Rapid evolution of leaf physiology in an introduced beach daisy.

Photosynthesis is a key biological process. However, we know little about whether plants change their photosynthetic strategy when introduced to a new range. We located the most likely source population for the South African beach daisy Arctotheca populifolia introduced to Australia in the 1930s, and ran a common-garden experiment measuring 10 physiological and morphological leaf traits associated with photosynthesis. Based on predictions from theory, and higher rainfall in the introduced range, we hypothesized that introduced plants would have a (i) higher photosynthetic rate, (ii) lower water-use efficiency (WUE) and (iii) higher nitrogen-use efficiency. However, we found that introduced A. populifolia had a lower photosynthetic rate, higher WUE and lower nitrogen-use efficiency than did plants from Arniston, South Africa. Subsequent site visits suggested that plants in Arniston may be able to access moisture on a rocky shelf, while introduced plants grow on sandy beaches where water can quickly dissipate. Our unexpected findings highlight that: (1) it is important to compare introduced species to their source population for an accurate assessment of evolutionary change; (2) rainfall is not always a suitable proxy for water availability and (3) introduced species often undergo evolutionary changes, but without detailed ecological information we may not be able to accurately predict the direction of these changes.

Rapid reshaping: the evolution of morphological changes in an introduced beach daisy.

Thousands of species have been introduced to new ranges worldwide. These introductions provide opportunities for researchers to study evolutionary changes in form and function in response to new environmental conditions. However, almost all previous studies of morphological change in introduced species have compared introduced populations to populations from across the species' native range, so variation within native ranges probably confounds estimates of evolutionary change. In this study, we used microsatellites to locate the source population for the beach daisy Arctotheca populifolia that had been introduced to eastern Australia. We then compared four introduced populations from Australia with their original South African source population in a common-environment experiment. Despite being separated for less than 100 years, source and introduced populations of A. populifolia display substantial heritable morphological differences. Contrary to the evolution of increased competitive ability hypothesis, introduced plants were shorter than source plants, and introduced and source plants did not differ in total biomass. Contrary to predictions based on higher rainfall in the introduced range, introduced plants had smaller, thicker leaves than source plants. Finally, while source plants develop lobed adult leaves, introduced plants retain their spathulate juvenile leaf shape into adulthood. These changes indicate that rapid evolution in introduced species happens, but not always in the direction predicted by theory.

Weedy and seedy: the rapid evolution of life-history characteristics in an introduced daisy.

Despite the importance of life-history characteristics in determining a species' success, we still lack basic information about some fundamental life-history elements found across the life cycle of introduced plants. Our study assesses rapid evolutionary divergence in life-history characteristics of the beach daisy Arctotheca populifolia by comparing introduced Australian and source South African plants and measuring eight key variables including seed mass, germination, reproductive output and survival. This is the first study that compares the life history of an introduced plant species with its single original source population, providing a precise and powerful method for detecting evolutionary divergence. We found that introduced A. populifolia has evolved a suite of weedy life-history characteristics in less than 90 years: the introduced plants use a live-fast die-young strategy of germination and survival and produce significantly more inflorescences and more seeds that germinate faster. This knowledge adds to the remarkable data that we already have on the rapid evolutionary divergence occurring in the morphology, physiology and defence of this introduced plant and highlights the speed and scope of evolutionary divergence possible in plants. To fully understand and manage the future of our plant species, we must consider their potential for ongoing change in key aspects of life history.

Evolution of defense and herbivory in introduced plants-Testing enemy release using a known source population, herbivore trials, and time since introduction.

The enemy release hypothesis is often cited as a potential explanation for the success of introduced plants; yet, empirical evidence for enemy release is mixed. We aimed to quantify changes in herbivory and defense in introduced plants while controlling for three factors that might have confounded past studies: using a wide native range for comparison with the introduced range, measuring defense traits without determining whether they affect herbivore preferences, and not considering the effect of time since introduction. The first hypothesis we tested was that introduced plants will have evolved lower levels of plant defense compared to their source population. We grew South African (source) and Australian (introduced) beach daisies (Arctotheca populifolia) in a common-environment glasshouse experiment and measured seven defense traits. Introduced plants had more ash, alkaloids, and leaf hairs than source plants, but were also less tough, with a lower C:N ratio and less phenolics. Overall, we found no difference in defense between source and introduced plants. To determine whether the feeding habits of herbivores align with changes in defense traits, we conducted preference feeding trials using five different herbivore species. Herbivores showed no overall preference for leaves from either group. The second hypothesis we tested was that herbivory on introduced plant species will increase through time after introduction to a new range. We recorded leaf damage on herbarium specimens of seven species introduced to eastern Australia and three native control species. We found no change in the overall level of herbivory experienced by introduced plants since arriving in Australia. CONCLUSION: In the field of invasion ecology, we need to rethink the paradigm that species introduced to a new range undergo simple decreases in defenses against herbivores. Instead, plants are likely to employ a range of defense traits that evolve in both coordinated and opposing ways in response to a plethora of different biotic and abiotic selective pressures.

High genetic diversity is not essential for successful introduction.

Some introduced populations thrive and evolve despite the presumed loss of diversity at introduction. We aimed to quantify the amount of genetic diversity retained at introduction in species that have shown evidence of adaptation to their introduced environments. Samples were taken from native and introduced ranges of Arctotheca populifolia and Petrorhagia nanteuilii. Using microsatellite data, we identified the source for each introduction, estimated genetic diversity in native and introduced populations, and calculated the amount of diversity retained in introduced populations. These values were compared to those from a literature review of diversity in native, confamilial populations and to estimates of genetic diversity retained at introduction. Gene diversity in the native range of both species was significantly lower than for confamilials. We found that, on average, introduced populations showing evidence of adaptation to their new environments retained 81% of the genetic diversity from the native range. Introduced populations of P. nanteuilii had higher genetic diversity than found in the native source populations, whereas introduced populations of A. populifolia retained only 14% of its native diversity in one introduction and 1% in another. Our literature review has shown that most introductions demonstrating adaptive ability have lost diversity upon introduction. The two species studied here had exceptionally low native range genetic diversity. Further, the two introductions of A. populifolia represent the largest percentage loss of genetic diversity in a species showing evidence of substantial morphological change in the introduced range. While high genetic diversity may increase the likelihood of invasion success, the species examined here adapted to their new environments with very little neutral genetic diversity. This finding suggests that even introductions founded by small numbers of individuals have the potential to become invasive.