Isolation by resistance explains genetic diversity in the Arremon brushfinches of northern Mesoamerica.

Genetic differentiation between and within natural populations is the result of the joint effects of neutral and adaptative processes. In addition, the spatial arrangement of the landscape promotes connectivity or creates barriers to gene flow, directly affecting speciation processes. In this study, we carried out a landscape genomics analysis using NextRAD data from a montane forest specialist bird complex, the Mesoamerican Chestnut-capped/Green-striped Brushfinch of the genus Arremon. Specifically, we examined population genomic structure using different assignment methods and genomic differentiation and diversity, and we tested alternative genetic isolation hypotheses at the individual level (e.g., isolation by barrier, IBB; isolation by environment, IBE; isolation by resistance, IBR). We found well-delimited genomic structuring (K = 5) across Mesoamerican montane forests in the studied group. Individual-level genetic distances among major montane ranges were mainly explained by IBR hypotheses in this sedentary Neotropical taxon. Our results uncover genetic distances/differentiation and patterns of gene flow in allopatric species that support the role of tropical mountains as spatial landscape drivers of biodiversity. IBR clearly supports a pattern of conserved niche-tracking of suitable habitat conditions and topographic complexity throughout glacial-interglacial dynamics.

The tangled evolutionary history of a long-debated Mesoamerican taxon: The Velazquez Woodpecker (Melanerpes santacruzi, Aves: Picidae).

The Velazquez Woodpecker Melanerpes santacruzi is a highly polytypic species distributed from east-central Mexico to northern Nicaragua. The ample variation in body size, barring of the plumage, and the coloration of nasal tufts, neck, and belly have fueled debates about the taxonomy and evolutionary history of the species; however, the processes generating these patterns of variation and the underlying population dynamics throughout the species' distribution remain poorly understood. Here, we employed reduced representation genome sequencing (NextRAD) and Ecological Niche Modeling methods to test the distinctiveness of the Velazquez Woodpecker based on this new set of genomic data and analyze the correspondence of the genetic structure and ecological differentiation with phenotypic variation and geographic distribution. From phylogenetic and demographic analyses including the Golden-Fronted (M. aurifrons) and Red-bellied Woodpecker (M. carolinus), we obtained results congruent with previous molecular phylogenies. The clades of M. santacruzi and M. carolinus-M. aurifrons are reciprocally monophyletic, although the sister group relationship of M. aurifrons is ambiguous. Using genetic and ecological analyses, we found that the species is structured into three genetically and ecologically differentiated groups comprising the subspecies (1) M. s. santacruzi, (2) M. s. dubius and (3) M. s. grateloupensis-polygrammus-veraecrucis. These groups diverged recently, with two splits between 250,000 and 150,000 years ago, and show a significant genetic admixture among them, especially in their current contact zones. Ecological and demographic models suggest the existence of intermittent areas of sympatry and connectivity among populations of M. santacruzi since the Last Interglacial period. We also found evidence of bi-directional gene flow between the species M. aurifrons and the nearby populations of M. santacruzi (M. s. grateloupensis), along the Sierra Madre Oriental in northeastern Mexico. Gene flow seems to be uneven, with prevalence of movement in the direction from M. aurifrons to M. s. grateloupensis.

The geography of evolutionary divergence in the highly endemic avifauna from the Sierra Madre del Sur, Mexico.

BACKGROUND: Mesoamerica is a remarkable region with a high geological and ecological complexity. Within northern Mesoamerica, the biotic province of the Sierra Madre del Sur (SMS) in southwestern Mexico harbors exceptionally high avian endemism and diversity. Herein, we searched for spatially and temporally concordant phylogeographic patterns, in four bird genera from three distinct avian orders co-distributed across Mesoamerica and investigated their causes through hypothesis testing regarding historical processes. Selected species include endemic and differentiated populations across the montane forests of Mesoamerica, and particularly within the SMS. RESULTS: We gathered mitochondrial DNA sequences for at least one locus from 177 individuals across all species. We assessed genetic structure, demographic history, and defined a framework for the coalescent simulations used in biogeographic hypothesis testing temporal and spatial co-variance. Our analyses suggested shared phylogeographic breaks in areas corresponding to the SMS populations, and between the main montane systems in Mesoamerica, with the Central Valley of Oaxaca and the Nicaragua Depression being the most frequently shared breaks among analyzed taxa. Nevertheless, dating analyses and divergence patterns observed were consistent with the hypothesis of broad vicariance across Mesoamerica derived from mechanisms operating at distinct times across taxa in the SMS. CONCLUSIONS: Our study provides a framework for understanding the evolutionary origins and historical factors enhancing speciation in well-defined regions within Mesoamerica, indicating that the evolutionary history of extant biota inhabiting montane forests is complex and often idiosyncratic.

Distributional patterns of Neotropical seasonally dry forest birds: a biogeographical regionalization.

Neotropical seasonally dry forests (NSDFs) are widely distributed and possess high levels of species richness and endemism; however, their biogeography remains only partially understood. Using species distribution modelling and parsimony analysis of endemicity, we analysed the distributional patterns of the NSDF avifauna in order to identify their areas of endemism and provide a better understanding of the historical relationships among those areas. The strict consensus trees revealed 17 areas of endemism for NSDFs, which involve four large regions: Baja California, Caribbean-Antilles islands, Mesoamerica and South America. These well-resolved clades are circumscribed by geographical and ecological barriers associated with the Gulf of California, the leading edge of the Caribbean plate, the Tehuantepec Isthmus, the Polochic-Motagua fault, the Nicaragua Depression, the Chocó forest, the Amazon basin and the Andean Cordillera. Relationships among groups of NSDFs found here suggest that evolution of their avifauna involved a mixture of vicariance and dispersal events. Our results support the idea of independent diversification patterns and biogeographical processes in each region, including those previously associated with the Pleistocene Arc Hypothesis for NSDFs of south-eastern South America. This study provides a biogeographical framework to open new lines of research related to the biotic diversification of NSDFs.

Assumption-versus data-based approaches to summarizing species' ranges.

For conservation decision making, species' geographic distributions are mapped using various approaches. Some such efforts have downscaled versions of coarse-resolution extent-of-occurrence maps to fine resolutions for conservation planning. We examined the quality of the extent-of-occurrence maps as range summaries and the utility of refining those maps into fine-resolution distributional hypotheses. Extent-of-occurrence maps tend to be overly simple, omit many known and well-documented populations, and likely frequently include many areas not holding populations. Refinement steps involve typological assumptions about habitat preferences and elevational ranges of species, which can introduce substantial error in estimates of species' true areas of distribution. However, no model-evaluation steps are taken to assess the predictive ability of these models, so model inaccuracies are not noticed. Whereas range summaries derived by these methods may be useful in coarse-grained, global-extent studies, their continued use in on-the-ground conservation applications at fine spatial resolutions is not advisable in light of reliance on assumptions, lack of real spatial resolution, and lack of testing. In contrast, data-driven techniques that integrate primary data on biodiversity occurrence with remotely sensed data that summarize environmental dimensions (i.e., ecological niche modeling or species distribution modeling) offer data-driven solutions based on a minimum of assumptions that can be evaluated and validated quantitatively to offer a well-founded, widely accepted method for summarizing species' distributional patterns for conservation applications.

Complex biogeographic scenarios revealed in the diversification of the largest woodpecker radiation in the New World.

Phylogenetic relationships and patterns of evolution within Melanerpes, one of the most diverse groups of New World woodpeckers (22-23 lineages), have been complicated due to complex plumages and morphological adaptations. In an attempt to resolve these issues, we obtained sequence data from four nuclear introns and two mitochondrial protein-coding genes for 22 of the 24 currently recognized species in the genus. We performed phylogenetic analyses involving Maximum Likelihood and Bayesian Inference, species-tree divergence dating, and biogeographic reconstructions. Tree topologies from the concatenated and species-tree analyses of the mtDNA and nDNA showed broadly similar patterns, with three relatively well-supported groups apparent: (a) the Sphyrapicus clade (four species); (b) the typical Melanerpes clade, which includes temperate and subtropical dry forest black-backed species; and (c) the mostly barred-backed species, here referred to as the "Centurus" clade. The phylogenetic position of Melanerpes superciliaris regarding the rest of Melanerpes is ambiguous as it is recovered as sister to the rest of Melanerpes or as sister to a group including Sphyrapicus+Melanerpes. Our species tree estimations recovered the same well-delimited highly-supported clades. Geographic range evolution (estimated in BioGeoBEARS) was best explained by a DIVALIKE+j model, which includes vicariance, founder effect speciation, and anagenetic dispersal (range expansion) as important processes involved in the diversification of the largest radiation of woodpeckers in the New World.

Rapid postglacial diversification and long-term stasis within the songbird genus Junco: phylogeographic and phylogenomic evidence.

Natural systems composed of closely related taxa that vary in the degree of phenotypic divergence and geographic isolation provide an opportunity to investigate the rate of phenotypic diversification and the relative roles of selection and drift in driving lineage formation. The genus Junco (Aves: Emberizidae) of North America includes parapatric northern forms that are markedly divergent in plumage pattern and colour, in contrast to geographically isolated southern populations in remote areas that show moderate phenotypic divergence. Here, we quantify patterns of phenotypic divergence in morphology and plumage colour and use mitochondrial DNA genes, a nuclear intron, and genomewide SNPs to reconstruct the demographic and evolutionary history of the genus to infer relative rates of evolutionary divergence among lineages. We found that geographically isolated populations have evolved independently for hundreds of thousands of years despite little differentiation in phenotype, in sharp contrast to phenotypically diverse northern forms, which have diversified within the last few thousand years as a result of the rapid postglacial recolonization of North America. SNP data resolved young northern lineages into reciprocally monophyletic lineages, indicating low rates of gene flow even among closely related parapatric forms, and suggesting a role for strong genetic drift or multifarious selection acting on multiple loci in driving lineage divergence. Juncos represent a compelling example of speciation in action, where the combined effects of historical and selective factors have produced one of the fastest cases of speciation known in vertebrates.

Geographic isolation drives divergence of uncorrelated genetic and song variation in the Ruddy-capped Nightingale-Thrush (Catharus frantzii; Aves: Turdidae).

Montane barriers influence the evolutionary history of lineages by promoting isolation of populations. The effects of these historical processes are evident in patterns of differentiation among extant populations, which are often expressed as genetic and behavioral variation between populations. We investigated the effects of geographic barriers on the evolutionary history of a Mesoamerican bird by studying patterns of genetic and vocal variation in the Ruddy-capped Nightingale-Thrush (Turdidae: Catharus frantzii), a non-migratory oscine bird that inhabits montane forests from central Mexico to Panama. We reconstructed the phylogeographic history and estimated divergence times between populations using Bayesian and maximum likelihood methods. We found strong support for the existence of four mitochondrial lineages of C. frantzii corresponding to isolated mountain ranges: Sierra Madre Oriental; Sierra Madre del Sur; the highlands of Chiapas, Guatemala, and El Salvador; and the Talamanca Cordillera. Vocal features in C. frantzii were highly variable among the four observed clades, but vocal variation and genetic variation were uncorrelated. Song variation in C. frantzii suggests that sexual selection and cultural drift could be important factors driving song differentiation in C. frantzii.

Response of the endangered tropical dry forests to climate change and the role of Mexican Protected Areas for their conservation.

Assuming that co-distributed species are exposed to similar environmental conditions, ecological niche models (ENMs) of bird and plant species inhabiting tropical dry forests (TDFs) in Mexico were developed to evaluate future projections of their distribution for the years 2050 and 2070. We used ENM-based predictions and climatic data for two Global Climate Models, considering two Representative Concentration Pathway scenarios (RCP4.5/RCP8.5). We also evaluated the effects of habitat loss and the importance of the Mexican system of protected areas (PAs) on the projected models for a more detailed prediction of TDFs and to identify hot spots that require conservation actions. We identified four major distributional areas: the main one located along the Pacific Coast (from Sonora to Chiapas, including the Cape and Bajío regions, and the Balsas river basin), and three isolated areas: the Yucatán peninsula, central Veracruz, and southern Tamaulipas. When considering the effect of habitat loss, a significant reduction (~61%) of the TDFs predicted area occurred, whereas climate-change models suggested (in comparison with the present distribution model) an increase in area of 3.0-10.0% and 3.0-9.0% for 2050 and 2070, respectively. In future scenarios, TDFs will occupy areas above its current average elevational distribution that are outside of its present geographical range. Our findings show that TDFs may persist in Mexican territory until the middle of the XXI century; however, the challenges about long-term conservation are partially addressed (only 7% unaffected within the Mexican network of PAs) with the current Mexican PAs network. Based on our ENM approach, we suggest that a combination of models of species inhabiting present TDFs and taking into account change scenarios represent an invaluable tool to create new PAs and ecological corridors, as a response to the increasing levels of habitat destruction and the effects of climate change on this ecosystem.

Multilocus analysis of intraspecific differentiation in three endemic bird species from the northern Neotropical dry forest.

Among-species phylogeographic concordance provides insight into the common processes driving lineage divergence in a particular region. However, identifying the processes that caused phylogeographic breaks is not always straight forward, and inferring past environmental conditions in combination with documented geologic events is sometimes necessary to explain current patterns. We searched for concordant phylogeographic patterns and investigated their causes in three bird species (Momotus mexicanus, Melanerpes chrysogenys, and Passerina leclancherii) that belong to three different avian orders and are endemic to the northernmost range of the Neotropical dry forest. We obtained mitochondrial DNA (ND2 and COI or cyt b) and nuclear DNA (20454, GAPDH, MUSK, and TGFB) sequences for at least one locus from 162 individuals across all species and defined climatically stable areas using environmental niche model projections for the last 130,000 years to have a paleoenvironmental framework for the phylogeographic results. All three species showed marked phylogeographic structure, with breaks found in roughly similar areas, such as the border between the Mexican states of Guerrero and Oaxaca, and between southern Jalisco and Michoacán. Both of these regions are known biogeographic breaks among other taxa. Patterns of genetic diversity and differentiation were partially compatible with climatically stable areas. Coalescent analyses revealed recent population growth and estimated the deeper haplogroup divergence of all three taxa to have occurred within the last 600,000 years. The phylogeographic patterns found are noteworthy because they are maintained in a relatively small area for bird species with continuous ranges, and highlight a unique situation when compared to phylogeographic patterns found in other studies of Neotropical birds that have stressed the role of geographic barriers to explain intraspecific differentiation. Our results point to a scenario of population isolation resulting in the present phylogeographic structure, likely a result of historical climate fluctuations that have fragmented and reconnected the Neotropical dry forest. This study contributes to a growing body of evidence indicating active diversification of endemic lineages in the northern Neotropical dry forest region.

Evolutionary diversification and speciation in rodents of the Mexican lowlands: the Peromyscus melanophrys species group.

Despite some studies of the species groups within the genus Peromyscus have been performed, both evolutionary relationships among species within groups and group composition have remained controversial. In this study, we address phylogenetic relationships among species in the Peromyscus melanophrys group (P. melanophrys, P. perfulvus, and P. mekisturus), using a molecular phylogenetic analysis. This analysis is the first to include the poorly known P. mekisturus. We conducted maximum likelihood and Bayesian inference analyses with the ND3, tRNA-Arginine, ND4L, and partial ND4 mitochondrial genes, and the GHR nuclear gene. We consistently recovered a P. melanophrys group that is monophyletic with respect to the set of outgroups. Also, we recovered two distinct clades within P. perfulvus and two within P. melanophrys, one of which contain P. mekisturus among other P. melanophrys, all with geographic consistency. According to our divergence time estimates, the P. melanophrys group diverged during the Pliocene and the main diversification events within the group occurred at the end of the Pliocene and through the Pleistocene.

Molecular evidence of the taxonomic status of western Mexican populations of Pliaethornis longirostris (Aves: Trochilidae).

Species diversity is largely underestimated by current taxonomy, precluding a precise understanding of evolutionary processes. Genetic data have increased our understanding of that cryptic diversity, and multilocus studies are now desirable. In this study, we used mitochondrial and nuclear DNA sequences to evaluate the taxonomic status of the western Mexico's populations of Phaethornis longirostris. We found differences of 4.2 % in mtDNA and different alleles for one nDNA locus between western and eastern Mexican populations. Molecular and morphological evidence support the separation of these populations (P. 1. mexicanus and P. 1. griseoventer) as the species Phaethornis mexicanus Hartert 1897. Phaethornis mexicanus is endemic to western Mexico and sister to the remaining populations of P. longirostris. The speciation of P mexicanus probably occurred around 880,000 years ago by a vicariant event involving climatic-vegetational changes.

[Biogeographic regionalization of the mammals of tropical evergreen forests in Mesoamerica]. / Regionalización biogeográfica de la mastofauna de los bosques tropicales perennifolios de Mesoamérica.

Mesoamerica is a biologically complex zone that expands from Southern Mexico to extreme Northern Colombia. The biogeographical patterns and relationships of the mammalian fauna associated to the Mesoamerican Tropical Evergreen Forest (MTEF) are poorly understood, in spite of the wide distribution of this kind of habitat in the region. We compiled a complete georeferenced database of mammalian species distributed in the MTEF of specimens from museum collections and scientific literature. This database was used to create potential distribution maps through the use of environmental niche models (ENMs) by using the Genetic Algorithm for Rule-Set Production (GARP) using 22 climatic and topographic layers. Each map was used as a representation of the geographic distribution of the species and all available maps were summed to obtain general patterns of species richness in the region. Also, the maps were used to construct a presence-absence matrix in a grid of squares of 0.5 degrees of side, that was analyzed in a Parsimony Analysis of Endemicity (PAE), which resulted in a hypothesis of the biogeographic scheme in the region. We compiled a total of 41 527 records of 233 species of mammals associated to the MTEF. The maximum concentration of species richness (104-138 species) is located in the areas around the Isthmus of Tehuantepec, Northeastern Chiapas-Western Guatemala, Western Honduras, Central Nicaragua to Northwestern Costa Rica and Western Panama. The proposed regionalization indicates that mammalian faunas associated to these forests are composed of two main groups that are divided by the Isthmus of Tehuantepec in Oaxaca in: a) a Northern group that includes Sierra Madre of Chiapas-Guatemala and Yucatan Peninsula; and b) an austral group, that contains the Pacific slope of Chiapas towards the South including Central America. Some individual phylogenetic studies of mammal species in the region support the relationships between the areas of endemism proposed, which suggest a common biogeographical history. In spite that Mesoamerica is considered one of the most important hotspots for biological conservation, the poor knowledge of the biogeographic patterns, the scarcity of protected areas, and the high rate of habitat transformation due to human activities, make prioritary the development of conservation strategies that include patterns of species richness, endemism, and mammalian associations.

Speciation in an avian complex endemic to the mountains of Middle America (Ergaticus, Aves: Parulidae).

The implementation of the phylogeographic approach for the study of biodiversity is critical in poorly sampled regions like the montane systems of Middle America, as complex evolutionary histories often result in the presence of independent lineages not properly considered by traditional taxonomy. Herein we sequenced 2370 bp of mtDNA (ND2, cyt b and ATPase) from 81 individuals of Ergaticus, a complex of birds endemic to the montane forests of Middle America. Although current taxonomy recognizes two species, the results reveal considerable genetic structure with the presence of four mtDNA lineages. Two of these lineages within Ergaticus ruber evidence the need of a revaluation of the species limits for this taxon. The general phylogeographic pattern can be explained as a consequence of relative isolation of the populations in different mountain ranges separated by low elevation barriers. Most population groups did not show signals of demographic expansion with the exception of the one corresponding to clade 1. The divergence time estimates point to the Pleistocene as an important time period for the diversification of this complex.

The role of historical and contemporary processes on phylogeographic structure and genetic diversity in the Northern Cardinal, Cardinalis cardinalis.

BACKGROUND: Earth history events such as climate change are believed to have played a major role in shaping patterns of genetic structure and diversity in species. However, there is a lag between the time of historical events and the collection of present-day samples that are used to infer contemporary population structure. During this lag phase contemporary processes such as dispersal or non-random mating can erase or reinforce population differences generated by historical events. In this study we evaluate the role of both historical and contemporary processes on the phylogeography of a widespread North American songbird, the Northern Cardinal, Cardinalis cardinalis. RESULTS: Phylogenetic analysis revealed deep mtDNA structure with six lineages across the species' range. Ecological niche models supported the same geographic breaks revealed by the mtDNA. A paleoecological niche model for the Last Glacial Maximum indicated that cardinals underwent a dramatic range reduction in eastern North America, whereas their ranges were more stable in México. In eastern North America cardinals expanded out of glacial refugia, but we found no signature of decreased genetic diversity in areas colonized after the Last Glacial Maximum. Present-day demographic data suggested that population growth across the expansion cline is positively correlated with latitude. We propose that there was no loss of genetic diversity in areas colonized after the Last Glacial Maximum because recent high-levels of gene flow across the region have homogenized genetic diversity in eastern North America. CONCLUSION: We show that both deep historical events as well as demographic processes that occurred following these events are critical in shaping genetic pattern and diversity in C. cardinalis. The general implication of our results is that patterns of genetic diversity are best understood when information on species history, ecology, and demography are considered simultaneously.

The differential effect of lowlands on the phylogeographic pattern of a Mesoamerican montane species (Lepidocolaptes affinis, Aves: Furnariidae).

We sequenced 1869 bp of mtDNA (cyt b and ND2) from 80 specimens of Lepidocolaptes affinis, a montane bird species of Mesoamerica, sampled at 34 localities from Mexico to Costa Rica. The species showed moderate genetic diversity (π=0.0045) and phylogeographic structure (Φ(ST)=0.12-0.95). The phylogeographic pattern indicated the Nicaragua Depression has prevented gene flow whereas populations on the two sides of the Isthmus of Tehuantepec did not show strong genetic differentiation. In Mexico, the population of the Sierra Madre Oriental was composed of two different lineages. In general, our data did not support a scenario of historical demographic expansion, and matched partially the phylogeographic patterns of other Mesoamerica montane species.

Molecular systematics and evolution of the Cyanocorax jays.

Phylogenetic relationships were studied in the genus Cyanocorax (Aves: Corvidae) and related genera, Psilorhinus and Calocitta, a diverse group of New World jays distributed from the southern United States south to Argentina. Although the ecology and behavior of some species in the group have been studied extensively, lack of a molecular phylogeny has precluded rigorous interpretations in an evolutionary framework. Given the diverse combinations of plumage coloration, size, and morphology, the taxonomy of the group has been inconsistent and understanding of biogeographic patterns problematic. Moreover, plumage similarity between two geographically disjuct species, the Tufted jay (Cyanocorax dickeyi) from western Mexico and the White-tailed jay (C. mystacalis) from western Ecuador and Peru, has puzzled ornithologists for decades. Here, a phylogeny of all species in the three genera is presented, based on study of two mitochondrial and three nuclear genes. Phylogenetic trees revealed the non-monophyly of Cyanocorax, and the division of the whole assemblage in two groups: "Clade A" containing Psilorhinus morio, both species in Calocitta,Cyanocorax violaceus, C. caeruleus, C. cristatellus, and C. cyanomelas, and "Clade B" consisting of the remaining species in Cyanocorax. Relationships among species in Clade A were ambiguous and, in general, not well resolved. Within Clade B, analyses revealed the monophyly of the "Cissilopha" jays and showed no evidence for a sister relationship between C. mystacalis and C. dickeyi. The phylogenetic complexity of lineages in the group suggests several complications for the understanding biogeographic patterns, as well as for proposing a taxonomy that is consistent with morphological variation. Although multiple taxonomic arrangements are possible, recommendations are for recognizing only one genus, Cyanocorax, with Psilorhinus and Calocitta as synonyms.

RELICT HUMID TROPICAL FOREST IN MEXICO PROMOTES DIFFERENTIATION IN BARRED WOODCREEPERS Dendrocolaptes (AVES: FURNARIIDAE).

Humid tropical forests in Mesoamerica are distributed along the Atlantic slope and, in scattered locations, along the Mexican Pacific slope. These poorly explored Mexican forests include microendemic bird species. Two species in the genus Dendrocolaptes occur in lowland and foothill humid tropical forests of Mesoamerica. One of these, D. sanctithomae, is comprised of four subspecies, of which the two that occur in Mexico, D. s. sanctithomae and D. s. sheffleri, are distinctly different morphologically, and the latter is a poorly known microendemic taxon of the Mexican Pacific humid tropical forest in the Sierra Madre del Sur. We used both nuclear (nDNA) and mitochondrial DNA (mtDNA) to evaluate the genetic variation of D. sanctithomae in Mexico, and complemented this with a quantitative analysis of phenotypic traits. We also conducted analyses of environmental niche models to test the hypothesis of niche differentiation of D. s. sheffleri from other taxa of D. sanctithomae. Our phylogenetic reconstructions of mtDNA consistently recovered D. s. sheffleri and D. s. sanctithomae as reciprocally monophyletic, while they shared alleles of nDNA. These mtDNA differences are comparable with differences reported between other Dendrocolaptes sister-taxa pairs. Our analysis of phenotypic traits also indicated that the taxa differ in measurements of hallux and feather barring. In contrast, niche differentiation tests suggest that the niches of both taxa are more similar than expected by chance. Our evidence leads us to propose species status for D. sheffleri. This is an additional example of recent speciation in Mexico that indicates active and peripatric evolutionary differentiation in the northern Neotropics.

Influence of phylogenetic structure and climate gradients on geographical variation in the morphology of Mexican flycatcher forests assemblages (Aves: Tyrannidae).

Morphological variation is strongly related to variation in the ecological characteristics and evolutionary history of each taxon. To explore how geographical variation in morphology is related to different climatic gradients and phylogenetic structure, we analyzed the variation of morphological traits (body size, bill, and wing) of 64 species of tyrant flycatchers (Tyrannidae) distributed in Mexico. We measured these morphological traits in specimens from biological collections and related them to the climatic and topographic data of each collection locality. We performed the analyses separately at two levels: (1) the regional level and (2) the assemblage level, which was split into (assemblage I) lowland forests and (assemblage II) highland forests and other vegetation types. We also calculated the phylogenetic structure of flycatchers of each locality in order to explore the influence of climatic variables and the phylogenetic structure on the morphological variation of tyrant flycatchers, by means of linear mixed-effects models. We mapped the spatial variation of the relationship between morphological traits and environmental gradients, taking into account the phylogenetic structure. Important climatic variables explaining the morphological variation were those of temperature ranges (seasonality) and the results suggest that the phylogenetic clustering increases towards the highlands of Sierra Madre Oriental and Sierra Madre del Sur, and the lowlands of Balsas Depression. For the regional level, the spatial distribution of body size showed a pattern coincident with Bergmann's rule, with increasing in size from south to north. In the tropical lowland forests assemblage, body size tend to increase in seasonally dry forests (western Mexico) and decrease in the humid ones (eastern Mexico). In the assemblage of highland forests and other types of vegetation, morphological trait values increased northeast to southwest. Phylogenetic structure helped to explain the variation of morphology at the assemblage level but not at the regional level. The patterns of trait variation in the lowland and highland assemblages suggest that parts of morphological variation are explained both by the climatic gradients and by the lineage relatedness of communities. Overall, our results suggest that morphological variation is best explained by a varied set of variables, and that regression models representing this variation, as well as integrating phylogenetic patterns at different community levels, provide a new understanding of the mechanisms underlying the links among biodiversity, its geographical setting, and environmental change.

Environmental heterogeneity explains coarse-scale ß-diversity of terrestrial vertebrates in Mexico.

We explored the hypothesis that high ß-diversity of terrestrial vertebrates of Mexico is associated with a high environmental heterogeneity (HEH) and identify the drivers of ß-diversity at different spatial scales. We used distribution range maps of 2,513 species of amphibians, reptiles, mammals, and birds occurring in Mexico. We estimated ß-diversity for each taxon at four spatial scales (grid cells of 2°, 1°, 0.5° and 0.25°) using the multiplicative formula of Whittaker ßw. For each spatial scale, we derived 10 variables of environmental heterogeneity among cells based on raw data of temperature, precipitation, elevation, vegetation and soil. We applied conditional autoregressive models (CAR) to identify the drivers of ß-diversity for each taxon at each spatial scale. CARs increased in explanatory power from fine-to-coarse spatial scales in amphibians, reptiles and mammals. The heterogeneity in precipitation including both, coefficient of variation (CV) and range of values (ROV), resulted in the most important drivers of ß-diversity of amphibians; the heterogeneity in temperature (CV) and elevation (ROV) were the most important drivers of ß-diversity for reptiles; the heterogeneity in temperature (ROV) resulted in the most important driver in ß-diversity for mammals. For birds, CARs resulted significant at fine scales (grid cells of 0.5° and 0.25°), and the precipitation (ROV and CV), temperature (ROV), and vegetation (H) and soil (H) were heterogeneity variables retained in the model. We found support for the hypothesis of environmental heterogeneity (HEH) for terrestrial vertebrates at coarse scales (grid cell of 2°). Different variables of heterogeneity, mainly abiotic, were significant for each taxon, reflecting physiological differences among terrestrial vertebrate groups. Our study revealed the importance of mountain areas in the geographic patterns of ß-diversity of terrestrial vertebrates in Mexico. At a coarse scale, specific variables of heterogeneity can be used as a proxy of ß-diversity for amphibians and reptiles.