Mark van Kleunen

A meta-analysis of trait differences between invasive and non-invasive plant species

A major aim in ecology is identifying determinants of invasiveness. We performed a meta-analysis of 117 field or experimental-garden studies that measured pair-wise trait differences of a total of 125 invasive and 196 non-invasive plant species in the invasive range of the invasive species. We tested whether invasiveness is associated with performance-related traits (physiology, leaf-area allocation, shoot allocation, growth rate, size and fitness), and whether such associations depend on type of study and on biogeographical or biological factors. Overall, invasive species had significantly higher values than non-invasive species for all six trait categories. More trait differences were significant for invasive vs. native comparisons than for invasive vs. non-invasive alien comparisons. Moreover, for comparisons between invasive species and native species that themselves are invasive elsewhere, no trait differences were significant. Differences in physiology and growth rate were larger in tropical regions than in temperate regions. Trait differences did not depend on whether the invasive alien species originates from Europe, nor did they depend on the test environment. We conclude that invasive alien species had higher values for those traits related to performance than non-invasive species. This suggests that it might become possible to predict future plant invasions from species traits.

Are invaders different? A conceptual framework of comparative approaches for assessing determinants of invasiveness

What determines invasiveness of alien organisms is among the most interesting and urgent questions in ecology. In attempts to answer this question, researchers compare invasive alien species either to native species or to non-invasive alien species, and this is done in either the introduced or native ranges. However, inferences that can be drawn from these comparisons differ considerably, and failure to recognize this could hamper the search for determinants of invasiveness. To increase awareness about this issue, we present a framework of the various comparisons that can be used to test for determinants of invasiveness, and the specific questions each comparison can address. Moreover, we discuss how different comparisons complement each other, and therefore should be used in concert. For progress in invasion biology, it is crucial to realize that different comparisons address different biological questions and that some questions can only be answered unambiguously by combining them.

Global exchange and accumulation of non-native plants

All around the globe, humans have greatly altered the abiotic and biotic environment with ever-increasing speed. One defining feature of the Anthropocene epoch is the erosion of biogeographical barriers by human-mediated dispersal of species into new regions, where they can naturalize and cause ecological, economic and social damage. So far, no comprehensive analysis of the global accumulation and exchange of alien plant species between continents has been performed, primarily because of a lack of data. Here we bridge this knowledge gap by using a unique global database on the occurrences of naturalized alien plant species in 481 mainland and 362 island regions. In total, 13,168 plant species, corresponding to 3.9% of the extant global vascular flora, or approximately the size of the native European flora, have become naturalized somewhere on the globe as a result of human activity. North America has accumulated the largest number of naturalized species, whereas the Pacific Islands show the fastest increase in species numbers with respect to their land area. Continents in the Northern Hemisphere have been the major donors of naturalized alien species to all other continents. Our results quantify for the first time the extent of plant naturalizations worldwide, and illustrate the urgent need for globally integrated efforts to control, manage and understand the spread of alien species.

Macrophysiology: A Conceptual Reunification

Widespread recognition of the importance of biological studies at large spatial and temporal scales, particularly in the face of many of the most pressing issues facing humanity, has fueled the argument that there is a need to reinvigorate such studies in physiological ecology through the establishment of a macrophysiology. Following a period when the fields of ecology and physiological ecology had been regarded as largely synonymous, studies of this kind were relatively commonplace in the first half of the twentieth century. However, such large‐scale work subsequently became rather scarce as physiological studies concentrated on the biochemical and molecular mechanisms underlying the capacities and tolerances of species. In some sense, macrophysiology is thus an attempt at a conceptual reunification. In this article, we provide a conceptual framework for the continued development of macrophysiology. We subdivide this framework into three major components: the establishment of macrophysiological patterns, determining the form of those patterns (the very general ways in which they are shaped), and understanding the mechanisms that give rise to them. We suggest ways in which each of these components could be developed usefully.

RAPD variation among and within small and large populations of the rare clonal plant Ranunculus reptans (Ranunculaceae)

In the pre-alpine region of Europe numbers and sizes of populations of the clonal lake shore plant Ranunculus reptans have declined because of the regulation of lake water levels. We investigated genetic variation among and within 17 populations of different size (cover 1-10 000 m) in R. reptans with RAPD (random amplified polymorphic DNA) profiles. We sampled 127 rosettes in 14 populations at Lake Constance and three populations at or near Lake Como. There was significant genetic variation between plants from the two lake regions (5.9%, analysis of molecular variance [AMOVA], P < 0.001), among populations within lake regions (20.4%, P < 0.001), and within populations (73.7%, P < 0.001). Under the assumptions of Wrights island model the variation among populations corresponds to a gene flow of N(e)m = 0.70. Within the 14 Lake Constance populations we detected significant genetic variation among subpopulations separated by only a few metres (4.0% of the within-population variation; P < 0.05). Molecular variance was 24% smaller in small populations covering <100 m area than in larger ones (P < 0.03), indicating that samples from large populations were genetically more variable than samples representing comparable areas of smaller populations. We conclude that gene flow among populations is very limited and that genetic drift has caused reduced genetic variability of smaller populations. Conservation of genetic variability in R. reptans requires persistence of large and also of small populations (because of population differentiation), and it could be enhanced by increasing the size of small populations (to counter genetic drift).

No saturation in the accumulation of alien species worldwide

Although research on human-mediated exchanges of species has substantially intensified during the last centuries, we know surprisingly little about temporal dynamics of alien species accumulations across regions and taxa. Using a novel database of 45,813 first records of 16,926 established alien species, we show that the annual rate of first records worldwide has increased during the last 200 years, with 37% of all first records reported most recently (1970–2014). Inter-continental and inter-taxonomic variation can be largely attributed to the diaspora of European settlers in the nineteenth century and to the acceleration in trade in the twentieth century. For all taxonomic groups, the increase in numbers of alien species does not show any sign of saturation and most taxa even show increases in the rate of first records over time. This highlights that past efforts to mitigate invasions have not been effective enough to keep up with increasing globalization.

ADAPTIVE EVOLUTION OF PLASTIC FORAGING RESPONSES IN A CLONAL PLANT

Stoloniferous rosette plants may show horizontal and vertical foraging responses, such as changes in branching frequency, stolon internode length, leaf length, and height growth of stolons. To study whether such plastic foraging responses constitute an adaptation to het- erogeneity in competition, we studied genetic variation in and fitness consequences of plastic foraging responses in the stoloniferous lake-shore plant Ranunculus reptans. Because plastic foraging responses are likely to have been more strongly selected for in heterogeneous envi- ronments, we used 15 genotypes from competitive, heterogeneous microhabitats and 15 from competition-free, homogeneous microhabitats from Lake Constance (central Europe). We planted vegetatively propagated rosettes of the 30 genotypes (totaling 236 rosettes) into a greenhouse environment with spatially heterogeneous competition. Four replicates of each genotype were planted into the competition-free halves of experimental trays, and four other replicates into the halves with the naturally co-occurring grass Agrostis stolonifera. We found significant variation among genotypes in vertical and horizontal foraging responses. In line with the hypothesis of adaptive plasticity, genotypes from the competitive heterogeneous microhabitats more strongly increased the vertical angle of the first stolon internode (126%) and the specific stolon length (166%) in response to competition than genotypes from the competition-free homogeneous microhabitats. Moreover, we found that genotypes that were more plastic in the vertical angle of the first stolon internode, stolon height, and specific internode length produced more rosettes and flowers than less plastic genotypes (all selection gradients for plasticity . 0.316). Our findings strongly suggest that plastic foraging responses constitute an adaptation to environmental heterogeneity, at least in the stoloniferous R. reptans.

Tradeoffs associated with constitutive and induced plant resistance against herbivory

Several prominent hypotheses have been posed to explain the immense variability among plant species in defense against herbivores. A major concept in the evolutionary ecology of plant defenses is that tradeoffs of defense strategies are likely to generate and maintain species diversity. In particular, tradeoffs between constitutive and induced resistance and tradeoffs relating these strategies to growth and competitive ability have been predicted. We performed three independent experiments on 58 plant species from 15 different plant families to address these hypotheses in a phylogenetic framework. Because evolutionary tradeoffs may be altered by human-imposed artificial selection, we used 18 wild plant species and 40 cultivated garden-plant species. Across all 58 plant species, we demonstrate a tradeoff between constitutive and induced resistance, which was robust to accounting for phylogenetic history of the species. Moreover, the tradeoff was driven by wild species and was not evident for cultivated species. In addition, we demonstrate that more competitive species—but not fast growing ones—had lower constitutive but higher induced resistance. Thus, our multispecies experiments indicate that the competition–defense tradeoff holds for constitutive resistance and is complemented by a positive relationship of competitive ability with induced resistance. We conclude that the studied genetically determined tradeoffs are indeed likely to play an important role in shaping the high diversity observed among plant species in resistance against herbivores and in life history traits.

NO EVIDENCE FOR AN EVOLUTIONARY INCREASED COMPETITIVE ABILITY IN AN INVASIVE PLANT

We tested whether Solidago canadensis, which was introduced from North America into Europe from the 17th century onward, has evolved an increased competitive ability (EICA) at the cost of tolerance against herbivory in its new range. We grew plants from nine introduced European and 10 native American populations in a common garden in Europe. In half of the plants, we simulated herbivory by removing 50% of the leaf area and by spraying them with jasmonic acid. Although plants from Europe had 30.5% larger leaves, they had 27.4% smaller inflorescences and tended to grow less tall (−7.0%) and produce fewer vegetative offspring (−5.0%) than plants from North America. The simulated herbivory treatment did not result in any significant differences in height, inflorescence biomass, or number of vegetative offspring between treatment and control plants. Moreover, there were no significant differences in the response of European and American plants to simulated herbivory, indicating that they did not differ in thei...

Phylogenetically independent associations between autonomous self-fertilization and plant invasiveness

Many plant species have been introduced from their native ranges to new continents, but few have become naturalized or, ultimately, invasive. It has been predicted that species that do not require the presence of compatible mates and the services of pollinators for reproduction will be favored in establishment after long‐distance dispersal. We tested whether this hypothesis, generally referred to as Baker’s law, holds for South African species of Iridaceae (iris family) that have been introduced in other regions for horticultural purposes. Fruit and seed production of flowers from which pollinators had been experimentally excluded was assessed for 10 pairs of species from nine different genera or subgenera. Each species pair comprised one naturalized and one nonnaturalized species, all of which are used in international horticulture. On average, species of Iridaceae that have become naturalized outside their native ranges showed a higher capacity for autonomous fruit and seed production than congeneric species that have not become naturalized. This was especially true for the naturalized species that are considered to be invasive weeds. These results provide strong evidence for the role of autonomous seed production in increasing potential invasiveness in plants.