Climate change and elevational range shifts in insects.

On mountains, unique in their steep and rapid climatic gradients, many insects are shifting their elevational range limits to track recent temperature change. In a review of the range shift literature to date, most of the 1478 montane insect populations tested so far are shifting to higher elevations, but there is conspicuous variation in the responses. We discuss the impact of study methodology as well as potential abiotic and biotic factors that may underlie this variation in climate change response. We encourage more empirical studies spanning greater insect biodiversity and directly testing how variation in species' traits, biogeography, and abiotic-biotic context shapes variation in range shift responses.

Unusually large upward shifts in cold-adapted, montane mammals as temperature warms.

The largest and tallest mountain range in the contiguous United States, the Southern Rocky Mountains, has warmed considerably in the past several decades due to anthropogenic climate change. Herein we examine how 47 mammal elevational ranges (27 rodent and 4 shrew species) have changed from their historical distributions (1886-1979) to their contemporary distributions (post 2005) along 2,400-m elevational gradients in the Front Range and San Juan Mountains of Colorado. Historical elevational ranges were based on more than 4,580 georeferenced museum specimen and publication records. Contemporary elevational ranges were based on 7,444 records from systematic sampling efforts and museum specimen records. We constructed Bayesian models to estimate the probability a species was present, but undetected, due to undersampling at each 50-m elevational bin for each time period and mountain range. These models leveraged individual-level detection probabilities, the number and patchiness of detections across 50-m bands of elevation, and a decaying likelihood of presence from last known detections. We compared 95% likelihood elevational ranges between historical and contemporary time periods to detect directional change. Responses were variable as 26 mammal ranges changed upward, 6 did not change, 11 changed downward, and 4 were extirpated locally. The average range shift was 131 m upward, while exclusively montane species shifted upward more often (75%) and displayed larger average range shifts (346 m). The best predictors of upper limit and total directional change were species with higher maximum latitude in their geographic range, montane affiliation, and the study mountain was at the southern edge of their geographic range. Thus, mammals in the Southern Rocky Mountains serve as harbingers of more changes to come, particularly for montane, cold-adapted species in the southern portion of their ranges.

Natural population variability may be masking the more-individuals hypothesis.

Species richness and productivity are correlated at global and regional scales, but the mechanisms linking them are inconclusive. The most commonly invoked mechanism, the more-individuals hypothesis (MIH), hypothesizes that increased productivity leads to increased food resource availability, which leads to an increased number of individuals supporting more species. Empirical evidence for the MIH remains mixed despite a substantial literature. Here we used simulations to determine whether interannual population variability could be masking a "true" MIH relationship. In each simulation, fixed linear relationships between productivity, richness, and 50-yr average abundance mimicked the MIH mechanism. Abundance was allowed to vary annually and sampled for 1-40 yr. Linear regressions of richness on sampled abundance assessed the probability of detecting the fixed MIH relationship. Medium to high population variability with short-term sampling (1-3 yr) led to poor detection of the fixed MIH relationship. Notably, this level of sampling and population variability describes nearly all MIH studies to date. Long-term sampling (5+ yr) led to improved detection of the fixed relationship; thus it is necessary to detect support for the MIH reliably. Such sampling duration is nonexistent in the MIH literature. Robust future studies of the MIH necessitate consideration of interannual population variability.

A taxonomically broad metagenomic survey of 339 species spanning 57 families suggests cystobasidiomycete yeasts are not ubiquitous across all lichens.

PREMISE: Lichens are fungi that enter into obligate symbioses with photosynthesizing organisms (algae, cyanobacteria). Traditional narratives of lichens as binary symbiont pairs have given way to their recognition as dynamic metacommunities. Basidiomycete yeasts, particularly of the genus Cyphobasidium, have been inferred to be widespread and important components of lichen metacommunities. Yet, the presence of basidiomycete yeasts across a wide diversity of lichen lineages has not previously been tested. METHODS: We searched for lichen-associated cystobasidiomycete yeasts in newly generated metagenomic data from 413 samples of 339 lichen species spanning 57 families and 25 orders. The data set was generated as part of a large-scale project to study lichen biodiversity gradients in the southern Appalachian Mountains Biodiversity Hotspot of southeastern North America. RESULTS: Our efforts detected cystobasidiomycete yeasts in nine taxa (Bryoria nadvornikiana, Heterodermia leucomelos, Lecidea roseotincta, Opegrapha vulgata, Parmotrema hypotropum, P. subsumptum, Usnea cornuta, U. strigosa, and U. subgracilis), representing 2.7% of all species sampled. Seven of these taxa (78%) are foliose (leaf-like) or fruticose (shrubby) lichens that belong to families where basidiomycete yeasts have been previously detected. In several of the nine cases, cystobasidiomycete rDNA coverage was comparable to, or greater than, that of the primary lichen fungus single-copy nuclear genomic rDNA, suggesting sampling artifacts are unlikely to account for our results. CONCLUSIONS: Studies from diverse areas of the natural sciences have led to the need to reconceptualize lichens as dynamic metacommunities. However, our failure to detect cystobasidiomycetes in 97.3% (330 species) of the sampled species suggests that basidiomycete yeasts are not ubiquitous in lichens.

Habitat quality and disturbance drive lichen species richness in a temperate biodiversity hotspot.

The impacts of disturbance on biodiversity and distributions have been studied in many systems. Yet, comparatively less is known about how lichens-obligate symbiotic organisms-respond to disturbance. Successful establishment and development of lichens require a minimum of two compatible yet usually unrelated species to be present in an environment, suggesting disturbance might be particularly detrimental. To address this gap, we focused on lichens, which are obligate symbiotic organisms that function as hubs of trophic interactions. Our investigation was conducted in the southern Appalachian Mountains, USA. We conducted complete biodiversity inventories of lichens (all growth forms, reproductive modes, substrates) across 47, 1-ha plots to test classic models of responses to disturbance (e.g., linear, unimodal). Disturbance was quantified in each plot using a standardized suite of habitat quality variables. We additionally quantified woody plant diversity, forest density, rock density, as well as environmental factors (elevation, temperature, precipitation, net primary productivity, slope, aspect) and analyzed their impacts on lichen biodiversity. Our analyses recovered a strong, positive, linear relationship between lichen biodiversity and habitat quality: lower levels of disturbance correlate to higher species diversity. With few exceptions, additional variables failed to significantly explain variation in diversity among plots for the 509 total lichen species, but we caution that total variation in some of these variables was limited in our study area. Strong, detrimental impacts of disturbance on lichen biodiversity raises concerns about conservation and land management practices that fail to incorporate complete estimates of biodiversity, especially from ecologically important organisms such as lichens.

Midpoint attractors and species richness: Modelling the interaction between environmental drivers and geometric constraints.

We introduce a novel framework for conceptualising, quantifying and unifying discordant patterns of species richness along geographical gradients. While not itself explicitly mechanistic, this approach offers a path towards understanding mechanisms. In this study, we focused on the diverse patterns of species richness on mountainsides. We conjectured that elevational range midpoints of species may be drawn towards a single midpoint attractor - a unimodal gradient of environmental favourability. The midpoint attractor interacts with geometric constraints imposed by sea level and the mountaintop to produce taxon-specific patterns of species richness. We developed a Bayesian simulation model to estimate the location and strength of the midpoint attractor from species occurrence data sampled along mountainsides. We also constructed midpoint predictor models to test whether environmental variables could directly account for the observed patterns of species range midpoints. We challenged these models with 16 elevational data sets, comprising 4500 species of insects, vertebrates and plants. The midpoint predictor models generally failed to predict the pattern of species midpoints. In contrast, the midpoint attractor model closely reproduced empirical spatial patterns of species richness and range midpoints. Gradients of environmental favourability, subject to geometric constraints, may parsimoniously account for elevational and other patterns of species richness.

A Systematic Review of Global Drivers of Ant Elevational Diversity.

Ant diversity shows a variety of patterns across elevational gradients, though the patterns and drivers have not been evaluated comprehensively. In this systematic review and reanalysis, we use published data on ant elevational diversity to detail the observed patterns and to test the predictions and interactions of four major diversity hypotheses: thermal energy, the mid-domain effect, area, and the elevational climate model. Of sixty-seven published datasets from the literature, only those with standardized, comprehensive sampling were used. Datasets included both local and regional ant diversity and spanned 80° in latitude across six biogeographical provinces. We used a combination of simulations, linear regressions, and non-parametric statistics to test multiple quantitative predictions of each hypothesis. We used an environmentally and geometrically constrained model as well as multiple regression to test their interactions. Ant diversity showed three distinct patterns across elevations: most common were hump-shaped mid-elevation peaks in diversity, followed by low-elevation plateaus and monotonic decreases in the number of ant species. The elevational climate model, which proposes that temperature and precipitation jointly drive diversity, and area were partially supported as independent drivers. Thermal energy and the mid-domain effect were not supported as primary drivers of ant diversity globally. The interaction models supported the influence of multiple drivers, though not a consistent set. In contrast to many vertebrate taxa, global ant elevational diversity patterns appear more complex, with the best environmental model contingent on precipitation levels. Differences in ecology and natural history among taxa may be crucial to the processes influencing broad-scale diversity patterns.

Body size and activity times mediate mammalian responses to climate change.

Model predictions of extinction risks from anthropogenic climate change are dire, but still overly simplistic. To reliably predict at-risk species we need to know which species are currently responding, which are not, and what traits are mediating the responses. For mammals, we have yet to identify overarching physiological, behavioral, or biogeographic traits determining species' responses to climate change, but they must exist. To date, 73 mammal species in North America and eight additional species worldwide have been assessed for responses to climate change, including local extirpations, range contractions and shifts, decreased abundance, phenological shifts, morphological or genetic changes. Only 52% of those species have responded as expected, 7% responded opposite to expectations, and the remaining 41% have not responded. Which mammals are and are not responding to climate change is mediated predominantly by body size and activity times (phylogenetic multivariate logistic regressions, P < 0.0001). Large mammals respond more, for example, an elk is 27 times more likely to respond to climate change than a shrew. Obligate diurnal and nocturnal mammals are more than twice as likely to respond as mammals with flexible activity times (P < 0.0001). Among the other traits examined, species with higher latitudinal and elevational ranges were more likely to respond to climate change in some analyses, whereas hibernation, heterothermy, burrowing, nesting, and study location did not influence responses. These results indicate that some mammal species can behaviorally escape climate change whereas others cannot, analogous to paleontology's climate sheltering hypothesis. Including body size and activity flexibility traits into future extinction risk forecasts should substantially improve their predictive utility for conservation and management.

Environmental harshness is positively correlated with intraspecific divergence in mammals and birds.

Life on Earth is conspicuously more diverse in the tropics. Although this intriguing geographical pattern has been linked to many biotic and abiotic factors, their relative importance and potential interactions are still poorly understood. The way in which latitudinal changes in ecological conditions influence evolutionary processes is particularly controversial, as there is evidence for both a positive and a negative latitudinal gradient in speciation rates. Here, we identify and address some methodological issues (how patterns are analysed and how latitude is quantified) that could lead to such conflicting results. To address these issues, we assemble a comprehensive data set of the environmental correlates of latitude (including climate, net primary productivity and habitat heterogeneity) and combine it with biological, historical and molecular data to explore global patterns in recent divergence events (subspeciation). Surprisingly, we find that the harsher conditions that typify temperate habitats (lower primary productivity, decreased rainfall and more variable and unpredictable temperatures) are positively correlated with greater subspecies richness in terrestrial mammals and birds. Thus, our findings indicate that intraspecific divergence is greater in regions with lower biodiversity, a pattern that is robust to both sampling variation and latitudinal biases in taxonomic knowledge. We discuss possible causal mechanisms for the link between environmental harshness and subspecies richness (faster rates of evolution, greater likelihood of range discontinuities and more opportunities for divergence) and conclude that this pattern supports recent indications that latitudinal gradients of diversity are maintained by simultaneously higher potentials for both speciation and extinction in temperate than tropical regions.

Latitude, elevational climatic zonation and speciation in New World vertebrates.

Many biodiversity hotspots are located in montane regions, especially in the tropics. A possible explanation for this pattern is that the narrow thermal tolerances of tropical species and greater climatic stratification of tropical mountains create more opportunities for climate-associated parapatric or allopatric speciation in the tropics relative to the temperate zone. However, it is unclear whether a general relationship exists among latitude, climatic zonation and the ecology of speciation. Recent taxon-specific studies obtained different results regarding the role of climate in speciation in tropical versus temperate areas. Here, we quantify overlap in the climatic distributions of 93 pairs of sister species of mammals, birds, amphibians and reptiles restricted to either the New World tropics or to the Northern temperate zone. We show that elevational ranges of tropical- and temperate-zone species do not differ from one another, yet the temperature range experienced by species in the temperate zone is greater than for those in the tropics. Moreover, tropical sister species tend to exhibit greater similarity in their climatic distributions than temperate sister species. This pattern suggests that evolutionary conservatism in the thermal niches of tropical taxa, coupled with the greater thermal zonation of tropical mountains, may result in increased opportunities for allopatric isolation, speciation and the accumulation of species in tropical montane regions. Our study exemplifies the power of combining phylogenetic and spatial datasets of global climatic variation to explore evolutionary (rather than purely ecological) explanations for the high biodiversity of tropical montane regions.

Assessing the threat to montane biodiversity from discordant shifts in temperature and precipitation in a changing climate.

Mountains are centres of global biodiversity, endemism and threatened species. Elevational gradients present opportunities for species currently living near their upper thermal limits to track cooler temperatures upslope in warming climates, but only if changes in precipitation are sufficiently in step with temperature. We model local population extirpation risk for a range of temperature and precipitation scenarios over the next 100 years for 16 848 vertebrate species populations distributed along 156 elevational gradients. Average population extirpation risks due to warming alone were < 5%, but increased 10-fold, on average, when changes in precipitation were also considered. Under the driest scenarios (minimum predicted precipitation), local extirpation risks increased sharply (50-60%) and were especially worrisome for hydrophilic amphibians and montane Latin America (c. 80%). Realistic assessment of risks urgently requires improved monitoring of precipitation, better regional precipitation models and more research on the effects of changes in precipitation on montane distributions.

Microbes do not follow the elevational diversity patterns of plants and animals.

The elevational gradient in plant and animal diversity is one of the most widely documented patterns in ecology and, although no consensus explanation exists, many hypotheses have been proposed over the past century to explain these patterns. Historically, research on elevational diversity gradients has focused almost exclusively on plant and animal taxa. As a result, we do not know whether microbes exhibit elevational gradients in diversity that parallel those observed for macroscopic taxa. This represents a key knowledge gap in ecology, especially given the ubiquity, abundance, and functional importance of microbes. Here we show that, across a montane elevational gradient in eastern Peru, bacteria living in three distinct habitats (organic soil, mineral soil, and leaf surfaces) exhibit no significant elevational gradient in diversity (r2<0.17, P>0.1 in all cases), in direct contrast to the significant diversity changes observed for plant and animal taxa across the same montane gradient (r2>0.75, P<0.001 in all cases). This finding suggests that the biogeographical patterns exhibited by bacteria are fundamentally different from those of plants and animals, highlighting the need for the development of more inclusive concepts and theories in biogeography to explain these disparities.

Niche conservatism as an emerging principle in ecology and conservation biology.

The diversity of life is ultimately generated by evolution, and much attention has focused on the rapid evolution of ecological traits. Yet, the tendency for many ecological traits to instead remain similar over time [niche conservatism (NC)] has many consequences for the fundamental patterns and processes studied in ecology and conservation biology. Here, we describe the mounting evidence for the importance of NC to major topics in ecology (e.g. species richness, ecosystem function) and conservation (e.g. climate change, invasive species). We also review other areas where it may be important but has generally been overlooked, in both ecology (e.g. food webs, disease ecology, mutualistic interactions) and conservation (e.g. habitat modification). We summarize methods for testing for NC, and suggest that a commonly used and advocated method (involving a test for phylogenetic signal) is potentially problematic, and describe alternative approaches. We suggest that considering NC: (1) focuses attention on the within-species processes that cause traits to be conserved over time, (2) emphasizes connections between questions and research areas that are not obviously related (e.g. invasives, global warming, tropical richness), and (3) suggests new areas for research (e.g. why are some clades largely nocturnal? why do related species share diseases?).

Metabolic theory and elevational diversity of vertebrate ectotherms.

The Metabolic Theory of Ecology (MTE) posits that the temperature-dependent kinetics of metabolism shape broad-scale patterns of biodiversity. Here we test whether the MTE accounts for patterns of diversity using 102 elevational diversity gradients of reptiles and amphibians. In particular, we examined the support for the two key predictions of the MTE: that the reciprocal of absolute temperature (1/kT) and diversity are linearly related and that the slope of that relationship is -0.65. We also tested two underlying assumptions of the MTE in cases with appropriate data, namely, that abundance is invariant among samples, and that behavioral thermoregulation influences the MTE predictions. We found that few studies supported the predictions of the MTE for the relationship between environmental temperature and elevational diversity using previous methods on individual gradients and using meta-analysis. The predominant relationship was curvilinear, and the slopes were steeper than predicted. In analyses of individual gradients, only 6% followed the MTE predictions in the strictest application, and 25% in the broadest. We found violations of the assumption of invariant abundances in all five test cases. All four herpetofaunal groups, regardless of behavioral thermoregulatory abilities, demonstrated poor fits to the MTE predictions. Even when arid gradients are removed, ameliorating the potential effects of water limitation, the MTE did not account for herpetofaunal elevational diversity. We conclude that an interplay of factors shapes elevational diversity gradients rather than the simple kinetics of biochemical reactions.

Phylogeny, niche conservatism and the latitudinal diversity gradient in mammals.

Biologists have long searched for mechanisms responsible for the increase in species richness with decreasing latitude. The strong correlation between species richness and climate is frequently interpreted as reflecting a causal link via processes linked to energy or evolutionary rates. Here, we investigate how the aggregation of clades, as dictated by phylogeny, can give rise to significant climate-richness gradients without gradients in diversification or environmental carrying capacity. The relationship between climate and species richness varies considerably between clades, regions and time periods in a global-scale phylogenetically informed analysis of all terrestrial mammal species. Many young clades show negative richness-temperature slopes (more species at cooler temperatures), with the ages of these clades coinciding with the expansion of temperate climate zones in the late Eocene. In carnivores, we find steeply positive richness-temperature slopes in clades with restricted distributions and tropical origins (e.g. cat clade), whereas widespread, temperate clades exhibit shallow, negative slopes (e.g. dog-bear clade). We show that the slope of the global climate-richness gradient in mammals is driven by aggregating Chiroptera (bats) with their Eutherian sister group. Our findings indicate that the evolutionary history should be accounted for as part of any search for causal links between environment and species richness.

Patterns and causes of species richness: a general simulation model for macroecology.

Understanding the causes of spatial variation in species richness is a major research focus of biogeography and macroecology. Gridded environmental data and species richness maps have been used in increasingly sophisticated curve-fitting analyses, but these methods have not brought us much closer to a mechanistic understanding of the patterns. During the past two decades, macroecologists have successfully addressed technical problems posed by spatial autocorrelation, intercorrelation of predictor variables and non-linearity. However, curve-fitting approaches are problematic because most theoretical models in macroecology do not make quantitative predictions, and they do not incorporate interactions among multiple forces. As an alternative, we propose a mechanistic modelling approach. We describe computer simulation models of the stochastic origin, spread, and extinction of species' geographical ranges in an environmentally heterogeneous, gridded domain and describe progress to date regarding their implementation. The output from such a general simulation model (GSM) would, at a minimum, consist of the simulated distribution of species ranges on a map, yielding the predicted number of species in each grid cell of the domain. In contrast to curve-fitting analysis, simulation modelling explicitly incorporates the processes believed to be affecting the geographical ranges of species and generates a number of quantitative predictions that can be compared to empirical patterns. We describe three of the 'control knobs' for a GSM that specify simple rules for dispersal, evolutionary origins and environmental gradients. Binary combinations of different knob settings correspond to eight distinct simulation models, five of which are already represented in the literature of macroecology. The output from such a GSM will include the predicted species richness per grid cell, the range size frequency distribution, the simulated phylogeny and simulated geographical ranges of the component species, all of which can be compared to empirical patterns. Challenges to the development of the GSM include the measurement of goodness of fit (GOF) between observed data and model predictions, as well as the estimation, optimization and interpretation of the model parameters. The simulation approach offers new insights into the origin and maintenance of species richness patterns, and may provide a common framework for investigating the effects of contemporary climate, evolutionary history and geometric constraints on global biodiversity gradients. With further development, the GSM has the potential to provide a conceptual bridge between macroecology and historical biogeography.

Vertebrate range sizes indicate that mountains may be 'higher' in the tropics.

In 1967, Daniel Janzen proposed the influential, but largely untested hypothesis, that tropical mountain passes are physiologically higher than temperate mountains. I test his key prediction, the one upon which all the others rely: namely, that elevational range sizes of organisms get larger on mountains at increasing latitudes. My analyses use 170 montane gradients spanning 36.5 degrees S to 48.2 degrees N latitude compiled from over 80 years of research and 16,500 species of rodents, bats, birds, lizards, snakes, salamanders, and frogs. In support of Janzen's prediction, I find that elevational range size increases with increasing latitude for all vertebrate groups except rodents. I document additional lines of evidence for temperature variability as a plausible mechanism for trends in vertebrate range size, including strong effects of thermoregulation and daily temperature variability, and a weak effect of precipitation.

Area and mammalian elevational diversity.

Elevational gradients hold enormous potential for understanding general properties of biodiversity. Like latitudinal gradients, the hypotheses for diversity patterns can be grouped into historical explanations, climatic drivers, and spatial hypotheses. The spatial hypotheses include the species-area effect and spatial constraint (mid-domain effect null models). I test these two spatial hypotheses using regional diversity patterns for mammals (non-volant small mammals and bats) along 34 elevational gradients spanning 24.4 degrees S-40.4 degrees N latitude. There was high variability in the fit to the species-area hypothesis and the mid-domain effect. Both hypotheses can be eliminated as primary drivers of elevational diversity. Area and spatial constraint both represent sources of error rather than mechanisms underlying these mammalian diversity patterns. Similar results are expected for other vertebrate taxa, plants, and invertebrates since they show comparable distributions of elevational diversity patterns to mammalian patterns.

Evolution and the latitudinal diversity gradient: speciation, extinction and biogeography.

A latitudinal gradient in biodiversity has existed since before the time of the dinosaurs, yet how and why this gradient arose remains unresolved. Here we review two major hypotheses for the origin of the latitudinal diversity gradient. The time and area hypothesis holds that tropical climates are older and historically larger, allowing more opportunity for diversification. This hypothesis is supported by observations that temperate taxa are often younger than, and nested within, tropical taxa, and that diversity is positively correlated with the age and area of geographical regions. The diversification rate hypothesis holds that tropical regions diversify faster due to higher rates of speciation (caused by increased opportunities for the evolution of reproductive isolation, or faster molecular evolution, or the increased importance of biotic interactions), or due to lower extinction rates. There is phylogenetic evidence for higher rates of diversification in tropical clades, and palaeontological data demonstrate higher rates of origination for tropical taxa, but mixed evidence for latitudinal differences in extinction rates. Studies of latitudinal variation in incipient speciation also suggest faster speciation in the tropics. Distinguishing the roles of history, speciation and extinction in the origin of the latitudinal gradient represents a major challenge to future research.