Species range limits involve many aspects of evolution and ecology, from species distribution and abundance to the evolution of niches. Theory suggests myriad processes by which range limits arise, including competitive exclusion, Allee effects, and gene swamping; however, most models remain empirically untested. Range limits are correlated with a number of abiotic and biotic factors, but further experimentation is needed to understand underlying mechanisms. Range edges are characterized by increased genetic isolation, genetic differentiation, and variability in individual and population performance, but evidence for decreased abundance and fitness is lacking. Evolution of range limits is understudied in natural systems; in particular, the role of gene flow in shaping range limits is unknown. Biological invasions and rapid distribution shifts caused by climate change represent large-scale experiments on the underlying dynamics of range limits. A better fusion of experimentation and theory will advance our understanding of the causes of range limits. 415 Annu. Rev. Ecol. Evol. Syst. 2009.40:415-436. Downloaded from www.annualreviews.org by Stanford University -Main Campus -Lane Medical Library on 10/08/12. For personal use only.

Gene flow among populations can enhance local adaptation if it introduces new genetic variants available for selection

The range of resources that a species uses (i.e. its niche breadth) might determine the geographical area it can occupy, but consensus on whether a niche breadth-range size relationship generally exists among species has been slow to emerge. The validity of this hypothesis is a key question in ecology in that it proposes a mechanism for commonness and rarity, and if true, may help predict species' vulnerability to extinction. We identified 64 studies that measured niche breadth and range size, and we used a meta-analytic approach to test for the presence of a niche breadth-range size relationship. We found a significant positive relationship between range size and environmental tolerance breadth (z = 0.49), habitat breadth (z = 0.45), and diet breadth (z = 0.28). The overall positive effect persisted even when incorporating sampling effects. Despite significant variability in the strength of the relationship among studies, the general positive relationship suggests that specialist species might be disproportionately vulnerable to habitat loss and climate change due to synergistic effects of a narrow niche and small range size. An understanding of the ecological and evolutionary mechanisms that drive and cause deviations from this niche breadth-range size pattern is an important future research goal.

How ecological niche breadth evolves is central to adaptation and speciation and has been a topic of perennial interest. Niche breadth evolution research has occurred within environmental, ecological, evolutionary, and biogeographical contexts, and although some generalities have emerged, critical knowledge gaps exist. Performance breadth trade-offs, although long invoked, may not be common determinants of niche breadth evolution or limits. Niche breadth can expand or contract from specialist or generalist lineages, and so specialization need not be an evolutionary dead end. Whether niche breadth determines diversification and distribution breadth and how niche breadth is partitioned among individuals and populations within a species are important but particularly understudied topics. Molecular genetic and phylogenetic techniques have greatly expanded understanding of niche breadth evolution, but field studies of how niche breadth evolves are essential for providing mechanistic details and allowing the development of comprehensive theory and improved prediction of biological responses under global change.

According to theory, gene flow to marginal populations may stall or aid adaptation at range limits by swamping peripheral populations with maladaptive gene flow or by enhancing genetic variability and reducing inbreeding depression, respectively. We tested these contrasting predictions by manipulating patterns of gene flow of the annual plant, Mimulus laciniatus, at its warm range limit. Gene flow was experimentally applied by using crosses within warm-limit populations (selfed and outcrossed), between warm-limit populations, and between warm-limit and central range populations across two elevational transects. We measured the fitness of offspring in a common garden at the warm-edge species range limit. All sources of gene flow increased seedling emergence at the range limit, suggesting local inbreeding depression at both range limit populations; however, lifetime reproductive success only increased significantly when pollen originated from another warm-limit population. Center-to-warm-edge gene flow was maladaptive by delaying time to development at this warm, fast-drying range limit, whereas edge-to-edge gene flow hastened emergence time and time to reproduction. By empirically testing theory on the effects of gene flow on the formation of geographic range limits, we find benefits of gene flow among populations to be greatest when gene flow is between populations occupying the same range limit. Our results emphasize the overlooked importance of gene flow among populations occurring near the same range limit and highlight the potential for prescriptive gene flow as a conservation option for populations at risk from climate change.climate adaptation | ecological gradients | natural selection | phenology | species range limits T heory on the evolution of range limits predicts that gene flow from large, central populations to edge populations at the range limit could create a flood of maladaptive, nonlocal genes, thereby stalling adaptation and niche expansion (1-3). Alternatively, gene flow from central populations may increase effective population size and genetic variation in edge populations, thereby ultimately increasing fitness at the range limit and perhaps contributing to range expansion (4-6). Overlooked in these models is gene flow among edge populations, which may be especially beneficial because it (i) supplies both favorable alleles or gene combinations that are adaptive at range limits (7) and (ii) enhances genetic variation upon which selection may act. These ideas have not been tested empirically in natural systems at geographic range limits.With rapid climate change, low genetic variation may constrain the ability of populations to adapt quickly to warming environments (8-10). Globally, climate warming is pushing species ranges upwards in elevation, leaving rear-edge populations to adapt, migrate, or perish (9, 10). Mimulus laciniatus (cutleaved monkeyflower), an annual plant, inhabits mossy areas on granite seeps between 975 and 3,270 m on the western slope of the California Sierra Nevada Mountai...