Legume effects in a native community invaded by alien Asteraceae in a multi-species comparison.

Facilitation has been a long-neglected type of interaction but received more attention recently. Legumes are commonly involved in facilitative interactions due to their nitrogen fixation. Facilitative interactions are so far underappreciated yet potentially important for biological invasions, especially given increasing numbers of alien species. In a common garden experiment using 30 annual Asteraceae species (neophytes, archaeophytes, plus some natives), grown in communities with or without legume presence, we measured functional traits and fitness in focal Asteraceae, as well as nitrogen characteristics of Asteraceae and two native community phytometer species. We investigated how legume presence affects relationships between trait and nitrogen concentration and Asteraceae fitness; and whether mechanisms of facilitation in legume presence and its effects on aboveground performance differ among native phytometer, neophyte, and archaeophyte Asteraceae using the δ15N natural abundance method. Lower specific leaf area was associated with higher aboveground biomass and seed production, with a stronger effect in legume absence. Nitrogen concentration had a positive relationship with biomass, but did not generally increase seed production. Our results hint at N facilitation for the native grass phytometer Festuca rupicola when growing in legume presence, whereas the forb Potentilla argentea and 27 alien Asteraceae species did not indicate facilitative effects. Intriguingly, direct legume facilitation in native phytometer species was only detected when growing with archaeophytes neighbors, not with neophytes. This hints at varied mechanisms of competition for nitrogen between natives and alien species of different residence time and deepens the understanding of altered facilitative leguminous effects in alien species presence.

Native and alien species suffer from late arrival, while negative effects of multiple alien species on natives vary.

Ongoing globalisation and climate change are causing plant species to invade new habitats and thereby alter biodiversity and ecosystem functioning. Since numbers of plant invasions continue to increase globally, it is crucial to investigate the effects of multiple co-occurring alien species on native communities. Furthermore, priority effects due to the earlier emergence of certain species affecting fitness of later arriving species can shape community structure and affect native species performance. We investigate in a common garden pot experiment the interactions among five alien-native species pairs. First we focus on the effect of growing with either one or two alien neighbour species on a native plant, second we alter the arrival time of the alien or native neighbour by 3 weeks. Generally, native species performance decreased when surrounded by two alien species compared to only one, although the magnitude of this effect varied depending on species, with one species even performing better with alien neighbours than in monoculture. Species performance greatly decreased when arriving second in the pot, for both native and alien species. In contrast, alien species tended to benefit more from arriving early. Given that we studied annual ruderal species, their potentially lower competitive ability might explain why we detected negative effects of late arrival. We highlight the need to further elucidate underlying mechanisms of small-scale invasion dynamics to achieve generalisations concerning the response of multiple alien and native plants given their species-specific differences in response to neighbour species and arrival time.

Predicting plant invasions under climate change: are species distribution models validated by field trials?

Climate change may facilitate alien species invasion into new areas, particularly for species from warm native ranges introduced into areas currently marginal for temperature. Although conclusions from modelling approaches and experimental studies are generally similar, combining the two approaches has rarely occurred. The aim of this study was to validate species distribution models by conducting field trials in sites of differing suitability as predicted by the models, thus increasing confidence in their ability to assess invasion risk. Three recently naturalized alien plants in New Zealand were used as study species (Archontophoenix cunninghamiana, Psidium guajava and Schefflera actinophylla): they originate from warm native ranges, are woody bird-dispersed species and of concern as potential weeds. Seedlings were grown in six sites across the country, differing both in climate and suitability (as predicted by the species distribution models). Seedling growth and survival were recorded over two summers and one or two winter seasons, and temperature and precipitation were monitored hourly at each site. Additionally, alien seedling performances were compared to those of closely related native species (Rhopalostylis sapida, Lophomyrtus bullata and Schefflera digitata). Furthermore, half of the seedlings were sprayed with pesticide, to investigate whether enemy release may influence performance. The results showed large differences in growth and survival of the alien species among the six sites. In the more suitable sites, performance was frequently higher compared to the native species. Leaf damage from invertebrate herbivory was low for both alien and native seedlings, with little evidence that the alien species should have an advantage over the native species because of enemy release. Correlations between performance in the field and predicted suitability of species distribution models were generally high. The projected increase in minimum temperature and reduced frosts with climate change may provide more suitable habitats and enable the spread of these species.

Alien plant fitness is limited by functional trade-offs rather than a long-term increase in competitive effects of native communities.

Alien plants experience novel abiotic conditions and interactions with native communities in the introduced area. Intra- and interspecific selection on functional traits in the new environment may lead to increased population growth with time since introduction (residence time). However, selection regimes might differ depending on the invaded habitat. Additionally, in high-competition habitats, a build-up of biotic resistance of native species due to accumulation of eco-evolutionary experience to aliens over time may limit invasion success. We tested if the effect of functional traits and the population dynamics of aliens depends on interspecific competition with native plant communities. We conducted a multi-species experiment with 40 annual Asteraceae that differ in residence time in Germany. We followed their population growth in monocultures and in interspecific competition with an experienced native community (varying co-existence times between focals and community). To more robustly test our findings, we used a naïve community that never co-existed with the focals. We found that high seed mass decreased population growth in monocultures but tended to increase population growth under high interspecific competition. We found no evidence for a build-up of competition-mediated biotic resistance by the experienced community over time. Instead, population growth of the focal species was similarly inhibited by the experienced and naïve community. By comparing the effect of experienced and naïve communities on population dynamics over 2 years across a large set of species with a high variation in functional traits and residence time, this study advances the understanding of the long-term dynamics of plant invasions. In our study system, population growth of alien species was not limited by an increase of competitive effects by native communities (one aspect of biotic resistance) over time. Instead, invasion success of alien plants may be limited because initial spread in low-competition habitats requires different traits than establishment in high-competition habitats.

Effects of Residence Time, Auto-Fertility and Pollinator Dependence on Reproductive Output and Spread of Alien and Native Asteraceae.

Alien plants benefit from auto-fertility to spread over areas where the lack of co-evolved mutualists would otherwise limit invasion success. However, the widespread generalists among mutualists and their large geographical ranges allow alien plants to be integrated into networks. The role of residence time also has to be accounted for, as it takes time for a species to spread and adapt to a new area. We investigated how residence time, auto-fertility and pollinator dependence affect reproductive output and invasion success of Asteraceae in Germany. We conducted a multi-species common-garden experiment along an alien-native continuum including 42 species of natives, archaeophytes and neophytes (casual and established), subjecting plant individuals either to free access or exclusion of pollinators. Pollinator dependence does not play a crucial role in invasion success, with most Asteraceae being able to self-fertilize. Surprisingly, both established neophytes and natives showed higher abilities to self-fertilize, while archaeophytes and casual neophytes were more attractive to pollinators. In contrast to casual neophytes, the established neophytes' strategy was associated with a large reproductive output. Yet, auto-fertility was not associated with range size, since archaeophytes reached the largest range sizes. Elucidating how breeding systems affect invasion success is crucial for predicting and managing invasions.