Temporal dynamics of seed excretion by wild ungulates: implications for plant dispersal

Dispersal is a key process in metapopulation dynamics as it conditions species' spatial responses to gradients of abiotic and biotic conditions and triggers individual and gene flows. In the numerous plants that are dispersed through seed consumption by herbivores (endozoochory), the distance and effectiveness of dispersal is determined by the combined effects of seed retention time in the vector's digestive system, the spatial extent of its movements, and the ability of the seeds to germinate once released. Estimating these three parameters from experimental data is therefore crucial to calibrate mechanistic metacommunity models of plant–herbivore interactions. In this study, we jointly estimated the retention time and germination probability of six herbaceous plants transported by roe deer (Capreolus capreolus), red deer (Cervus elaphus), and wild boar (Sus scrofa) through feeding experiments and a Bayesian dynamic model. Retention time was longer in the nonruminant wild boar (>36 h) than in the two ruminant species (roe deer: 18–36 h, red deer: 3–36 h). In the two ruminants, but not in wild boar, small and round seeds were excreted faster than large ones. Low germination probabilities of the excreted seeds reflected the high cost imposed by endozoochory on plant survival. Trait‐mediated variations in retention time and germination probability among animal and plant species may impact plant dispersal distances and interact with biotic and abiotic conditions at the release site to shape the spatial patterns of dispersed plant species.     

Abiotic conditions shape the relationship between indigenous and exotic species richness in a montane biodiversity hotspot

Montane ecosystems are more prone to invasions by exotic plant species than previously thought. Besides abiotic factors, such as climate and soil properties, plant-plant interactions within communities are likely to affect the performance of potential invaders in their exotic range. The biotic resistance hypothesis predicts that high indigenous species richness hampers plant invasions. The biotic acceptance hypothesis, on the other hand, predicts a positive relationship between indigenous and exotic species richness. We tested these two hypotheses using observational data along an elevational gradient in a southern African biodiversity hotspot. Species composition data of indigenous and exotic plants were recorded in 20 road verge plots along a gradient of 1775–2775 m a.s.l. in the Drakensberg, South Africa. Plots were 2?×?50 m in size and positioned at 50 m elevational intervals. We found a negative correlation between indigenous and exotic richness for locations with poorly developed mineral soils, suggesting biotic resistance through competitive interactions. A strong positive correlation for plots with very shallow soils at high elevations indicated a lack of biotic resistance and the possibility of facilitating interactions in harsher environments. These results suggest that biotic resistance is restricted to the lower and mid elevations while biotic acceptance prevails in presence of severe abiotic stress, potentially increasing the risk of plant invasions into montane biodiversity hotspots.

     

Management of invasive plants through ecological resistance

Despite debates on the real impact of plant invasion on native biodiversity, there remain many situations where exotic invasive plants must be managed and habitats restored. Restoration practices that build on plant community assembly principles could be useful to delay or prevent re-invasion after control, but there are still few syntheses of the biodiversity theory, ecological mechanisms and experimental evidence relevant to invasive plant management, possibly delaying applications. To provide such a synthesis, we review current knowledge on three key determinants of invasion success: biotic resistance, abiotic constraints, and propagule pressure. We elaborate on the ecological mechanisms at play for each determinant and emphasize, using case studies, their relevance for invasive plant management and ecological restoration. We find evidence that restoring a plant cover can enhance invasion resistance, but the challenge for both research and field applications is to understand how multiple determinants interact in relation to species traits in the fields. Failure to recognize these interactions and their effect on community assembly processes may explain some of the mixed species responses observed. While we need control and restoration case studies with local species at different sites, the development of a coherent, dynamic and adaptive framework around biotic/ecological resistance will have to go beyond the idiosyncrasy of the many species and systems being tested. Emphasizing the functional diversity of the restored community seems a promising approach when facing potentially multiple invaders and/or fluctuating abiotic conditions.     

Long‐term consistency in spatial patterns of primate seed dispersal

Seed dispersal is a key ecological process in tropical forests, with effects on various levels ranging from plant reproductive success to the carbon storage potential of tropical rainforests. On a local and landscape scale, spatial patterns of seed dispersal create the template for the recruitment process and thus influence the population dynamics of plant species. The strength of this influence will depend on the long‐term consistency of spatial patterns of seed dispersal. We examined the long‐term consistency of spatial patterns of seed dispersal with spatially explicit data on seed dispersal by two neotropical primate species, Leontocebus nigrifrons and Saguinus mystax (Callitrichidae), collected during four independent studies between 1994 and 2013. Using distributions of dispersal probability over distances independent of plant species, cumulative dispersal distances, and kernel density estimates, we show that spatial patterns of seed dispersal are highly consistent over time. For a specific plant species, the legume Parkia panurensis, the convergence of cumulative distributions at a distance of 300 m, and the high probability of dispersal within 100 m from source trees coincide with the dimension of the spatial–genetic structure on the embryo/juvenile (300 m) and adult stage (100 m), respectively, of this plant species. Our results are the first demonstration of long‐term consistency of spatial patterns of seed dispersal created by tropical frugivores. Such consistency may translate into idiosyncratic patterns of regeneration.     

Seed rain and environmental controls on invasion of <Emphasis Type="Italic">Picea abies</Emphasis> into grassland

Although changes in land-use, climate, and the spread of introduced tree species have increased the global importance of tree invasions into grasslands, our ability to predict any particular invasion is limited. To elucidate mechanisms driving tree invasions of grasslands, we studied in detail how seed dispersal and fine-scale environment control the expansion of an introduced Picea abies Karst. (Norway spruce) population into Western Carpathian grassland. We mapped invading trees and measured tree size, fecundity, seed rain, seedling density, plant community composition, and light and soil environment within a 200 × 60 m belt across the invasion front. Maximum likelihood estimates of dispersal kernels suggested peak seed deposition directly underneath tree crowns where germination was poor, but mean dispersal distances were sufficiently large to generate overlapping seed shadows from multiple trees that saturated the invasion front with seeds further away from seed-dispersing trees. Partial Mantel tests indicated that germinant density was affected considerably less by seed rain than by moss cover (r = 0.54), overstory tree influence (r = −0.32), soil moisture (r = 0.21), grass cover (r = −0.15), and diffuse radiation (r = 0.13). However, these variables were not independent but formed complex multivariate gradients within the invasion front. Moss cover and soil moisture were negatively correlated with overstory tree influence and the resulting gradient clearly affected germinant density (partial Mantel r = 0.45). In contrast, positively correlated light and grass cover defined a gradient related weakly to germinant density (partial Mantel r = 0.05) as it integrated opposing effects of these variables on germinants. Seedlings had similar environmental associations, but except for the lasting positive effects of moss these tended to weaken with seedling size. Although a few seedlings may establish and survive in the more adverse environment of the outer edges of the invasion front, a significant population expansion may require a gradual build-up of the critical density of invading trees to reduce grass cover and facilitate germination on moist mossy seedbeds within uncolonized areas. Thus, Picea abies appears more likely to spread within temperate grasslands by gradual expansion of its population frontier rather than by advanced groups.     

Pisonia grandis monocultures limit the spread of an invasive ant—a case of carbohydrate quality?

The mechanisms by which invasive species are able to spread into and dominate natural communities are poorly understood and remain a focus of invasion research. In this quest, studying invasions that are limited by a controlling factor will be more informative than will studies documenting unabated spread and impacts. Some ant species are very successful invaders, and research demonstrating abiotic and biotic factors limiting their success has aided the understanding of invasion ecology. We report here a study showing the highly invasive African big headed ant Pheidole megacephala having a novel distribution on coral cays within Australia’s Great Barrier Reef. These patterns displayed a clear limitation of its distribution with monocultures of the tree Pisonia grandis. This distribution was contrary to the known environmental limitations of the ant, and the limitation could not be associated with an underlying abiotic determinant of the vegetation type. We present these distributional patterns, and following consideration of all known biotic and abiotic limitations of ant invasions we discuss the potential that the peculiar ecophysiology of P. grandis is the causal factor. Specifically, we suggest that the quality of carbohydrate supply to ants is a limitation to invasive spread in much the same way that carbohydrate quantity is known to affect ant population densities in other ecosystems.     

Invasive plants may promote predator‐mediated feedback that inhibits further invasion

Understanding the impacts of invasive species requires placing invasion within a full community context. Plant invaders are often considered in the context of herbivores that may drive invasion by avoiding invaders while consuming natives (enemy escape), or inhibit invasion by consuming invaders (biotic resistance). However, predators that attack those herbivores are rarely considered as major players in invasion. Invasive plants often promote predators, generally by providing improved habitat. Here, we show that predator‐promoting invaders may initiate a negative feedback loop that inhibits invasion. By enabling top‐down control of herbivores, predator‐promoting invaders lose any advantage gained through enemy escape, indirectly favoring natives. In cases where palatable invaders encounter biotic resistance, predator promotion may allow an invader to persist, but not dominate. Overall, results indicate that placing invaders in a full community context may reveal reduced impacts of invaders compared to expectations based on simple plant–plant or plant–herbivore subsystems.     

Spatial autocorrelation and the analysis of invasion processes from distribution data: a study with the crayfish <Emphasis Type="Italic">Procambarus clarkii</Emphasis>

Complex spatial dynamics are frequent in invasive species; analyzing distribution patterns can help to understand the mechanisms driving invasions. We used different spatial regression techniques to evaluate processes determining the invasion of the red swamp crayfish Procambarus clarkii. We evaluated four a priori hypotheses on processes that may determine crayfish invasion: landscape alteration, connectivity, wetland suitability for abiotic and biotic features. We assessed the distribution of P. clarkii in 119 waterbodies in a recently invaded area. We used spatially explicit statistical techniques (spatial eigenvector mapping, generalized additive models, Bayesian intrinsic conditional autoregressive models) within an information-theoretic framework to assess the support of hypotheses; we also analyzed the pattern of spatial autocorrelation of data, model residuals, and eigenvectors. We found strong agreement between the results of spatial eigenvector mapping and Bayesian autoregressive models. Procambarus clarkii was significantly associated with the largest, permanent wetlands. Additive models suggested also association with human-dominated landscapes, but tended to overfit data. The results indicate that abiotic wetlands features and landscape alteration are major drivers of the species’ distribution. Species distribution data, residuals of ordinary least squares regression, and spatial eigenvectors all showed positive and significant spatial autocorrelation at distances up to 2,500 m; this may be caused by the dispersal ability of the species. Our analyses help to understand the processes determining the invasion and to identify the areas most at risk where screening and early management efforts can be focused. The comparison of multiple spatial techniques allows a robust assessment of factors determining complex distribution patterns.     

A 10-year study of changes in forest vegetation on Silhouette island, Seychelles

A 10 year study of forest communities on Silhouette island, Seychelles demonstrates stability of forest composition in most areas over this time-scale. Areas with heavy invasion by alien species were found to be regenerating, particularly with the rapid loss of Clidemia hirta. This is attributed to the abundance of well-adapted native plants allowing competitive exclusion to take place, throughout competition for light. It was noted that invasive plant species tend to be unstable on the rocky slopes covered by native high forest. A high rate of tree fall and limited seed dispersal may reduce the impact of the invasive tree Paraserianthes falcataria in the future. Other species such as Cinnamomum verum and Psidium cattleianum may persist as major invaders due to wider seed dispersal.     

Division within the North American boreal forest: Ecological niche divergence between the Bicknell's Thrush (Catharus bicknelli) and Gray‐cheeked Thrush (C. minimus)

Sister species that diverged in allopatry in similar environments are expected to exhibit niche conservatism. Using ecological niche modeling and a multivariate analysis of climate and habitat data, I test the hypothesis that the Bicknell's Thrush (Catharus bicknelli) and Gray‐cheeked Thrush (C. mimimus), sister species that breed in the North American boreal forest, show niche conservatism. Three tree species that are important components of breeding territories of both thrush species were combined with climatic variables to create niche models consisting of abiotic and biotic components. Abiotic‐only, abiotic+biotic, and biotic‐only models were evaluated using the area under the curve (AUC) criterion. Abiotic+biotic models had higher AUC scores and did not over‐project thrush distributions compared to abiotic‐only or biotic‐only models. From the abiotic+biotic models, I tested for niche conservatism or divergence by accounting for the differences in the availability of niche components by calculating (1) niche overlap from ecological niche models and (2) mean niche differences of environmental values at occurrence points. Niche background similarity tests revealed significant niche divergence in 10 of 12 comparisons, and multivariate tests revealed niche divergence along 2 of 3 niche axes. The Bicknell's Thrush breeds in warmer and wetter regions with a high abundance of balsam fir (Abies balsamea), whereas Gray‐cheeked Thrush often co‐occurs with black spruce (Picea mariana). Niche divergence, rather than conservatism, was the predominant pattern for these species, suggesting that ecological divergence has played a role in the speciation of the Bicknell's Thrush and Gray‐cheeked Thrush. Furthermore, because niche models were improved by the incorporation of biotic variables, this study validates the inclusion of relevant biotic factors in ecological niche modeling to increase model accuracy.     

Abiotic and biotic resistance to grass invasion in serpentine annual plant communities

Biological invasions severely impact native plant communities, causing dramatic shifts in species composition and the restriction of native species to spatially isolated refuges. Competition from resident species and the interaction between resource limitation and competition have been overlooked as mechanisms of community resistance in refugia habitats. We examined the importance of these factors in determining the resistance of California serpentine plant communities to invasion by three common European grasses, Avena barbata, Bromus diandrus, and Hordeum murinum. We added seeds of each of these grasses to plots subjected to six levels of resource addition (N, P, Ca, H₂O, all resources together, and a no-addition control) and two levels of competition (with resident community present or removed). Resource limitation and competition had strong effects on the biomass and reproduction of the three invaders. The addition of all resources together combined with the removal of the resident community yielded individual plants that were fourfold to 20-fold larger and sixfold to 20-fold more fecund than plants from control plots. Competitor removal alone yielded invaders that were twofold to sevenfold larger and twofold to ninefold more fecund. N addition alone or in combination with other resources led to a twofold to ninefold increase in the biomass and fecundity of the invaders. No other resource alone significantly affected native or invader performance, suggesting that N was the key limiting resource during our experiment. We found a significant interaction between abiotic and biotic resistance for Bromus, which experienced increased competitive suppression in fertilized plots. The threefold increase in resident biomass with N addition was likely responsible for this result. Our results confirm that serpentine plant communities are severely N limited, which, in combination with competition from resident species, promotes the resistance of these systems to invasions. Our work suggests that better understanding the relative sensitivities of invaders and residents to the physical environment is critical to predicting how abiotic and biotic factors interact to determine community resistance.     

Resistance of Island Rainforest to Invasion by Alien Plants: Influence of Microhabitat and Herbivory on Seedling Performance

Experiments have promise in determining mechanisms by which communities resist invasion. Growth and survivorship of transplanted seedlings of introduced tree species (Leucaena leucocephala, Muntingia calabura, Adenanthera pavonia, and Clausena excavata) were used to assess abiotic (light regime) and biotic resistance (herbivory) to invasion of rainforest on Christmas Island (Indian Ocean). At four sites, seedlings were transplanted into the forest edge along roadside verges and into adjacent intact forest; half were caged to prevent access by the dominant seedling consumer, the red land crab (Gecarcoidea natalis). Red crab densities did not differ between roadside and forest plots. Red crabs initially reduced survivorship of Leucaena in both edge and interior plots but virtually all seedlings in the forest interior were dead after 41 weeks. Survivorship of Muntingia was also initially reduced by crabs in the forest edge, but again, all but one seedling died by the end of the experiment. Seedlings of Adenanthera and Clausena fared much better overall, surviving well in both locations. Red crabs had no overall effect on Adenanthera survival but significantly reduced survival of Clausena in forest plots. For both species, seedling performance was greater in the forest edge than in the forest interior. Red crabs had no effect on height increment for either species; however, for Clausena, red crabs reduced seedling mass in the forest interior. Both Adenanthera and Clausena were able to persist in the intact forest. Clausena is now actively invading intact rainforest, but Adenanthera appears dispersal-limited. Resistance factors in intact forest appear hierarchical: biotic resistance afforded by land crabs can impede establishment of some plant invaders but seedling responses to abiotic factors (e.g., the light regime) largely overwhelm its effect. Together, these two community attributes are likely to restrict the range of plant invaders to a small suite of species that can successfully establish in intact rainforest on the island. However, increased propagule pressure from a variety of shade-tolerant species and further declines in the abundance of the dominant seedling consumer may lead to increasing invasion success in this island rainforest. This revised version was published online in July 2006 with corrections to the Cover Date.     

A meta-analysis of biotic resistance to exotic plant invasions

Biotic resistance describes the ability of resident species in a community to reduce the success of exotic invasions. Although resistance is a well‐accepted phenomenon, less clear are the processes that contribute most to it, and whether those processes are strong enough to completely repel invaders. Current perceptions of strong, competition‐driven biotic resistance stem from classic ecological theory, Elton's formulation of ecological resistance, and the general acceptance of the enemies‐release hypothesis. We conducted a meta‐analysis of the plant invasions literature to quantify the contribution of resident competitors, diversity, herbivores and soil fungal communities to biotic resistance. Results indicated large negative effects of all factors except fungal communities on invader establishment and performance. Contrary to predictions derived from the natural enemies hypothesis, resident herbivores reduced invasion success as effectively as resident competitors. Although biotic resistance significantly reduced the establishment of individual invaders, we found little evidence that species interactions completely repelled invasions. We conclude that ecological interactions rarely enable communities to resist invasion, but instead constrain the abundance of invasive species once they have successfully established.     

Co-invasion of similar invaders results in analogous ecological impact niches and no synergies

Exotic species can cause changes to their invaded ecosystems, which can be large and long lasting. Despite most landscapes being invaded by multiple exotic plant species, >90 % of impact studies only characterize the impacts of single species. Therefore, our knowledge of invasive plant impacts does not reflect the co-invaded nature of most landscapes, potentially ignoring complex interactions among exotic species. Our objective was to characterize potential invader synergies (positive interactions) on biotic and abiotic ecological parameters among the important forest invaders Japanese stiltgrass (Microstegium vimineum) and wavyleaf basketgrass (Oplismenus undulatifolius), which co-invade eastern US deciduous forests. To characterize synergies, we used a factorial selective removal study, as well as an observational study to further explore invader cover-impact relationships. Although both invaders can reduce native plant richness by 70 % individually or in combination, there were no impact synergies. Total cover of any combination of the two invaders had a negative quadratic effect on total, exotic, and native plant richness; i.e., all community metrics were greatest at intermediate levels of total invader cover and lowest at maximum invader cover. Native richness was more greatly affected than exotic richness by the co-invasion. Soil metrics had no clear trend in either study. Japanese stiltgrass and wavyleaf basketgrass appear to have overlapping impact niches—the number, magnitude, and direction of biotic and abiotic changes to the invaded ecosystem—that only vary in impact magnitude, not breadth. As a result of their overlapping impact niches and non-synergies in this co-invaded system, the addition of the recent invader wavyleaf basketgrass has not resulted in additional changes to the invaded forests. Future impact studies should focus on multiple species and identifying synergies, especially as they relate to invader cover, which informs ecological interactions and management prioritization.     

Accounting for dispersal and biotic interactions to disentangle the drivers of species distributions and their abundances

Although abiotic factors, together with dispersal and biotic interactions, are often suggested to explain the distribution of species and their abundances, species distribution models usually focus on abiotic factors only. We propose an integrative framework linking ecological theory, empirical data and statistical models to understand the distribution of species and their abundances together with the underlying community assembly dynamics. We illustrate our approach with 21 plant species in the French Alps. We show that a spatially nested modelling framework significantly improves the model's performance and that the spatial variations of species presence-absence and abundances are predominantly explained by different factors. We also show that incorporating abiotic, dispersal and biotic factors into the same model bring new insights to our understanding of community assembly. This approach, at the crossroads between community ecology and biogeography, is a promising avenue for a better understanding of species co-existence and biodiversity distribution.     

Twenty‐five years of plant community dynamics and invasion in New Zealand tussock grasslands

Understanding how plant communities respond to plant invasions is important both for understanding community structure and for predicting future ecosystem change. In a system undergoing intense plant invasion for 25 years, we investigated patterns of community change at a regional scale. Specifically, we sought to quantify how tussock grassland plant community structure had changed and whether changes were related to increases in plant invasion. Frequency data for all vascular plants were recorded on 124, permanent transects in tussock grasslands across the lower eastern South Island of New Zealand measured three times over a period of 25 years. Multivariate analyses of species richness were used to describe spatial and temporal patterns in the vegetation. Linear mixed‐effects models were used to relate temporal changes in community structure to the level and rate of invasion of three dominant invasive species in the genus Hieracium while accounting for relationships with other biotic and abiotic variables. There was a strong compositional gradient from exotic‐ to native‐dominated plant communities that correlated with increasing elevation. Over the 25 years, small‐scale species richness significantly decreased and then increased again; however, these changes differed in different plant communities. Exotic species frequency consistently increased on some transects and consistently declined on others. Species richness changes were correlated with the level of Hieracium invasion and abiotic factors, although the relationship with Hieracium changed from negative to positive over time. Compositional changes were not related to measured predictors. Our results suggest that observed broad‐scale fluctuations in species richness and community composition dynamics were not driven by Hieracium invasion. Given the relatively minor changes in community composition over time, we conclude that there is no evidence for widespread degradation of these grasslands over the last 25 years. However, because of continuing weed invasion, particularly at lower elevations, impacts may emerge in the longer term.     

Ungulates increase forest plant species richness to the benefit of non‐forest specialists

Large wild ungulates are a major biotic factor shaping plant communities. They influence species abundance and occurrence directly by herbivory and plant dispersal, or indirectly by modifying plant‐plant interactions and through soil disturbance. In forest ecosystems, researchers’ attention has been mainly focused on deer overabundance. Far less is known about the effects on understory plant dynamics and diversity of wild ungulates where their abundance is maintained at lower levels to mitigate impacts on tree regeneration. We used vegetation data collected over 10 years on 82 pairs of exclosure (excluding ungulates) and control plots located in a nation‐wide forest monitoring network (Renecofor). We report the effects of ungulate exclusion on (i) plant species richness and ecological characteristics, (ii) and cover percentage of herbaceous and shrub layers. We also analyzed the response of these variables along gradients of ungulate abundance, based on hunting statistics, for wild boar (Sus scrofa), red deer (Cervus elaphus) and roe deer (Capreolus capreolus). Outside the exclosures, forest ungulates maintained higher species richness in the herbaceous layer (+15%), while the shrub layer was 17% less rich, and the plant communities became more light‐demanding. Inside the exclosures, shrub cover increased, often to the benefit of bramble (Rubus fruticosus agg.). Ungulates tend to favour ruderal, hemerobic, epizoochorous and non‐forest species. Among plots, the magnitude of vegetation changes was proportional to deer abundance. We conclude that ungulates, through the control of the shrub layer, indirectly increase herbaceous plant species richness by increasing light reaching the ground. However, this increase is detrimental to the peculiarity of forest plant communities and contributes to a landscape‐level biotic homogenization. Even at population density levels considered to be harmless for overall plant species richness, ungulates remain a conservation issue for plant community composition.     

Identifying the abiotic and biotic drivers behind the elevational distribution shift of a parasitic plant

• Climate change will alter the biotic and abiotic environment and dissipate ecological barriers, reorganising maps of current distribution of parasites and their hosts. In this study, we analyse the population dynamics of the parasitic plant Viscum album subsp. austriacum and explore key biotic (host availability and seed dispersal) as well as abiotic (temperature) factors influencing elevational distribution.
• The study was conducted along an elevational gradient of a Mediterranean mountain, covering the distribution belts of three potential pine hosts: Pinus halepensis (1300–1500 m), P. nigra (1300–1900 m) and P. sylvestris var. nevadensis (1600–2000 m). Along this gradient, we measured multiple variables of mistletoe population (prevalence, abundance and demographic profile) and different factors that might define the current mistletoe distribution (host suitability and availability, temperature and seed dispersal services).
• We found a decline in mistletoe prevalence and abundance with increasing elevation, detecting larger values of both variables at lower elevations of the most suitable host (Pinus nigra). Pinus sylvestris var. nevadensis was a suboptimal but suitable host for the parasite at high elevations. Mistletoe found suitable temperatures and seed dispersal services all along the gradient, being able to recruit at any site.
• With warming temperatures, the presence of suitable vectors for parasite dispersion, and the presence of a sub‐optimal host (P. sylvestris var. nevadensis) at the mountain top, mistletoe currently has a window of opportunity to expand its present geographic distribution to the summits.

     

Lack of genetic isolation by distance,similar genetic structuring but different demographic histories in a fig‐pollinating wasp mutualism

Historical abiotic factors such as climatic oscillations and extreme climatic events as well as biotic factors have shaped the structuring of species' genetic diversity. In obligate species‐specific mutualisms, the biogeographic histories of the interacting species are tightly linked. This could be particularly true for nuclear genes in the Ficus‐pollinating wasp mutualistic association as the insects disperse pollen from their natal tree. In this study, we compare spatial genetic structure of plant and pollinator for the Ficus hirta–Valisia javana association throughout southeast China including Hainan Island, for both nuclear and cytoplasmic markers. We show that dispersal of the insect leads to plant and insect presenting similar signatures of lack of genetic isolation by distance for nuclear genes on the continent over a distance of 1000 km. But we also show that the demographic histories of plant and insect are strikingly different. This is in agreement with extreme climatic events leading to transient regional extinctions of the insects, associated with local survival of the plants. We also observe evidence of genetic differentiation for both wasps and fig‐tree between the continent and Hainan Island, although the Qiongzhou Strait is only on average 30 km wide, suggesting that geographic isolation by itself has not been sufficient to generate this differentiation. Hence, our results suggest that in highly dispersive mutualistic systems, isolation‐by‐dispersal limitation across a geographic barrier could be supplemented by isolation by adaptation, and maybe by coevolution, allowing further genetic divergence. In such systems, species may frequently be composed of a single population.     

Running off the road: roadside non-native plants invading mountain vegetation

Prevention is regarded as a cost-effective management action to avoid unwanted impacts of non-native species. However, targeted prevention can be difficult if little is known about the traits of successfully invading non-native species or habitat characteristics that make native vegetation more resistant to invasion. Here, we surveyed mountain roads in seven regions worldwide, to investigate whether different species traits are beneficial during primary invasion (i.e. spread of non-native species along roadside dispersal corridors) and secondary invasion (i.e. percolation from roadsides into natural adjacent vegetation), and to determine if particular habitat characteristics increase biotic resistance to invasion. We found primary invasion up mountain roads tends to be by longer lived, non-ruderal species without seed dispersal traits. For secondary invasion, we demonstrate that both traits of the non-native species and attributes of the receiving natural vegetation contribute to the extent of invasion. Non-native species that invade natural adjacent vegetation tend to be shade and moisture tolerant. Furthermore, non-native species invasion was greater when the receiving vegetation was similarly rich in native species. Our results show how mountain roads define which non-native species are successful; first by favouring certain traits in mountain roadsides (the key dispersal pathway to the top), and secondly by requiring a different set of traits when species invade the natural adjacent vegetation. While patterns in species traits were observed at a global level, regional abiotic and biotic variables largely generated region-specific levels of response, suggesting that management should be regionally driven.