Distributional shifts – not geographic isolation – as a probable driver of montane species divergence

As biodiversity hotspots, montane regions have been a focus of research to understand the divergence process. Like their oceanic counterparts, the diversity of the ‘sky islands’ might be ascribed to geographic isolation of mountaintops. However, because the sky islands, and especially those in northern latitudes, are subject to extreme climatic events such as the glacial cycles that drove both altitudinal and geographical shifts in species’ distributions, the dynamic colonization process is also a possible factor driving divergence. Here we test these two hypotheses (i.e. isolation versus colonization) in a flightless montane grasshopper, Melanoplus oregonensis, which is a member of a diverse group that radiated across the Rocky Mountains of North America. Using approximate Bayesian computation (ABC) and spatially explicit simulations that account for spatial heterogeneity and temporal shifts in species distributions, we show that a colonization model of the sky islands from refugial populations provides a significantly better fit to the empirical genetic data than a model of the geographic isolation among sky islands. Moreover, support for the colonization model holds irrespective of whether the movement of individuals was modeled as a diffusion process or was informed by differences in habitat suitabilities across the landscape. With validation analyses to confirm the models provide a good fit to the data, as well as general power and quality analyses, the research not only adds to a growing body of work on the complex dynamics underlying montane biodiversity, but it also provides much needed evaluation of competing hypotheses based on explicit models of the divergence process, as opposed to inferences about diversification drivers from species diversity patterns.     

Climatic Niche Dynamics and Its Role in the Insular Endemism of <Emphasis Type="Italic">Anolis</Emphasis> Lizards

Insular systems are usually characterized by have a high species diversity, endemism, and evolutionary uniqueness. Although ecological and evolutionary factors shaping insular diversity and endemism are relatively well established, there is a little understanding about climatic niche dynamics for many insular adaptive radiations. Here, we evaluate the tempo and mode of climatic niche evolution in an iconic insular radiation of lizards. By using an extensive dataset of phylogenetic and coarse-grain climatic niches, we evaluated phylogenetic niche divergence and niche conservatism across temporal and spatial scales in the Caribbean Anolis lizard radiation. We found several instances of niche shifts during the anole radiation across islands. Many of these niche shifts converged to similar climatic regimes between different islands. Furthermore, we find evidence that single-island endemic species are more limited by low suitability of climatic conditions outside its native islands than oceanic barriers due to the high climatic heterogeneity observed at least between Greater Antillean islands. These results suggest that within-lineage climatic niche conservatism has been prevalent in short time scales and likely played a role driving the exceptional insular endemism observed today.     

Population genomic evidence for multiple Pliocene refugia in a montane‐restricted harvestman (Arachnida,Opiliones, Sclerobunus robustus) from the southwestern United States

The integration of ecological niche modelling into phylogeographic analyses has allowed for the identification and testing of potential refugia under a hypothesis‐based framework, where the expected patterns of higher genetic diversity in refugial populations and evidence of range expansion of nonrefugial populations are corroborated with empirical data. In this study, we focus on a montane‐restricted cryophilic harvestman, Sclerobunus robustus, distributed throughout the heterogeneous Southern Rocky Mountains and Intermontane Plateau of southwestern North America. We identified hypothetical refugia using ecological niche models (ENMs) across three time periods, corroborated these refugia with population genetic methods using double‐digest RAD‐seq data and conducted population‐level phylogenetic and divergence dating analyses. ENMs identify two large temporally persistent regions in the mid‐latitude highlands. Genetic patterns support these two hypothesized refugia with higher genetic diversity within refugial populations and evidence for range expansion in populations found outside hypothesized refugia. Phylogenetic analyses identify five to six genetically divergent, geographically cohesive clades of S. robustus. Divergence dating analyses suggest that these separate refugia date to the Pliocene and that divergence between clades pre‐dates the late Pleistocene glacial cycles, while diversification within clades was likely driven by these cycles. Population genetic analyses reveal effects of both isolation by distance (IBD) and isolation by environment (IBE), with IBD more important in the continuous mountainous portion of the distribution, while IBE was stronger in the populations inhabiting the isolated sky islands of the south. Using model‐based coalescent approaches, we find support for postdivergence migration between clades from separate refugia.     

Lineage diversification and historical demography of a sky island salamander,Plethodon ouachitae,from the Interior Highlands

Sky islands provide ideal opportunities for understanding how climatic changes associated with Pleistocene glacial cycles influenced species distributions, genetic diversification, and demography. The salamander Plethodon ouachitae is largely restricted to high‐elevation, mesic forest on six major mountains in the Ouachita Mountains. Because these mountains are separated by more xeric, low‐elevation valleys, the salamanders appear to be isolated on sky islands where gene flow among populations on different mountains may be restricted. We used DNA sequence data along with ecological niche modelling and coalescent simulations to test several hypotheses related to diversifications in sky island habitats. Our results revealed that P. ouachitae is composed of seven well‐supported lineages structured across six major mountains. The species originated during the Late Pliocene, and lineage diversification occurred during the Middle Pleistocene in a stepping stone fashion with a cyclical pattern of dispersal to a new mountain followed by isolation and divergence. Diversification occurred primarily on an east–west axis, which is likely related to the east–west orientation of the Ouachita Mountains and the more favourable cooler and wetter environmental conditions on north slopes compared to south‐facing slopes and valleys. All non‐genealogical coalescent methods failed to detect significant population expansion in any lineages. Bayesian skyline plots showed relatively stable population sizes over time, but indicated a slight to moderate amount of population growth in all lineages starting approximately 10 000–12 000 years ago. Our results provide new insight into sky island diversifications from a previously unstudied region, and further demonstrate that climatic changes during the Pleistocene had profound effects on lineage diversification and demography, especially in species from environmentally sensitive habitats in montane regions.     

The role of niche divergence and phenotypic adaptation in promoting lineage diversification in the Sage Sparrow (Artemisiospiza belli,Aves: Emberizidae)

Niche divergence or conservatism and phenotypic adaptation are important in lineage diversification. We used mitochondrial DNA (mtDNA), morphology and ecological niche models to examine these processes in three subspecies of Sage Sparrow (Artemisiospiza belli belli, A. b. canescens and A. b. nevadensis) that breed in bioclimatically diverse ecoregions in western North America. Overall, mtDNA and morphology are congruent with subspecies, ecoregion and bioclimatic niche. Niche divergence, rather than niche conservatism, accompanied by phenotypic adaptation, is associated with lineage diversification between subspecies. This diversification has occurred with and without physical barriers or accompanying genetic divergence. Populations of A. b. canescens are divided by a montane barrier into two bioclimatic regions (San Joaquin Valley, Mojave Desert), where they are indistinguishable phenotypically, but show distinctive genetic patterns. Although there is no physical barrier between A. b. canescens in the San Joaquin Valley and A. b. belli in the Coast Ranges, these populations occupy different bioclimatic niches and are phenotypically, but not genetically, diagnosable. Niche overlap is greatest between A. b. canescens from the Mojave Desert and A. b. nevadensis from the Great Basin, yet these subspecies maintain distinctive phenotypes and mtDNA, even in local secondary contact and sympatry. Palaeoclimatic niche models for the Last Glacial Maximum (c. 21 000 bp ) and the Last Interglacial (c. 120 000 bp ) suggest that ecoregionally distinct populations of Artemisiospiza belli experienced different Pleistocene range fluctuations and glacial refugia, with temporal niche conservatism. Populations probably reached their current distributions as favourable climates and habitats expanded after the last glaciation. © 2012 The Linnean Society of London, Biological Journal of the Linnean Society, 2012, ??, ??–??.     

Genetic diversity and ecological niche modelling of the restricted Recordia reitzii (Verbenaceae) from southern Brazilian Atlantic forest

Genetic diversity analyses, coupled with ecological niche modelling (ENM) of species with a restricted distribution, may provide valuable information for understanding diversification patterns in endangered areas. We analyzed the genetic diversity of Recordia reitzii, a tree restricted to the threatened and highly fragmented Brazilian Atlantic forest, using three intergenic cpDNA spacers and ten microsatellite (SSR) loci. To assess the historical processes that may have influenced the distribution of extant R. reitzii populations, the current potential distributions of R. reitzii and Recordia boliviana, a closely related species, were modelled and projected onto the Last Glacial Maximum (LGM) and Last Interglacial (LIG) periods. Niche divergence was quantified between these two. The cpDNA and SSR data showed a north–south pattern of the diversity distribution and structured populations, suggesting that gene flow is probably limited. According to our data, R. reitzii exhibits low genetic diversity, which may be a result of a founder or distribution‐reduction effect, narrow distribution or small population size. The ecological niche models showed a wider palaeodistribution during the LIG and a retraction during the LGM for both species. Tests of niche divergence and conservatism indicated that bioclimatic factors might have influenced the diversification of these Recordia species. © 2014 The Linnean Society of London, Botanical Journal of the Linnean Society, 2014, 176 , 332–348.     

Niche conservatism and phylogenetic clustering in a tribe of arid‐adapted marsupial mice,the Sminthopsini

The progressive expansion of the Australian arid zone during the last 20 Ma appears to have spurred the diversification of several families of plants, vertebrates and invertebrates, yet such taxonomic groups appear to show limited niche radiation. Here, we test whether speciation is associated with niche conservatism (constraints on ecological divergence) or niche divergence in a tribe of marsupial mice (Sminthopsini; 23 taxa) that includes the most speciose genus of living dasyurids, the sminthopsins. To that end, we integrated phylogenetic data with ecological niche modelling, to enable us to reconstruct the evolution of climatic suitability within Sminthopsini. Niche overlap among species was low‐moderate (but generally higher than expected given environmental background similarity), and the degree of phylogenetic clustering increased with aridity. Climatic niche reconstruction illustrates that there has been little apparent evolution of climatic tolerance within clades. Accordingly, climatic disparity tends to be accumulated among clades, suggesting considerable niche conservatism. Our results also indicate that evolution of climatic tolerances has been heterogeneous across different dimensions of climate (temperature vs. precipitation) and across phylogenetic clusters (Sminthopsis murina group vs. other groups). Although some results point to the existence of shifts in climatic niches during the speciation of sminthopsins, our study provides evidence for substantial phylogenetic niche conservatism in the group. We conclude that niche diversification had a low impact on the speciation of this tribe of small, but highly mobile marsupials.     

Patterns of speciation are similar across mountainous and lowland regions for a Neotropical plant radiation (Costaceae: Costus)

High species richness and endemism in tropical mountains are recognized as major contributors to the latitudinal diversity gradient. The processes underlying mountain speciation, however, are largely untested. The prevalence of steep ecogeographic gradients and the geographic isolation of populations by topographic features are predicted to promote speciation in mountains. We evaluate these processes in a species-rich Neotropical genus of understory herbs that range from the lowlands to montane forests and have higher species richness in topographically complex regions. We ask whether climatic niche divergence, geographic isolation, and pollination shifts differ between mountain-influenced and lowland Amazonian sister pairs inferred from a 756-gene phylogeny. Neotropical Costus ancestors diverged in Central America during a period of mountain formation in the last 3 million years with later colonization of Amazonia. Although climatic divergence, geographic isolation, and pollination shifts are prevalent in general, these factors do not differ between mountain-influenced and Amazonian sister pairs. Despite higher climatic niche and species diversity in the mountains, speciation modes in Costus appear similar across regions. Thus, greater species richness in tropical mountains may reflect differences in colonization history, diversification rates, or the prevalence of rapidly evolving plant life forms, rather than differences in speciation mode.     

Diversification of montane species via elevation shifts: the case of the Kaçkar cricket Phonochorion (Orthoptera)

Localized up‐down altitudinal shifts and subsequent isolation–admixture of montane species in response to glacial cycles has been proposed as a mechanism for the high diversity along Anatolian mountains. However, specific predictions of the proposed mechanism (the elevation shift model) have yet to be tested. Here, we provide a first assessment of this model for promoting inter‐ and intraspecific genetic diversity in the bush‐cricket genus Phonochorion endemic to the West Lesser Caucasus hotspot. Mitochondrial genes were analysed by Bayesian Markov Chain Monte Carlo inferences and coalescent simulations. Timing of diversification was estimated using a multispecies coalescent model. Divergence with gene flow was tested using an isolation with migration model. Population genetic parameters and genetic structuring were determined using Bayesian coalescent methods and spatial analysis. Demographic history was assessed using mismatch distributions and extended Bayesian skyline plots. Speciation events corresponded both to the Miocene and Pleistocene while intraspecific divergence was Pleistocene based. There was evidence for moderate levels of gene flow between species during diversification; however, incomplete lineage sorting could explain the data as well as gene flow. Overall diversification patterns within the genus Phonochorion agree with the predictions of the elevations shift model. Genetic patterns of diversification were driven mainly by Pleistocene glacial cycles and reflected the nature and distribution of sky islands. There was also some albeit weak evidence of demographic expansions coinciding with glacial cooling. However, evidence for divergence with gene flow was inconclusive.     

Evolutionary differentiation in the Neotropical montane region: molecular phylogenetics and phylogeography of Buarremon brush-finches (Aves, Emberizidae)

Studies on Neotropical phylogeography have largely focused on lowland organisms. Because lowland and highland biotas have different histories and are likely affected by different processes influencing population differentiation, understanding Neotropical diversification requires detailed studies on montane taxa. We present the most comprehensive analysis of population differentiation conducted so far on a widespread group of Neotropical montane organisms, focusing on the evolutionary relationships and phylogeography of Buarremon brush-finches (Aves: Emberizidae) in montane areas from Mexico through Argentina. Sequences of mitochondrial and nuclear genes demonstrate that Buarremon is not monophyletic with respect to Arremon and Lysurus. Genetic structure revealed by mtDNA is strong in both B. brunneinucha and B. torquatus. Gene genealogies and nucleotide diversity indicate that B. brunneinucha originated in Mexico and later expanded to South America, where it followed one colonization route through the east, and one through the west of the continent. Differentiation among populations of B. torquatus was substantial, reaching 8% uncorrected sequence divergence within South America. Relationships among major lineages of B. torquatus were not fully resolved owing to rapid differentiation, but the occurrence of closely related taxa in distant locations suggests a complex history of diversification. Some Colombian populations of B. brunneinucha have affinities with populations from Venezuela and the East Andean slope of Ecuador and Peru, and others with those from the Pacific slope of Ecuador. Moreover, five divergent lineages of B. torquatus occur within Colombia, highlighting the importance of dense sampling in northwest South America for studies on diversification of widespread Neotropical lineages.     

Tracing the origins of widespread highland species: a case of Neogene diversification across the Mexican sierras in an endemic lizard

The evolutionary history of the Mexican sierras has been shaped by various geological and climatic events over the past several million years. The relative impacts of these historical events on diversification in highland taxa, however, remain largely uncertain owing to a paucity of studies on broadly‐distributed montane species. We investigated the origins of genetic diversification in widely‐distributed endemic alligator lizards in the genus Barisia to help develop a better understanding of the complex processes structuring biological diversity in the Mexican highlands. We estimated lineage divergence dates and the diversification rate from mitochondrial DNA sequences, and combined divergence dates with reconstructions of ancestral geographical ranges to track lineage diversification across geography through time. Based on our results, we inferred ten geographically structured, well supported mitochondrial lineages within Barisia. Diversification of a widely‐distributed ancestor appears tied to the formation of the Trans‐Mexican Volcanic Belt across central Mexico during the Miocene and Pliocene. The formation of filter barriers such as major river drainages may have later subdivided lineages. The results of the present study provide additional support for the increasing number of studies that suggest Neogene events heavily impacted genetic diversification in widespread montane taxa. © 2011 The Linnean Society of London, Biological Journal of the Linnean Society, 2012, 105 , 382–394.     

Pliocene forest dynamics as a primary driver of African bird speciation

Aim Montane tropics are areas of high endemism, and mechanisms driving this endemism have been receiving increasing attention at a global scale. A general trend is that climatic factors do not explain the species richness of species with small to medium‐sized geographic ranges, suggesting that geological and evolutionary processes must be considered. On the African continent, several hypotheses including both refugial and geographic uplift models have been advanced to explain avian speciation and diversity in the lowland forest and montane regions of central and eastern Africa; montane regions in particular are recognized as hotspots of vertebrate endemism. Here, we examine the possible role of these models in driving speciation in a clade of African forest robins. Location Africa. Methods We constructed the first robustly supported molecular phylogenetic hypothesis of forest robins. On this phylogeny, we reconstructed habitat‐based distributions and geographic distributions relative to the Albertine Rift. We also estimated the timing of lineage divergences via a molecular clock. Results Robust estimates of phylogenetic relationships and clock‐based divergences reject Miocene tectonic uplift and Pleistocene forest refugia as primary drivers of speciation in forest robins. Instead, our data suggest that most forest robin speciation took place in the Late Pliocene, from 3.2 to 2.2 Ma. Distributional patterns are complex, with the Albertine Rift region serving as a general east–west break across the group. Montane distributions are inferred to have evolved four times. Main conclusions Phylogenetic divergence dates coincide with a single period of lowland forest retraction in the late Pliocene, suggesting that most montane speciation resulted from the rapid isolation of populations in montane areas, rather than montane areas themselves being drivers of speciation. This conclusion provides additional evidence that Pliocene climate change was a major driver of speciation in broadly distributed African animal lineages. We further show that lowland forest robins are no older than their montane relatives, suggesting that lowland areas are not museums which house ‘ancient’ taxa; rather, for forest robins, montane areas should be viewed as living museums of a late Pliocene diversification event. A forest refugial pattern is operating in Africa, but it is not constrained to the Pleistocene.     

Sky island diversification meets the multispecies coalescent – divergence in the spruce‐fir moss spider (Microhexura montivaga,Araneae, Mygalomorphae) on the highest peaks of southern Appalachia

Microhexura montivaga is a miniature tarantula‐like spider endemic to the highest peaks of the southern Appalachian mountains and is known only from six allopatric, highly disjunct montane populations. Because of severe declines in spruce‐fir forest in the late 20th century, M. montivaga was formally listed as a US federally endangered species in 1995. Using DNA sequence data from one mitochondrial and seven nuclear genes, patterns of multigenic genetic divergence were assessed for six montane populations. Independent mitochondrial and nuclear discovery analyses reveal obvious genetic fragmentation both within and among montane populations, with five to seven primary genetic lineages recovered. Multispecies coalescent validation analyses [guide tree and unguided Bayesian Phylogenetics and Phylogeography (BPP), Bayes factor delimitation (BFD)] using nuclear‐only data congruently recover six or seven distinct lineages; BFD analyses using combined nuclear plus mitochondrial data favour seven or eight lineages. In stark contrast to this clear genetic fragmentation, a survey of secondary sexual features for available males indicates morphological conservatism across montane populations. While it is certainly possible that morphologically cryptic speciation has occurred in this taxon, this system may alternatively represent a case where extreme population genetic structuring (but not speciation) leads to an oversplitting of lineage diversity by multispecies coalescent methods. Our results have clear conservation implications for this federally endangered taxon and illustrate a methodological issue expected to become more common as genomic‐scale data sets are gathered for taxa found in naturally fragmented habitats.     

Sky islands as foci for divergence of fig trees and their pollinators in southwest China

The dynamics of populations and their divergence over time have shaped current levels of biodiversity and in the case of the “sky islands” of mountainous southwest (SW) China have resulted in an area of exceptional botanical diversity. Ficus tikoua is a prostrate fig tree subendemic to the area that displays unique intraspecific diversity, producing figs typical of different pollination modes in different parts of its range. By combining climate models, genetic variation in populations of the tree's obligate fig wasp pollinators and distributions of the different plant phenotypes, we examined how this unusual situation may have developed. We identified three genetically distinct groups of a single Ceratosolen pollinator species that have largely parapatric distributions. The complex topography of the region contributed to genetic divergence among the pollinators by facilitating geographical isolation and providing refugia. Migration along elevations in response to climate oscillations further enhanced genetic differentiation of the three pollinator groups. Their distributions loosely correspond to the distributions of the functionally significant morphological differences in the male figs of their host plants, but postglacial expansion of one group has not been matched by spread of its associated plant phenotype, possibly due to a major river barrier. The results highlight how interplay between the complex topography of the “sky island” complex and climate change has shaped intraspecies differentiation and relationships between the plant and its pollinator. Similar processes may explain the exceptional botanical diversity of SW China.     

Investigating niche and lineage diversification in widely distributed taxa: phylogeography and ecological niche modeling of the Peromyscus maniculatus species group

The relationship between lineage formation and variation in the ecological niche is a fundamental evolutionary question. Two prevailing hypotheses reflect this relationship: niche conservatism and niche divergence. Niche conservatism predicts a pattern where sister taxa will occupy similar niche spaces; whereas niche divergence predicts that sister taxa will occupy different niche spaces. Widely distributed species often show distinct phylogeographic structure, but little research has been conducted on how the environment may be related to these phylogenetic patterns. We investigated the relationship between lineage divergence and environmental space for the closely related species Peromyscus maniculatus and P. polionotus utilizing phylogenetic techniques and ecological niche modeling (ENM). We estimated the phylogenetic relationship among individuals based on complete cytochrome b sequences that represent individuals from a majority of the species ranges. Niche spaces that lineages occupy were estimated by using 12 environmental layers. Differences in niche space were tested using multivariate statistics based on location data, and ENMs were employed using maximum entropy algorithms. Two similarity indices estimated significant divergence in environmental space based on the ENM. Six geographically structured lineages were identified within P. maniculatus. Nested within P. maniculatus we found that P. polionotus recently diverged from a clade occupying central and western United States. We estimated that the majority of the genetic lineages occupy distinct environmental niches, which supports a pattern of niche divergence. Two sister taxa showed niche divergence and represent different ecomorphs, suggesting morphological, genetic and ecological divergence between the two lineages. Two other sister taxa were observed in the same environmental space based on multivariate statistics, suggesting niche conservatism. Overall our results indicate that a widely distributed species may exhibit both niche conservatism and niche divergence, and that most lineages seem to occupy distinct environmental niches.     

Genomic insight into “sky island” species diversification in a mountainous biodiversity hotspot

“Sky island” species diversification contributes greatly to mountainous biodiversity. However, the underlying genomic divergence and the inferred drivers remain largely unknown. In this study, we examined the diversification history of five diploid species with three exclusively endemic to the sky islands (mountains) of the Himalaya–Hengduan Mountains biodiversity hotspot. All of them together comprise a clade of the genus Eutrema (Brassicaceae). We resequenced genomes of multiple individuals of the found populations for each species. We recovered the inconsistent phylogenetic relationships between plastome and nuclear‐genome trees for one species. Based on nuclear population genomic data, we detected high genetic divergence between five species with limited gene flow. Four species seemed to diverge mainly through geographical isolation, whereas one arose through hybrid origin. The origins of the sampled five species were dated to within the late Miocene when mountains were uplifted and climates oscillated. All species decreased their population sizes since the inferred origin of each species initially, but only two of them expanded after the Last Glacial Maximum. Together, these findings suggest that geographic isolation plays an important role in driving the sky island species diversification of the sampled species in addition to the occasional gene flow that might have led to the hybrid origin of some sky island species, similar to the species diversification of sea islands.     

From pampa to puna: Biogeography and diversification of a group of Neotropical obligate grassland birds (Anthus: Motacillidae)

The evolution of Neotropical birds of open landscapes remains largely unstudied. We investigate the diversification and biogeography of a group of Neotropical obligate grassland birds (Anthus: Motacillidae). We use a multilocus phylogeny of 22 taxa of Anthus to test the hypothesis that these birds radiated contemporaneously with the development of grasslands in South America. We employ the R package DDD to analyze the dynamics of Anthus diversification across time in Neotropical grasslands, explicitly testing for shifts in dynamics associated with the Miocene development of grasslands, the putative Pleistocene expansion of arid lowland biomes, and Pleistocene sundering of Andean highland grasslands. A lineage‐through‐time plot revealed increases in the number of lineages, and DDD detected shifts to a higher clade‐level carrying capacity during the late Miocene, indicating an early burst of diversification associated with grassland colonization. However, we could not corroborate the shift using power analysis, probably reflecting the small number of tips in our tree. We found evidence of a divergence at ~1 Mya between northern and southern Amazonian populations of Anthus lutescens, countering Haffer's idea of Pleistocene expansion of open biomes in the Amazon Basin. We used BioGeoBears to investigate ancestral areas and directionality of colonization of Neotropical grasslands. Members of the genus diversified into, out of, and within the Andes, within‐Andean diversification being mostly Pleistocene in origin.     

Climatic niche conservatism and ecological opportunity in the explosive radiation of arvicoline rodents (Arvicolinae,Cricetidae)

Climatic niche conservatism shapes patterns of diversity in many taxonomic groups, while ecological opportunity (EO) can trigger rapid speciation that is less constrained by the amount of time a lineage has occupied a given habitat. These two processes are well studied, but limited research has considered their joint and relative roles in shaping diversity patterns. We characterized climatic and biogeographic variables for 102 species of arvicoline rodents (Arvicolinae, Cricetidae), testing the effects of climatic niche conservatism and EO on arvicoline diversification as lineages transitioned between biogeographic regions. We found that the amount of time a lineage has occupied a precipitation niche is positively correlated with diversity along a precipitation gradient, suggesting climatic niche conservatism. In contrast, shift in diversification rate explained diversity patterns along a temperature gradient. Our results suggest that an indirect relationship exists between temperature and diversification that is associated with EO as arvicoline rodents colonized warm Palearctic environments. Climatic niche conservatism alone did not fully explain diversity patterns under density‐dependence, highlighting the additional importance of EO‐related processes in promoting the explosive radiation in arvicoline rodents and shaping diversity pattern among biogeographic regions and along climatic gradients.     

CLIMATE‐DRIVEN DIVERSIFICATION AND PLEISTOCENE REFUGIA IN PHILIPPINE BIRDS: EVIDENCE FROM PHYLOGEOGRAPHIC STRUCTURE AND PALEOENVIRONMENTAL NICHE MODELING

Avian diversification in oceanic archipelagos is largely attributed to isolation across marine barriers. During glacial maxima, lowered sea levels resulted in repeated land connections between islands joined by shallow seas. Consequently, such islands are not expected to show endemism. However, if climate fluctuations simultaneously caused shifts in suitable environmental conditions, limiting populations to refugia, then occurrence on and dispersal across periodic land bridges are not tenable. To assess the degree to which paleoclimate barriers, rather than marine barriers, drove avian diversification in the Philippine Archipelago, we produced ecological niche models for current‐day, glacial maxima, and interglacial climate scenarios to infer potential Pleistocene distributions and paleoclimate barriers. We then tested marine and paleoclimate barriers for correspondence to geographic patterns of population divergence, inferred from DNA sequences from eight codistributed bird species. In all species, deep‐water channels corresponded to zones of genetic differentiation, but six species exhibited deeper divergence associated with a periodic land bridge in the southern Philippines. Ecological niche models for these species identified a common paleoclimate barrier that coincided with deep genetic structure among populations. Although dry land connections joined southern Philippine islands during low sea level stands, unfavorable environmental conditions limited populations within landmasses, resulting in long‐term isolation and genetic differentiation. These results highlight the complex nature of diversification in archipelagos: marine barriers, changes in connectivity due to sea level change, and climate‐induced refugia acted in concert to produce great species diversity and endemism in the Philippines.