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      Common garden comparisons of native and introduced plant populations: latitudinal clines can obscure evolutionary inferences

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          Abstract

          Common garden studies are increasingly used to identify differences in phenotypic traits between native and introduced genotypes, often ignoring sources of among-population variation within each range. We re-analyzed data from 32 common garden studies of 28 plant species that tested for rapid evolution associated with biological invasion. Our goals were: (i) to identify patterns of phenotypic trait variation among populations within native and introduced ranges, and (ii) to explore the consequences of this variation for how differences between the ranges are interpreted. We combined life history and physiologic traits into a single principal component (PC ALL) and also compared subsets of traits related to size, reproduction, and defense (PC SIZE, PC REP, and PC DEF, respectively). On average, introduced populations exhibited increased growth and reproduction compared to native conspecifics when latitude was not included in statistical models. However, significant correlations between PC-scores and latitude were detected in both the native and introduced ranges, indicating population differentiation along latitudinal gradients. When latitude was explicitly incorporated into statistical models as a covariate, it reduced the magnitude and reversed the direction of the effect for PC ALL and PC SIZE. These results indicate that unrecognized geographic clines in phenotypic traits can confound inferences about the causes of evolutionary change in invasive plants.

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          Phenotypic and genetic differentiation between native and introduced plant populations.

          Plant invasions often involve rapid evolutionary change. Founder effects, hybridization, and adaptation to novel environments cause genetic differentiation between native and introduced populations and may contribute to the success of invaders. An influential idea in this context has been the Evolution of Increased Competitive Ability (EICA) hypothesis. It proposes that after enemy release plants rapidly evolve to be less defended but more competitive, thereby increasing plant vigour in introduced populations. To detect evolutionary change in invaders, comparative studies of native versus introduced populations are needed. Here, we review the current empirical evidence from: (1) comparisons of phenotypic variation in natural populations; (2) comparisons of molecular variation with neutral genetic markers; (3) comparisons of quantitative genetic variation in a common environment; and (4) comparisons of phenotypic plasticity across different environments. Field data suggest that increased vigour and reduced herbivory are common in introduced plant populations. In molecular studies, the genetic diversity of introduced populations was not consistently different from that of native populations. Multiple introductions of invasive plants appear to be the rule rather than the exception. In tests of the EICA hypothesis in a common environment, several found increased growth or decreased resistance in introduced populations. However, few provided a full test of the EICA hypothesis by addressing growth and defence in the same species. Overall, there is reasonable empirical evidence to suggest that genetic differentiation through rapid evolutionary change is important in plant invasions. We discuss conceptual and methodological issues associated with cross-continental comparisons and make recommendations for future research. When testing for EICA, greater emphasis should be put on competitive ability and plant tolerance. Moreover, it is important to address evolutionary change in characteristics other than defence and growth that could play a role in plant invasions.
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            Phenotypic similarity and the evolutionary significance of countergradient variation.

            Countergradient variation is a geographical pattern of genotypes (with respect to environments) in which genetic influences on a trait oppose environmental influences, thereby minimizing phenotypic change along the gradient. Phenotypic similarity across changing environments ought to be of intense interest because it belies considerable genotypic change. When it occurs in characters that are positively associated with fitness, countergradient variation conflicts with the hypothesis that local adaptation to one environment trades off against performance in another environment. Cases of countergradient variation therefore offer unique insight into the mechanisms that produce and maintain phenotypic similarity and/or differences along environmental gradients. Copyright © 1995. Published by Elsevier Ltd.
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              Rapid evolution of a geographic cline in size in an introduced fly.

              The introduction and rapid spread of Drosophila subobscura in the New World two decades ago provide an opportunity to determine the predictability and rate of evolution of a geographic cline. In ancestral Old World populations, wing length increases clinally with latitude. In North American populations, no wing length cline was detected one decade after the introduction. After two decades, however, a cline has evolved and largely converged on the ancestral cline. The rate of morphological evolution on a continental scale is very fast, relative even to rates measured within local populations. Nevertheless, different wing sections dominate the New versus Old World clines. Thus, the evolution of geographic variation in wing length has been predictable, but the means by which the cline is achieved is contingent.
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                Author and article information

                Journal
                Evol Appl
                Evol Appl
                eva
                Evolutionary Applications
                Blackwell Publishing Ltd (Oxford, UK )
                1752-4571
                1752-4571
                May 2009
                23 December 2008
                : 2
                : 2
                : 187-199
                Affiliations
                [1 ]simpleDepartment of Ecology & Evolutionary Biology, University of Toronto Toronto, ON, Canada
                [2 ]simpleDivision of Biological Sciences, University of Montana Missoula, MT, USA
                Author notes
                Robert I. Colautti, Department of Ecology & Evolutionary Biology, University of Toronto, 25 Willcocks St., Toronto, ON, Canada M5S 3B2. Tel.: 416-978-5603; fax: 416-978-5878; e-mail: rob.colautti@ 123456utoronto.ca
                Article
                10.1111/j.1752-4571.2008.00053.x
                3352372
                25567860
                175a7cd9-6535-4dec-959b-8ce5291ff950
                © 2008 The Authors. Journal compilation © 2008 Blackwell Publishing Ltd
                History
                : 21 August 2008
                : 24 November 2008
                Categories
                Synthesis

                Evolutionary Biology
                common garden,biogeographical comparisons,introduced plants,evolution of invasive species,latitudinal gradients,clinal variation,eica

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