearchCite this article: Ramírez-Barahona S, Barrera-Redondo J, Eguiarte LE. 2016 Rates of ecological divergence and body size evolution are correlated with species diversification in scaly tree ferns. Proc. R. Soc. Variation in species richness across regions and between different groups of organisms is a major feature of evolution. Several factors have been proposed to explain these differences, including heterogeneity in the rates of species diversification and the age of clades. It has been frequently assumed that rapid rates of diversification are coupled to high rates of ecological and morphological evolution, leading to a prediction that remains poorly explored for most species: the positive association between ecological niche divergence, morphological evolution and species diversification. We combined a time-calibrated phylogeny with distribution, ecological and body size data for scaly tree ferns (Cyatheaceae) to test whether rates of species diversification are predicted by the rates at which clades have evolved distinct ecological niches and body sizes. We found that rates of species diversification are positively correlated with rates of ecological and morphological evolution, with rapidly diversifying clades also showing rapidly evolving ecological niches and body sizes. Our results show that rapid diversification of scaly tree ferns is associated with the evolution of species with comparable morphologies that diversified into similar, yet distinct, environments. This suggests parallel evolutionary pathways opening in different tropical regions whenever ecological and geographical opportunities arise. Accordingly, rates of ecological niche and body size evolution are relevant to explain the current patterns of species richness in this 'ancient' fern lineage across the tropics.

Aim We investigated changes in distribution of cloud forests during the last 130 kyr, and tested whether these changes explain the spatial patterns of genetic diversity of the tree fern Alsophila firma (Cyatheaceae), a species restricted to this habitat. Location Mexican cloud forests. Methods We sampled 204 individuals from 16 localities. Genetic data consisted of DNA sequences for five chloroplast microsatellites and one nuclear gene. We used distribution modelling to predict the historical distribution of cloud forests during the last glacial period, using two palaeoclimate models: the Model for Interdisciplinary Research on Climate (MIROC) and the Community Climate System Model (CCSM). We tested the correlation between temporal cloud forest stability and genetic diversity and used an approximate Bayesian computation (ABC) framework to test two plausible demographic scenarios. Results The range fluctuations observed for cloud forests during the last 130 kyr are key factors affecting the distribution of genetic variation in A. firma. Increased genetic diversity in areas with high temporal environmental suitability is probably the result of increased population sizes and higher interpopulation connectivity. In accordance with the expansion of cloud forests predicted by CCSM, the genetic data supported the scenario of a population expansion occurring c. 110 ka, followed by population divergence c. 20 ka. However, population dynamics involving expansion of suitable microclimates could reconcile the stability of cloud forests predicted by MIROC and the observed genetic patterns. Main conclusions The predicted changes in the distribution of cloud forests were congruent with the population genetics of A. firma. However, the choice of palaeoclimate model has a substantial impact on the inferences drawn from the observed genetic and demographic patterns. The use of alternative palaeoclimate hypotheses and biome modelling can provide a common analytical framework for evaluating the historical cohesiveness of forest communities.

Aim: The formation of the Trans-Mexican Volcanic Belt (TMVB) played an important role in driving inter-and intraspecific diversification at high elevations. However, Pleistocene climate changes and ecological factors might also contribute to plant genetic structuring along the volcanic belt. Here, we analysed phylogeographical patterns of the parrot-mistletoe Psittacanthus calyculatus to determine the relative contribution of these different factors. Location: Trans-Mexican Volcanic Belt. Methods: Using nuclear and chloroplast DNA sequence data for 370 individuals, we investigate the genetic differentiation of 35 populations across the species range.We conducted phylogenetic, population and spatial genetic analyses of P. calyculatus sequences along with ecological niche modelling and Bayesian inference methods to gain insight into the structuring of genetic variation of these populations.Results: Our analyses revealed population structure with three genetic groups corresponding to individuals from Oaxaca and those from the central-eastern and western TMVB regions. A significant genetic signal of demographic expansion, an eastto-west expansion predicted by species distribution modelling, and approximate Bayesian computation analyses strongly supported a scenario of habitat isolation and invasion of TMVB by P. calyculatus during the late-Pleistocene.Main conclusions: The genetic differentiation of P. calyculatus may be explained by the combined effects of (1) geographical isolation linked to the effects of the glacial/interglacial cycles and environmental factors, driving genetic differentiation from congeners into more xeric vegetation and (2) the invasion of TMVB from east to west, suggesting a role for both colonization and glacial/interglacial cycles models.

Existing global regionalization schemes for plants consider the compositional affinities among biotas, but these have not explicitly considered phylogenetic information. Here, we present for the first time, a phytogeographical delineation of the global vascular flora based on species-level evolutionary relationships.We analysed 8737 820 geographical occurrence records for vascular plants together with a time-calibrated phylogeny including 67 269 species. We constructed a global phylogenetic regionalization by estimating species composition and phylogenetic beta diversity among 200 km 9 200 km grid cells across the world.We identified de novo 16 phytogeographical units that are deeply split into two clusters: Laurasian and Gondwanan. Our regionalization broadly matches previous schemes, but also highlights the separation of the Gondwanan biota into an Holotropical cluster and an Australian-Neozealandic-Patagonian cluster. In contrast, no clear split among Laurasian and Gondwanan biotas was retrieved when omitting phylogenetic information.The integration of phylogenetic and geographical information provides new insights into the delineation of phytogeographical areas and their historical relationships, enabling the identification of three large, clearly differentiated biotas, here referred to as kingdoms: Holarctic, Holotropical, and Austral. Our results provide further evidence for delineating transition zones and show a clear latitudinal pattern of increasing evolutionary distinctiveness towards the poles.