THE EVOLUTION OF ENVIRONMENTAL TOLERANCE AND RANGE SIZE: A COMPARISON OF GEOGRAPHICALLY RESTRICTED AND WIDESPREAD MIMULUS

The geographic ranges of closely related species can vary dramatically, yet we do not fully grasp the mechanisms underlying such variation. The niche breadth hypothesis posits that species that have evolved broad environmental tolerances can achieve larger geographic ranges than species with narrow environmental tolerances. In turn, plasticity and genetic variation in ecologically important traits and adaptation to environmentally variable areas can facilitate the evolution of broad environmental tolerance. We used five pairs of western North American monkeyflowers to experimentally test these ideas by quantifying performance across eight temperature regimes. In four species pairs, species with broader thermal tolerances had larger geographic ranges, supporting the niche breadth hypothesis. As predicted, species with broader thermal tolerances also had more within‐population genetic variation in thermal reaction norms and experienced greater thermal variation across their geographic ranges than species with narrow thermal tolerances. Species with narrow thermal tolerance may be particularly vulnerable to changing climatic conditions due to lack of plasticity and insufficient genetic variation to respond to novel selection pressures. Conversely, species experiencing high variation in temperature across their ranges may be buffered against extinction due to climatic changes because they have evolved tolerance to a broad range of temperatures.     

Distance,elevation and environment as drivers of diversity and divergence in bumble bees across latitude and altitude

Identifying drivers of dispersal limitation and genetic differentiation is a key goal in biogeography. We examine patterns of population connectivity and genetic diversity using restriction site‐associated DNA sequencing (RADseq) in two bumble bee species, Bombus vosnesenskii and Bombus bifarius, across latitude and altitude in mountain ranges from California, Oregon and Washington, U.S.A. Bombus vosnesenskii, which occurs across a broader elevational range at most latitudes, exhibits little population structure while B. bifarius, which occupies a relatively narrow higher elevation niche across most latitudes, exhibits much stronger population differentiation, although gene flow in both species is best explained by isolation with environmental niche resistance. A relationship between elevational habitat breadth and genetic diversity is also apparent, with B. vosnesenskii exhibiting relatively consistent levels of genetic diversity across its range, while B. bifarius has reduced genetic diversity at low latitudes, where it is restricted to high‐elevation habitat. The results of this study highlight the importance of the intersect between elevational range and habitat suitability in influencing population connectivity and suggest that future climate warming will have a fragmenting effect even on populations that are presently well connected, as they track their thermal niches upward in montane systems.     

Over the top: do thermal barriers along elevation gradients limit biotic similarity?

Organismal dispersal through mountain passes should be more constrained by temperature‐related differences between lowland and highland sites in montane environments. This may lead to higher rates of diversification through isolation of existing lineages toward the tropics. This mechanism, proposed by Janzen, could influence broad‐scale patterns of biodiversity across mountainous regions and more broadly across latitudinal gradients. We constructed two complementary analyses to test this hypothesis. First, we measured topographically‐derived thermal gradients using recently‐developed climatic data across the Americas, reviewing the main expectations from Janzen's climatic model. Then, we evaluated whether thermal barriers predict assemblage similarity for amphibians and mammals along elevational gradients across most of their latitudinal extent in the Americas. Thermal barriers between low and high elevation areas, initially proposed to be unique to tropical environments, are comparably strong in some temperate regions, particularly along the western slopes of North American dividing ranges. Biotic similarity for both mammals and amphibians decreases between sites that are separated by elevation‐related thermal barriers. That is, the stronger the thermal barrier separating pairs of sites across the latitudinal gradient, the lower the similarity of their species assemblages. Thermal barriers explain 10–35% of the variation in latitudinal gradients of biotic similarity, effects that were stronger in comparisons of sites at high elevations. Mammals' stronger dispersal capacities and homeothermy may explain weaker effects of thermal barriers on gradients of assemblage similarity than among amphibians. Understanding how temperature gradients have shaped gradients of montane biological diversity in the past will improve understanding of how changing environments may affect them in the future.     

Limits to elevational distributions in two species of emberizine finches: disentangling the role of interspecific competition, autoecology, and geographic variation in the environment

When species' elevational ranges are wider where putative competitors are absent, researchers have concluded that interspecific competition influences elevational distributions. This overlooks the distinction between factors that limit distributions directly and factors that only influence organisms indirectly through covarying regulators or resources. Because elevation affects organisms indirectly, testing whether competition influences elevational ranges relies on the heretofore untested assumption that the relationship between elevation and factors influencing organisms directly is similar across geography. Focusing on Buarremon brush-finches (Aves: Emberizidae), a group in which distributions represent one of the best examples of the potential role of competition limiting elevational ranges, we show that when distributions are compared along axes of climatic variation, some patterns of elevational range variation do appear to be consistent with predictions of the hypothesis that release from competition underlies expanded elevational ranges in allopatry. However, other patterns of expanded elevational ranges in the absence of putative competitors are better explained by hypothesis related to species' autoecology and geographic variation in the environment. This latter finding cautions against using elevation uncritically as a dimension of ecological niches, and suggests that classical examples of interspecific competition may need re-evaluation.     

Divergent selection along elevational gradients promotes genetic and phenotypic disparities among small mammal populations

Species distributed along mountain slopes, facing contrasting habitats in short geographic scale, are of particular interest to test how ecologically based divergent selection promotes phenotypic and genetic disparities as well as to assess isolation‐by‐environment mechanisms. Here, we conduct the first broad comparative study of phenotypic variation along elevational gradients, integrating a large array of ecological predictors and disentangling population genetic driver processes. The skull form of nine ecologically distinct species distributed over a large altitudinal range (100–4200 m) was compared to assess whether phenotypic divergence is a common phenomenon in small mammals and whether it shows parallel patterns. We also investigated the relative contribution of biotic (competition and predation) and abiotic parameters on phenotypic divergence via mixed models. Finally, we assessed the population genetic structure of a rodent species (Niviventer confucianus) via analysis of molecular variance and F_ST along three mountain slopes and tested the isolation‐by‐environment hypothesis using Mantel test and redundancy analysis. We found a consistent phenotypic divergence and marked genetic structure along elevational gradients; however, the species showed mixed patterns of size and skull shape trends across mountain zones. Individuals living at lower altitudes differed greatly in both phenotype and genotype from those living at high elevations, while middle‐elevation individuals showed more intermediate forms. The ecological parameters associated with phenotypic divergence along elevation gradients are partly related to species' ecological and evolutionary constraints. Fossorial and solitary animals are mainly affected by climatic factors, while terrestrial and more gregarious species are influenced by biotic and abiotic parameters. A novel finding of our study is that predator richness emerged as an important factor associated with the intraspecific diversification of the mammalian skull along elevational gradients, a previously overlooked parameter. Population genetic structure was mainly driven by environmental heterogeneity along mountain slopes, with no or a week spatial effect, fitting the isolation‐by‐environment scenario. Our study highlights the strong and multifaceted effects of heterogeneous steep habitats and ecologically based divergent selective forces in small mammal populations.     

Climatic zonation drives latitudinal variation in speciation mechanisms

Many groups of organisms show greater species richness in the tropics than in the temperate zone, particularly in tropical montane regions. Forty years ago, Janzen suggested that more limited temperature seasonality in the tropics leads to greater climatic zonation and more climatic barriers to organismal dispersal along elevational gradients in the tropics relative to temperate regions. These factors could lead to differences in how species arise in tropical versus temperate regions and possibly contribute to greater tropical diversity. However, no studies have compared the relationships among climate, elevational distribution and speciation in a group inhabiting both tropical and temperate regions. Here, we compare elevational and climatic divergence among 30 sister-species pairs (14 tropical, 16 temperate) within a single family of salamanders (Plethodontidae) that reaches its greatest species richness in montane Mesoamerica. In support of Janzen's hypothesis, we find that sister species are more elevationally and climatically divergent in the tropics than in the temperate zone. This pattern seemingly reflects regional variation in the role of climate in speciation, with niche conservatism predominating in the temperate zone and niche divergence in the tropics. Our study demonstrates how latitudinal differences in elevational climatic zonation may increase opportunities for geographical isolation, speciation and the associated build-up of species diversity in the tropics relative to the temperate zone.     

Plant invasion and speciation along elevational gradients on the oceanic island La Palma,Canary Islands

Ecosystems that provide environmental opportunities but are poor in species and functional richness generally support speciation as well as invasion processes. These processes are expected not to be equally effective along elevational gradients due to specific ecological, spatial, and anthropogenic filters, thus controlling the dispersal and establishment of species. Here, we investigate speciation and invasion processes along elevational gradients. We assess the vascular plant species richness as well as the number and percentage of endemic species and non‐native species systematically along three elevational gradients covering large parts of the climatic range of La Palma, Canary Islands. Species richness was negatively correlated with elevation, while the percentage of Canary endemic species showed a positive relationship. However, the percentage of Canary–Madeira endemics did not show a relationship with elevation. Non‐native species richness (indicating invasion) peaked at 500 m elevation and showed a consistent decline until about 1,200 m elevation. Above that limit, no non‐native species were present in the studied elevational gradients. Ecological, anthropogenic, and spatial filters control richness, diversification, and invasion with elevation. With increase in elevation, richness decreases due to species–area relationships. Ecological limitations of native ruderal species related to anthropogenic pressure are in line with the absence of non‐native species from high elevations indicating directional ecological filtering. Increase in ecological isolation with elevation drives diversification and thus increased percentages of Canary endemics. The best preserved eastern transect, including mature laurel forests, is an exception. The high percentage of Canary–Madeira endemics indicates the cloud forest's environmental uniqueness—and thus ecological isolation—beyond the Macaronesian islands.     

An alternative explanation for global trends in thermal tolerance

Ectotherms from higher latitudes can generally perform over broader temperature ranges than tropical ectotherms. This pattern is thought to reflect trends in temperature variability: tropical ectotherms evolve to be ‘thermal specialists’ because their environment is thermally stable. However, the tropics are also hotter, and most physiological rates increase exponentially with temperature. Using a dataset spanning diverse ectotherms, we show that the temperature ranges ectotherms tolerate (the difference between lower and upper critical temperatures, and between optimum and upper critical temperatures) generally represents the same range of equivalent biological rates (e.g. metabolism) for cool‐ and warm‐adapted species, and independent of latitude or elevation. This suggests that geographical trends in temperature variability may not be the ultimate mechanism underlying latitudinal and elevational trends in thermal tolerance. Rather, we propose that tropical ectotherms can perform over a narrower range of temperatures than species from higher latitudes because the tropics are hotter.     

Leaf trait associations with environmental variation in the wide‐ranging shrub Dodonaea viscosa subsp. angustissima (Sapindaceae)

Intra‐species variation in specific leaf area (SLA) and leaf area (LA) provides mechanistic insight into the persistence and function of plants, including their likely success under climate change and their suitability for revegetation. We measured SLA and LA in 101 Australian populations of the perennial shrub Dodonaea viscosa (L.) Jacq. subsp. angustissima (narrow‐leaf hop‐bush) (Sapindaceae). Populations were located across about a 1000 km north–south gradient, with climate grading from arid desert to mesic Mediterranean. We also measured leaves from 11 populations across an elevational gradient (300–800 m asl), where aridity and temperature decrease with elevation. We used regression and principal component analyses to relate leaf traits to the abiotic environment. SLA displayed clinal variation, increasing from north to south and correlated with latitude and the first principal component of joint environmental variables. Both SLA and LA correlated positively with most climatic and edaphic variables. Across latitude, LA showed more variability than SLA. Changes in leaf density and thickness may have caused the relative stability of SLA. Only LA decreased with elevation. The absence of a SLA response to elevation could be a consequence of abiotic conditions that favour low SLA at both ends of the elevational gradient. We demonstrated that the widely distributed narrow‐leaf hop‐bush shows considerable variability in LA and SLA, which allows it to persist in a broad environmental envelope. As this shrub is widely used for revegetation in Australia, South America and the Asia‐Pacific region, our results are consistent with the notion that seed used to revegetate mesic environments could be sourced from more arid areas to increase seed suitability to future climate change.     

Can Rapoport's rule explain tree species richness along the Himalayan elevation gradient,Nepal?

Rapoport's rule applied to an elevation gradient predicts a positive correlation between elevation ranges and elevation. This is supposed to be caused by the increasing magnitude of the climatic extremes at higher elevations, and thus, it is deduced that species richness should decrease with increasing elevation. The distribution of 614 tree species was used to test Rapoport's elevational rule along a gradient from 100 to 4300 m a.s.l., in the Nepalese Himalaya. The relationship between species richness and elevation was analysed by using generalized linear models (GLM). Generalized additive models (GAM) were used to examine the relationship between elevational range and the elevational mid-point of a species along the gradient. The widest elevation ranges are observed at mid-elevations, and narrow elevation ranges are observed at both ends of the gradient. This does not support Rapoport's elevation rule, as proposed by Stevens. There is a peak in species richness between 900 and 1000 m, and not in the tropical lowland as projected by Rapoport's elevation rule.     

Idiosyncratic responses to climate‐driven forest fragmentation and marine incursions in reed frogs from Central Africa and the Gulf of Guinea Islands

Organismal traits interact with environmental variation to mediate how species respond to shared landscapes. Thus, differences in traits related to dispersal ability or physiological tolerance may result in phylogeographic discordance among co‐distributed taxa, even when they are responding to common barriers. We quantified climatic suitability and stability, and phylogeographic divergence within three reed frog species complexes across the Guineo‐Congolian forests and Gulf of Guinea archipelago of Central Africa to investigate how they responded to a shared climatic and geological history. Our species‐specific estimates of climatic suitability through time are consistent with temporal and spatial heterogeneity in diversification among the species complexes, indicating that differences in ecological breadth may partly explain these idiosyncratic patterns. Likewise, we demonstrated that fluctuating sea levels periodically exposed a land bridge connecting Bioko Island with the mainland Guineo‐Congolian forest and that habitats across the exposed land bridge likely enabled dispersal in some species, but not in others. We did not find evidence that rivers are biogeographic barriers across any of the species complexes. Despite marked differences in the geographic extent of stable climates and temporal estimates of divergence among the species complexes, we recovered a shared pattern of intermittent climatic suitability with recent population connectivity and demographic expansion across the Congo Basin. This pattern supports the hypothesis that genetic exchange across the Congo Basin during humid periods, followed by vicariance during arid periods, has shaped regional diversity. Finally, we identified many distinct lineages among our focal taxa, some of which may reflect incipient or unrecognized species.     

Neither species geographic range size,climatic envelope,nor intraspecific leaf trait variability capture habitat specialization in a hyperdiverse Amazonian forest

Many plant species exhibit strong association with topographic habitats at local scales. However, the historical biogeographic and physiological drivers of habitat specialization are still poorly understood, and there is a need for relatively easy‐to‐measure predictors of species habitat niche breadth. Here, we explore whether species geographic range, climatic envelope, or intraspecific variability in leaf traits is related to the degree of habitat specialization in a hyperdiverse tropical tree community in Amazonian Ecuador. Contrary to our expectations, we find no effect of the size of species geographic ranges, the diversity of climate a species experiences across its range, or intraspecific variability in leaf traits in predicting topographic habitat association in the ~300 most common tropical tree species in a 25‐ha tropical forest plot. In addition, there was no phylogenetic signal to habitat specialization. We conclude that species geographic range size, climatic niche breadth, and intraspecific variability in leaf traits fail to capture the habitat specialization patterns observed in this highly diverse tropical forest.     

Physiological Limits along an Elevational Gradient in a Radiation of Montane Ground Beetles

A central challenge in ecology and biogeography is to determine the extent to which physiological constraints govern the geographic ranges of species along environmental gradients. This study tests the hypothesis that temperature and desiccation tolerance are associated with the elevational ranges of 12 ground beetle species (genus Nebria) occurring on Mt. Rainier, Washington, U.S.A. Species from higher elevations did not have greater cold tolerance limits than lower-elevation species (all species ranged from -3.5 to -4.1°C), despite a steep decline in minimum temperature with elevation. Although heat tolerance limits varied among species (from 32.0 to 37.0°C), this variation was not generally associated with the relative elevational range of a species. Temperature gradients and acute thermal tolerance do not support the hypothesis that physiological constraints drive species turnover with elevation. Measurements of intraspecific variation in thermal tolerance limits were not significant for individuals taken at different elevations on Mt. Rainier, or from other mountains in Washington and Oregon. Desiccation resistance was also not associated with a species’ elevational distribution. Our combined results contrast with previously-detected latitudinal gradients in acute physiological limits among insects and suggest that other processes such as chronic thermal stress or biotic interactions might be more important in constraining elevational distributions in this system.     

Linking species thermal tolerance to elevational range shifts in upland dung beetles

Climate warming has been proposed as the main cause of the recent range shifts seen in many species. Although species' thermal tolerances are thought to play a key role in determining responses to climate change, especially in ectotherms, empirical evidence is still limited. We investigate the connection between species' thermal tolerances, elevational range and shifts in the lower elevational limit of dung beetle species (Coleoptera, Aphodiidea) in an upland region in the northwest of England. We measured thermal tolerances in the laboratory, and used current and historical distribution data to test specific hypotheses about the area's three dominant species, particularly the species most likely to suffer from warming: Agollinus lapponum. We found marked differences between species in their minimum and maximum thermal tolerance and in their elevational range and patterns of abundance. Overall, differences in thermal limits among species matched the abundance patterns along the elevation gradient expected if distributions were constrained by climate. Agollinus lapponum abundance increased with elevation and this species showed lower maximum and minimum thermal limits than Acrossus depressus, for which abundance declined with elevation. Consistent with lower tolerance to high temperature, we recorded an uphill retreat of the low elevation limit of A. lapponum (177 m over 57 yr) in line with the increase in summer temperature observed in the region over the same period. Moreover, this species has been replaced at low and mid‐elevations by the other two warm‐tolerant species (A. depressus and Agrilinus ater). Our results provide empirical evidence that species' thermal tolerance constrains elevational ranges and contributes to explain the observed responses to climate warming. A mechanistic understanding of how climate change directly affects species, such as the one presented here, will provide a robust base to inform predictions of how individual species and whole assemblages may change in the future.     

Support for the elevational Rapoport's rule among seed plants in Nepal depends on biogeographical affinities and boundary effects

As one of the most important hypotheses on biogeographical distribution, Rapoport's rule has attracted attention around the world. However, it is unclear whether the applicability of the elevational Rapoport's Rule differs between organisms from different biogeographical regions. We used Stevens’ method, which uses species diversity and the averaged range sizes of all species within each (100 m) elevational band to explore diversity‐elevation, range‐elevation, and diversity‐range relationships. We compared support for the elevational Rapoport's rule between tropical and temperate species of seed plants in Nepal. Neither tropical nor temperate species supported the predictions of the elevational Rapoport's rule along the elevation gradient of 100–6,000 m a.s.l. for any of the studied relationships. However, along the smaller 1,000–5,000 m a.s.l. gradient (4,300 m a.s.l. for range‐elevation relationships) which is thought to be less influenced by boundary effects, we observed consistent support for the rule by tropical species, although temperate species did not show consistent support. The degree of support for the elevational Rapoport's rule may not only be influenced by hard boundary effects, but also by the biogeographical affinities of the focal taxa. With ongoing global warming and increasing variability of temperature in high‐elevation regions, tropical taxa may shift upward into higher elevations and expand their elevational ranges, causing the loss of temperate taxa diversity. Relevant studies on the elevational Rapoport's rule with regard to biogeographical affinities may be a promising avenue to further our understanding of this rule.     

Resources determine frugivore assemblages and fruit removal along an elevational gradient

Geographic variation in the diversity, abundance or composition of plant and frugivore assemblages may have consequences for seed dispersal processes. Such variations may be related to climatic conditions as well as habitat characteristics such as fruit availability and forest complexity. Studying frugivore assemblages and seed dispersal processes along an elevational gradient can help to elucidate the interplay between the extent of dispersal services provided by frugivores and the geographic variability of the food resources. We studied frugivore assemblages on and fruit removal from 28 rowan trees (Sorbus aucuparia) along an elevational gradient in the Bavarian Forest, Germany. Both, the number of frugivore species and the number of frugivore individuals were significantly enhanced by high fruit availability. In both cases we found a slight interaction between elevation and fruit availability indicating a higher attractiveness of fruits for frugivores at lower than at higher elevations. A high number of frugivore individuals in turn significantly increased fruit removal from rowan trees. Here, we found a significant interaction between elevation and the number of frugivore individuals suggesting that the number of frugivores is of major importance for fruit removal particularly at lower elevations. Path analysis corroborated that the number of frugivore individuals indirectly mediated the effect of fruit availability on fruit removal. These findings suggest that fruit removal is rather influenced by changes in habitat characteristics than in climatic conditions across space.     

Population genetic structure reveals terrestrial affinities for a headwater stream insect

1. The spatial distribution of stream‐dwelling organisms is often considered to be limited primarily according to the hierarchical structure of the hydrologic network, and previous conceptual models of population genetic structure have reflected this generality. Headwater specialists, however, are confined to short upstream sections of the network, and therefore are unlikely to respond in the same way as species with a broader range of habitat tolerance. 2. Here, we propose a model to describe spatial patterns of genetic diversity in headwater specialists with a limited ability for among‐stream dispersal. The headwater model predicts a partitioning of genetic variance according to higher‐elevation ‘islands’ of terrestrial habitat that provide required headwater stream conditions. The model therefore expects a geographic pattern of genetic variance similar to that expected for low‐dispersal terrestrial species occupying the adjacent habitat. 3. Using a 1032‐bp mitochondrial DNA fragment encompassing parts of the COI and COII genes, we demonstrate that Madrean Sky Islands populations of the giant water bug Abedus herberti conform to the proposed headwater model. Furthermore, they exhibit phylogeographic patterns broadly concordant with those shown for several terrestrial species in the region, including a major zone of discontinuity in the Chiricahua mountain range. 4. Overall, populations are highly isolated from one another, and a nested clade analysis suggested that A. herberti population structure, similarly to terrestrial Sky Islands species studied previously, has been influenced by Pleistocene climatic cycles causing expansion and contraction of temperate woodland habitat. 5. Because they have no ability to disperse among present‐day mountaintop habitat islands, A. herberti and other headwater species with limited dispersal ability are vulnerable to the projected increasing rate of climatic warming in this region.     

Identifying indicator species of elevation: Comparing the utility of woody plants,ants and moths for long‐term monitoring

Ecologists have found the distributions of many groups of organisms to be elevationally stratified. Consequently, various taxa (or species) have been proposed as indicators for inclusion within long‐term monitoring programmes to quantify the ecological impacts of future climatic change. Ideal indicators should be restricted to a particular elevational range (i.e. have high specificity) and be readily detectable across space and time (i.e. have high fidelity). This, however, has not been rigorously tested for elevational studies. We employed a spatially and temporally replicated sampling design to test the utility of tree, ant, and canopy and understorey moth species as indicators of elevation within continuous subtropical rainforest of eastern Australia. Using the classical indicator value protocol, we tested (i) whether the number of indicator species (all taxa) found in the observed data was significantly greater than the number obtained by chance; (ii) whether the indicator species (ants and moths) identified from one sampling season responded to elevation in a similar way in samples obtained from other seasons; and (iii) whether the indicator species (ants) identified from one elevational transect responded to elevation in a similar way in a nearby transect that incorporated similar elevational ranges. All groups had significantly greater numbers of indicator species than expected by chance. Temporal fidelity of moth indicator species was lower than that of ants as the suite of moth indicator species showed high seasonal variation. In contrast, ants showed high spatial and temporal fidelity. Most ant indicator species were, however, indicative of low and mid‐elevations, and only one species was indicative of the highest elevation, suggesting their relatively low conservation significance in relation to climate warming in this region. It is essential that we understand how spatial and temporal variation affects the distributions of different taxonomic groups when incorporating multiple taxa for long‐term monitoring programmes.     

Break zones in the distributions of alleles and species in alpine plants

Aim We test for the congruence between allele‐based range boundaries (break zones) in silicicolous alpine plants and species‐based break zones in the silicicolous flora of the European Alps. We also ask whether such break zones coincide with areas of large elevational variation. Location The European Alps. Methods On a regular grid laid across the entire Alps, we determined areas of allele‐ and species‐based break zones using respective clustering algorithms, identifying discontinuities in cluster distributions (breaks), and quantifying integrated break densities (break zones). Discontinuities were identified based on the intra‐specific genetic variation of 12 species and on the floristic distribution data from 239 species, respectively. Coincidence between the two types of break zones was tested using Spearman’s correlation. Break zone densities were also regressed on topographical complexity to test for the effect of elevational variation. Results We found that two main break zones in the distribution of alleles and species were significantly correlated. Furthermore, we show that these break zones are in topographically complex regions, characterized by massive elevational ranges owing to high mountains and deep glacial valleys. We detected a third break zone in the distribution of species in the eastern Alps, which is not correlated with topographic complexity, and which is also not evident from allelic distribution patterns. Species with the potential for long‐distance dispersal tended to show larger distribution ranges than short‐distance dispersers. Main conclusions We suggest that the history of Pleistocene glaciations is the main driver of the congruence between allele‐based and species‐based distribution patterns, because occurrences of both species and alleles were subject to the same processes (such as extinction, migration and drift) that shaped the distributions of species and genetic lineages. Large elevational ranges have had a profound effect as a dispersal barrier for alleles during post‐glacial immigration. Because plant species, unlike alleles, cannot spread via pollen but only via seed, and thus disperse less effectively, we conclude that species break zones are maintained over longer time spans and reflect more ancient patterns than allele break zones.     

Disentangling the historical routes to community assembly in the global epicentre of biodiversity

Aim

The exceptional turnover in biota with elevation and number of species coexisting at any elevation makes tropical mountains hotspots of biodiversity. However, understanding the historical processes through which species arising in geographical isolation (i.e. allopatry) assemble along the same mountain slope (i.e. sympatry) remains a major challenge. Multiple models have been proposed including (1) the sorting of already elevationally divergent species, (2) the displacement of elevation upon secondary contact, potentially followed by convergence, or (3) elevational conservatism, in which ancestral elevational ranges are retained. However, the relative contribution of these processes to generating patterns of elevational overlap and turnover is unknown.

Location

Tropical mountains of Central- and South-America.

Time Period

The last 12 myr.

Major Taxa Studied

Birds.

Methods

We collate a dataset of 165 avian sister pairs containing estimates of phylogenetic age, geographical and regional elevational range overlap. We develop a framework based on continuous-time Markov models to infer the relative frequency of different historical pathways in explaining present-day overlap and turnover of sympatric species along elevational gradients.

Results

We show that turnover of closely related bird species across elevation can predominantly be explained by displacement of elevation ranges upon contact (81%) rather than elevational divergence in allopatry (19%). In contrast, overlap along elevation gradients is primarily (88%) explained by conservatism of elevational ranges rather than displacement followed by elevational expansion (12%).

Main Conclusions

Bird communities across elevation gradients are assembled through a mix of processes, including the sorting, displacement and conservatism of species elevation ranges. The dominant role of conservatism in explaining co-occurrence of species on mountain slopes rejects more complex scenarios requiring displacement followed by expansion. The ability of closely related species to coexist without elevational divergence provides a direct and faster pathway to sympatry and helps explain the exceptional species richness of tropical mountains.