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. K E Y W O R D S :Climatic variability hypothesis, geographic range size, genetic variation, niche breadth, specialist-generalist tradeoffs, thermal performance curve.

Summary Geographic range size has long fascinated ecologists and evolutionary biologists, yet our understanding of the factors that cause variation in range size among species and across space remains limited. Not only does geographic range size inform decisions about the conservation and management of rare and nonindigenous species due to its relationship with extinction risk, rarity, and invasiveness, but it also provides insights into fundamental processes such as dispersal and adaptation. There are several features unique to plants (e.g. polyploidy, mating system, sessile habit) that may lead to distinct mechanisms explaining variation in range size. Here, we highlight key studies testing intrinsic and extrinsic hypotheses about geographic range size under contrasting scenarios where species' ranges are static or change over time. We then present results from a meta‐analysis of the relative importance of commonly hypothesized determinants of range size in plants. We show that our ability to infer the relative importance of these determinants is limited, particularly for dispersal ability, mating system, ploidy, and environmental heterogeneity. We highlight avenues for future research that merge approaches from macroecology and evolutionary ecology to better understand how adaptation and dispersal interact to facilitate niche evolution and range expansion.

Determining how species' geographic ranges are governed by current climates and how they will respond to rapid climatic change poses a major biological challenge. Geographic ranges are often spatially fragmented and composed of genetically differentiated populations that are locally adapted to different thermal regimes. Tradeoffs between different aspects of thermal performance, such as between tolerance to high temperature and tolerance to low temperature or between maximal performance and breadth of performance, suggest that the performance of a given population will be a subset of that of the species. Therefore, species-level projections of distribution might overestimate the species' ability to persist at any given location. However, current approaches to modeling distributions often do not consider variation among populations. Here, we estimated genetically-based differences in thermal performance curves for growth among 12 populations of the scarlet monkeyflower, Mimulus cardinalis, a perennial herb of western North America. We inferred the maximum relative growth rate (RGR(max)), temperature optimum (T(opt)), and temperature breadth (T(breadth)) for each population. We used these data to test for tradeoffs in thermal performance, generate mechanistic population-level projections of distribution under current and future climates, and examine how variation in aspects of thermal performance influences forecasts of range shifts. Populations differed significantly in RGR(max) and had variable, but overlapping, estimates of T(opt) and T(breadth). T(opt) declined with latitude and increased with temperature of origin, consistent with tradeoffs between performances at low temperatures versus those at high temperatures. Further, T(breadth) was negatively related to RGR(max), as expected for a specialist-generalist tradeoff. Parameters of the thermal performance curve influenced properties of projected distributions. For both current and future climates, T(opt) was negatively related to latitudinal position, while T(breadth) was positively related to projected range size. The magnitude and direction of range shifts also varied with T(opt) and T(breadth), but sometimes in unexpected ways. For example, the fraction of habitat remaining suitable increased with T(opt) but decreased with T(breadth). Northern limits of all populations were projected to shift north, but the magnitude of shift decreased with T(opt) and increased with T(breadth). Median latitude was projected to shift north for populations with high T(breadth) and low T(opt), but south for populations with low T(breadth) and high T(opt). Distributions inferred by integrating population-level projections did not differ from a species-level projection that ignored variation among populations. However, the species-level approach masked the potential array of divergent responses by populations that might lead to genotypic sorting within the species' range. Thermal performance tradeoffs among populations within the species' range had important, but sometimes coun...

Species distribution models (SDMs) are widely used in ecology. In theory, SDMs capture (at least part of ) species' ecological niches and can be used to make inferences about the distribution of suitable habitat for species of interest. Because habitat suitability is expected to influence population demography, SDMs have been used to estimate a variety of population parameters, from occurrence to genetic diversity. However, a critical look at the ability of SDMs to predict independent data across different aspects of population biology is lacking. Here, we systematically reviewed the literature, retrieving 201 studies that tested predictions from SDMs against independent assessments of occurrence, abundance, population performance, and genetic diversity. Although there is some support for the ability of SDMs to predict occurrence (~53% of studies depending on how support was assessed), the predictive performance of these models declines progressively from occurrence to abundance, to population mean fitness, to genetic diversity. At the same time, we observed higher success among studies that evaluated performance for single versus multiple species, pointing to a possible publication bias. Thus, the limited accuracy of SDMs reported here may reflect the best-case scenario. We discuss the limitations of these models and provide specific recommendations for their use for different applications going forward. However, we emphasize that predictions from SDMs, especially when used to inform conservation decisions, should be treated as hypotheses to be tested with independent data rather than as stand-ins for the population parameters we seek to know.

As climate change shifts species' climatic envelopes across the landscape, equilibrium between geographic ranges and niches is likely diminishing due to time lags in demography and dispersal. If a species' range and niche are out of equilibrium, then population performance should decrease from cool, "leading" range edges, where populations are expanding into recently ameliorated habitats, to warm, "trailing" range edges, where populations are contracting from newly unsuitable areas.Population contraction signals that compensatory changes in vital rates are insufficient to buffer population growth from deteriorating environments. Life history theory predicts tradeoffs between fast development, high reproduction, and short longevity at low latitudes and slow development, less frequent but multiple bouts of reproduction, and long lifespan at high latitudes. If demographic compensation is driven by life history evolution, compensatory negative correlations in vital rates may be associated with this fast-slow continuum. An outstanding question is whether range limits and range contractions reflect inadequate compensatory life history shifts along environmental gradients, causing population growth rates to fall below replacement levels at range edges. We surveyed demography of 32 populations of the scarlet monkeyflower (Erythranthe cardinalis) spanning 11˚ latitude in western North America and used integral projection models to infer population dynamics and assess demographic compensation. Population growth rates decreased from north to south, consistent with leading-trailing dynamics. Southern populations are declining due to reduced survival, growth, and recruitment, despite compensatory increases in reproduction and faster life history characteristics, suggesting that demographic compensation will not rescue populations at the trailing range edge.Publication 581, Washington, D. C.). 23.Geber MA, Eckhart VM (2005) Experimental studies of adaptation in Clarkia xantiana. II. Fitness variation across a supspecies border. Evolution 59(3):521-531. 24. Angert AL, Schemske DW (2005) The evolution of species' distributions: reciprocal transplants across the elevation ranges of Mimulus cardinalis and M. lewisii. Evolution 59(8):222-235. 25. Sexton JP, Dickman EE (2016) What can local and geographic population limits tell us about distributions? Am J Bot 103(1):129-39. 26. Vaupel A, Matthies D (2012) Abundance, reproduction, and seed predation of an alpine plant decrease from the center toward the range limit. Ecology 93(10):2253-2262. 27. Jump AS, Woodward FI (2003) Seed production and population density decline approaching the range-edge of Cirsium species. New Phytol 160(2):349-358. 28. Eckhart VM, et al. (2011) The geography of demography: Long-term demographic studies and species distribution models reveal a species border limited by adaptation. Am Nat 178(S1):S26-S43. 29. Stanton-Geddes J, Tiffin P, Shaw RG (2012) Role of climate and competitors in limiting fitness across range edges of an annual plant. Ecology 93(7):160...