Ecologists often consider how environmental factors limit a species' geographic range. However, recent models suggest that geographic distribution also may be determined by a species' ability to adapt to novel environmental conditions. In this study, we empirically tested whether further evolution would be necessary for northern expansion of the weedy annual cocklebur (Xanthium strumarium) in its native North American range. We transplanted seedlings beyond the northern border and photoperiodically manipulated reproductive timing, a trait important for adaptation to shorter growing seasons at higher latitudes within the range, to determine whether further evolution of this trait would result in a phenotype viable beyond the range. Earlier reproductive induction enabled plants to produce mature seeds beyond the range and to achieve a reproductive output similar to those grown within the range. Therefore, evolution of earlier reproduction in marginal populations would be necessary for northward range expansion. This study is the first to empirically show that evolution in an ecologically important trait would enable a species to survive and reproduce beyond its current range. These results suggest that relatively few traits may limit a species' range and that identifying evolutionary constraints on such traits could be important for predicting geographic distribution.

Adaptation to local environments may be an important determinant of species’ geographic range. However, little is known about which traits contribute to adaptation or whether their further evolution would facilitate range expansion. In this study, we assessed the adaptive value of stress avoidance traits in the common annual Cocklebur (Xanthium strumarium) by performing a reciprocal transplant across a broad latitudinal gradient extending to the species’ northern border. Populations were locally adapted and stress avoidance traits accounted for most fitness differences between populations. At the northern border where growing seasons are cooler and shorter, native populations had evolved to reproduce earlier than native populations in the lower latitude gardens. This clinal pattern in reproductive timing corresponded to a shift in selection from favouring later to earlier reproduction. Thus, earlier reproduction is an important adaptation to northern latitudes and constraint on the further evolution of this trait in marginal populations could potentially limit distribution.

Summary• Here we examined species differences in perception and response to two distinct types of shade cue, reduced photosynthetically active radiation (PAR) with and without reduced red : far red ratio (R : FR), in Polygonum persicaria and Polygonum hydropiper , two closely related annuals of contrasting ecological breadth.• We compared plasticity data for light-gathering traits from glasshouse experiments at equivalently reduced PAR under neutral shade (R : FR 1.03) and green shade (R : FR 0.702).• Species shared the ability to distinguish between the two types of shade, as shown by the ability of each to respond differently to neutral vs green shade for one or more traits. However, the species' responses to these cues differed significantly. Polygonum persicaria expressed stronger shade-tolerance responses (increased leaf allocation and leaf area ratio) to reduced PAR alone than to green shade. By contrast, P. hydropiper expressed slightly less plasticity for these traits in neutral than in green shade.• The pronounced plastic response of P. persicaria to neutral shade may contribute to the range of habitats this widespread species can occupy, which includes neutralshade environments such as urban settings.

Factors promoting the evolution of specialists versus generalists have been little studied in ecological context. In a large‐scale comparative field experiment, we studied genotypes from naturally evolved populations of a closely related generalist/specialist species pair (Polygonum persicaria and P. hydropiper), reciprocally transplanting replicates of multiple lines into open and partially shaded sites where the species naturally co‐occur. We measured relative fitness, individual plasticity, herbivory, and genetic variance expressed in the contrasting light habitats at both low and high densities. Fitness data confirmed that the putative specialist out‐performed the generalist in only one environment, the favorable full sun/low‐density environment to which it is largely restricted in nature, while the generalist had higher lifetime reproduction in both canopy and dense neighbor shade. The generalist, P. persicaria, also expressed greater adaptive plasticity for biomass allocation and leaf size in shaded conditions than the specialist. We found no evidence that the ecological specialization of P. hydropiper reflects either genetically based fitness trade‐offs or maintenance costs of plasticity, two types of genetic constraint often invoked to prevent the evolution of broadly adaptive genotypes. However, the patterns of fitness variance and herbivore damage revealed how release from herbivory in a new range can cause an introduced species to evolve as a specialist in that range, a surprising finding with important implications for invasion biology. Patterns of fitness variance between and within sites are also consistent with a possible role for the process of mutation accumulation (in this case, mutations affecting shade‐expressed phenotypes) in the evolution and/or maintenance of specialization in P. hydropiper.

Adaptive differences among species are often thought to result from developmentally constant trait differences that enhance fitness in alternative environments. Species differences in patterns of individual phenotypic plasticity can also have ecological consequences. Indeed, functionally related constant and plastic traits may interact to determine the phenotype's adaptive value in particular conditions. We compared juvenile shade avoidance traits (height and its components, internode length and node number) across two field density treatments in Polygonumpersicaria and P. hydropiper, annual plant species that co‐occur in pastures comprised of a mosaic of plant densities. We used selection analyses to test trait contributions to fitness in alternative density treatments. Seedlings of both species expressed plasticity for internode elongation in response to density; P. persicaria plants increased internode length and consequently height significantly more in high density than did those of P. hydropiper. As predicted by the shade avoidance hypothesis, increased height was adaptive for both species in high density stands, so P. persicaria plants had higher fitness in this environment. By contrast, node numbers were relatively constant across density treatments in both species: P. hydropiper seedlings consistently produced more nodes than did those of P. persicaria. This constant trait difference contributed to P. hydropiper's greater relative fitness at low density, where more nodes and hence leaves enable plants to better exploit available light. Differences between species in these juvenile shade‐avoidance traits did not result from the evolutionary constraints of lack of heritable variation or costs of plasticity. We discuss how these interspecific trait differences may have been generated by divergent selective histories resulting from differences in herbivore resistance. These results illustrate how adaptive differences in both plastic and constantly expressed traits may jointly contribute to ecological distribution, including coexistence in patchy habitats.