The evaluation of matrilineal admixtures in “Cerambyx spp.” with opposite protection statuses “Cerambyx Cerdo (protected) and Cerambyx welensii (unprotected pest)” highlights the conflicting nature of the management of these insects in the UNESCO protected mediterranean Dehesa forest (Extremadura, Spain)

In a recent article in Ecology, Leffler et al. (2014) presented a potentially new perspective on the importance of trait differences between native and invasive exotic plants in explaining invasions in local native communities. The new perspective brought forward is that, if trait differences between invasive and native species are likely to be important in explaining exotic plant invasion, the differences must be larger than those observed between native species in the new community. A meta-analysis of previous studies searching for trait differences was presented, with the general finding that the magnitudes of trait differences between invasive and native species tend not to differ from those observed between native species only. Leffler et al. (2014) interpret this result as evidence that trait differences are highly context dependent, and that mechanisms other than trait differences are likely to be more important in most cases of invasion. We acknowledge that there is no universal explanation of successful exotic invasion into native communities. Moreover, we do not believe that invasive plant species always have trait values that differ substantially from the traits present in the native community, or that trait differences are important for invasion in all cases. However, we cannot agree with the criterion stipulated by Leffler et al. (2014), namely that a trait difference between invasive and native species can only be important to invasion success if it is greater than the differences among natives. Leffler et al. (2014) do not explain the logic behind the criterion, but a flaw of the criterion is that it will discount cases when a successfully invading species has intermediate trait values that are not represented by native species. Leffler et al. (2014) seem to focus on trait differences as representing niche differences among species. Consider the scenarios of niche differences among native and exotic invasive species in Fig. 1. If a trait is related to the niche space occupied by native species in the community and the invader, for a trait difference to be important in invasion success under the criterion of Leffler et al. (2014), only the scenario in Fig. 1a would qualify. Here, the invader occupies a niche at the extreme of the niche space, compared to native species. The average niche-related trait difference between the invasive species and the natives will be greater than the average difference among natives. However, consider Fig. 1b. Here, the invader occupies a vacant niche that is intermediate between the native species (Stachowicz and Tilman 2005), and the invader would have an intermediate, niche-related trait value not represented by the native community. However, the average trait difference between the invader and native species in Fig. 1b will be smaller than the difference among native species, and under the criterion proposed, the native-invasive trait difference would be considered unimportant. Thus, the criterion proposed by Leffler et al. (2014) cannot distinguish between cases where trait values may lie between those of native species but are still distinct and cases where they are very similar to native species. Exotic species may not only invade a community by having different niche-related traits compared to native species. Some of the traits considered in the metaanalysis of Leffler et al. (2014), e.g., biomass, are arguably traits related to fitness. Such fitness-related traits also do not have to be more different between invasive and native species than among natives, for them to be important for invasions. All that is required is for the trait difference to be large enough for invasive species to have greater fitness than the native species (Fig. 1c). If this occurs and there is niche overlap between the invasive species and a native species, then the invasive species should displace the native species (MacDougall et al. 2009). The trait difference between invasive and native species should always be greater than the average native-native difference only when the trait is related to niche space and the invader is occupying a vacant niche at the extremes of the niche space available to the whole community. Thus, cases that meet the Leffler et al. (2014) criterion could be viewed as representing only one of three possible scenarios where differences in traits between native and invasive species are potentially important, and the only scenario where native-native differences are relevant. The challenge is to understand which of the many traits we can measure are actually related to fitness and niches of invasive and native species, and then to identify whether fitness or niche differences (or perhaps even both) have led to invasion. Manuscript received 15 July 2014; revised 7 August 2014; accepted 10 September 2014. Corresponding Editor: D. C. Laughlin. 1 Ecology Lab, Department of Biology, University of Konstanz, Universitaetsstrassse 10, Konstanz D78457 Germany. 2 E-mail: wayne.dawson@uni-konstanz.de

Plant responses to changes in temperature can be a key factor in predicting the presence and managing invasive plant species while conserving resident native plant species in dryland ecosystems. Climate can influence germination, establishment, and seedling biomass of both native and invasive plant species. We tested the hypothesis that common and widely distributed native and an invasive plant species in dryland ecosystems in California respond differently to increasing temperatures. To test this, we examined the effects of temperature variation on germination, establishment, and per capita seedling biomass of three native and one invasive plant species (Bromus rubens) in independent 6 week growth trial experiments in a controlled greenhouse. Higher relative temperatures reduced the germination and establishment of the tested invasive species and two tested native species, however, per capita biomass was not significantly affected. Specifically, germination and establishment of the invasive species B. rubens and the native species Phacelia tanacetifolia was significantly reduced. This invasive species can often outcompete natives, but increasing temperature could potentially shift the balance between the germination and establishment of natives. A warming climate will likely have negative impacts on native annual plant species in California tested here because increasing temperatures can co-occur with drought. This study shows that our tested native annual plant species tested here have some resilience to relatively significant increases in temperature, and this can favor at least one native species relative to at least one highly noxious invasive plant species.

Trait divergence and the ecosystem impacts of invading species

Invasive plant species are major drivers of biodiversity losses, especially on islands which are prone to invasions and extinctions. In the “endemic montane forest” of Robinson Crusoe Island (Pacific Ocean, Chile) invasive exotic plant species threaten conservation efforts by establishing in gaps and outcompeting native tree species regeneration. We compared gap attributes and ground vegetation cover in three gap types: those dominated by native species ( 30 % cover by invasive species), and treated gaps (invasive species removed). We examined (a) which gap attributes favored native and exotic species, (b) the relationship between gap size and species richness, and (c) species responses to invasion and treatment. Gaps ranged in size from 46 to 777 m2 caused mainly by uprooted and snapped trees. Multi response permutation procedures showed a different floristic composition between natural, invaded and treated gaps. The presence of Myrceugeniafernandeziana (native species) and Aristotelia chilensis (invasive species) as gap border trees was positively and negatively correlated with native species richness, respectively. New gaps had more native species than old gaps, and smaller gaps contained relatively more native species than larger ones. An increase in invasive species cover was related to a decline in native species cover and richness. 1–6 years after treatment gaps tended to recover their native floristic composition. Highly effective conservation management programs will concentrate on monitoring gap creation, early control of invasive species, and by treating smaller gaps first.

Habitat fragmentation differentially affects invasive and native plant diversity in a human-dominated wetland island system

Biological invasion has attracted widespread attention because invasive species threaten native biodiversity and weaken ecosystem services. Based on field investigation of vegetation in Nujiang River valley, Northwest Yunnan, we analyzed the spatial patterns of native and invasive species richness, and the effects of topography, climate, and roadside habitat disturbance on the invasive versus native plant species richness. We recorded 26 exotic invasive plant species that belong to 13 families and 21 genera, and 1,145 native plant species, belonging to 158 families and 628 genera. Along the Nujiang River valley, species richness of invasive plants decreased with increasing latitude and altitude, while species richness of native plants increased with increasing latitude, and showed a hump-shaped pattern with elevation. A generalized linear model was used to estimate the roles of roadside disturbance, climate, topography and soil nutrients on the distribution of both native and invasive species richness. Results of hierarchical variation partitioning revealed that roadside habitat disturbance had primary impact on the distribution of two groups of species. Pre·研究报告· 390 生 物 多 样 性 Biodiversity Science 第 24 卷 cipitation was the climatic determinant of invasive species diversity, and small-scale topographic factors, especially aspect, mainly affected native species diversity. It is likely that native species became drought-resistant in the evolutionary process while invasive species failed to adapt themselves to the local arid environments due to the short colonization time. This research supports the hypothesis that resource availability is the main factor limiting plant invasion, and highlights the negative effects of human activity on biodiversity. In addition, results of structural equation modelling revealed that native communities aren’t resistant to plant invasion. The negative relationship between invasive and native species richness reflects the different responses of the two group species to environmental factors.

The idea that dominant invasive plant species outperform neighboring native species through higher rates of carbon assimilation and growth is supported by several analyses of global data sets. However, theory suggests that native and invasive species occurring in low-resource environments will be functionally similar, as environmental factors restrict the range of observed physiological and morphological trait values. We measured resource-use traits in native and invasive plant species across eight diverse vegetation communities distributed throughout the five mediterranean-climate regions, which are drought prone and increasingly threatened by human activities, including the introduction of exotic species. Traits differed strongly across the five regions. In regions with functional differences between native and invasive species groups, invasive species displayed traits consistent with high resource acquisition; however, these patterns were largely attributable to differences in life form. We found that species invading mediterranean-climate regions were more likely to be annual than perennial: three of the five regions were dominated by native woody species and invasive annuals. These results suggest that trait differences between native and invasive species are context dependent and will vary across vegetation communities. Native and invasive species within annual and perennial groups had similar patterns of carbon assimilation and resource use, which contradicts the widespread idea that invasive species optimize resource acquisition rather than resource conservation. .

Effects of co-occurring non-native invasive plant species on old-field succession

Diversified grasslands that contain native plant species are being recognized as important elements of agricultural landscapes and for production of biofuel feedstocks as well as a variety of other ecosystem services. Unfortunately, establishment of such grasslands is often difficult, unpredictable, and highly vulnerable to interference and invasion by weeds. Evidence suggests that soil-microbial “legacies” of invasive perennial species can inhibit growth of native grassland species. However, previous assessments of legacy effects of soil occupancy by invasive species that invade grasslands have focused on single invasive species and on responses to invasive soil occupancy in only a few species. In this study, we tested the hypothesis that legacy effects of invasive species differ qualitatively from those of native grassland species. In a glasshouse, three invasive and three native grassland perennials and a native perennial mixture were grown separately through three cycles of growth and soil conditioning in soils with and without arbuscular mycorrhizal fungi (AMF), after which we assessed seedling growth in these soils. Native species differed categorically from invasives in their response to soil conditioning by native or invasive species, but these differences depended on the presence of AMF. When AMF were present, native species largely had facilitative effects on invasive species, relative to effects of invasives on other invasives. Invasive species did not facilitate native growth; neutral effects were predominant, but strong soil-mediated inhibitory effects on certain native species occurred. Our results support the hypothesis that successful plant invaders create biological legacies in soil that inhibit native growth, but suggest also this mechanism of invasion will have nuanced effects on community dynamics, as some natives may be unaffected by such legacies. Such native species may be valuable as nurse plants that provide cost-effective restoration of soil conditions needed for efficient establishment of diversified grasslands.

The effects of biological invasions are most evident in isolated oceanic islands such as the Hawaiian Archipelago, where invasive plant species are rapidly changing the composition and function of plant communities. In this study, we compared the specific leaf area (SLA), leaf tissue construction cost (CC), leaf nutrient concentration, and net CO2 assimilation (A) of 83 populations of 34 native and 30 invasive species spanning elevation and substrate age gradients on Mauna Loa volcano in the island of Hawaii. In this complex environmental matrix, where annual precipitation is higher than 1500 mm, we predicted that invasive species, as a group, will have leaf traits, such as higher SLA and A and lower leaf CC, which may result in more efficient capture of limiting resources (use more resources at a lower carbon cost) than native species. Overall, invasive species had higher SLA and A, and lower CC than native species, consistent with our prediction. SLA and foliar N and P were 22.5%, 30.5%, and 37.5% higher, respectively, in invasive species compared to native ones. Light-saturated photosynthesis was higher for invasive species (9.59 μmol m-2 s-1) than for native species (7.31 μmol m-2 s-1), and the difference was larger when A was expressed on a mass basis. Leaf construction costs, on the other hand, were lower for the invasive species (1.33 equivalents of glucose g-1) than for native species (1.37). This difference was larger when CC was expressed on an area basis. The trends in the above traits were maintained when groups of ecologically equivalent native and invasive species (i.e., sharing similar life history traits and growing in the same habitat) were compared. Foliar N and P were significantly higher in invasive species across all growth forms. Higher N may partially explain the higher A of invasive species. Despite relatively high N, the photosynthetic nitrogen use efficiency of invasive species was 15% higher than that of native species. These results suggest that invasive species may not only use resources more efficiently than native species, but may potentially demonstrate higher growth rates, consistent with their rapid spread in isolated oceanic islands.

Invasive, non-native plant species often outcompete native species and reduce biodiversity. Invasive plants frequently begin growth before native plants, yet few studies have examined whether invasives win in competition partly by colonizing disturbed sites more quickly or by beginning growth earlier in the season than native plants (i.e. due to priority effects). We hypothesized that invasive plant species would benefit more from priority effects than would comparable native species and that earlier growth of invasive species would decrease plant biodiversity. To test this hypothesis, we grew three pairs of invasive and native plant species from three different functional groups/plant families (C3 grasses/Poaceae, non-leguminous forbs/Asteraceae, and legumes/Fabaceae). We seeded each of the species 3 weeks before seeding the other five species into large pots in a greenhouse. Consistent with our hypothesis, we found much stronger priority effects with invasive than native species. Each invasive species formed a near-monocultures when seeded first (97.5 % of total biomass, on average) whereas native species did not similarly dominate (29.8 % of total biomass, on average). Similarly, Simpson’s species diversity was 81 % higher when the initially sown species was native rather than invasive. The literature suggests that invasive species in the field often begin growth earlier in the spring than native species and that climate change may increasingly allow invasives to begin growth before native species, indicating invasive priority effects may become increasingly common.

Invasive species may outperform native species by acquiring more resources or by efficiently using limited resources. Studies comparing leaf traits as a metric of carbon capture strategies in native and invasive species have come to different conclusions. Some studies suggest that invasive species are better at acquiring resources, but that native and invasive species use resources similarly. Other studies have found that native and invasive species differ in resource use efficiency, which implies different biochemical or physiological mechanisms of carbon capture. To resolve this debate, we examined relationships among four leaf traits (photosynthetic rate, specific leaf area, foliar nitrogen, foliar phosphorus) in co-occurring native and invasive species from eight plant communities across five Mediterranean-climate ecosystems. We performed standardized major axis regression for all trait combinations within and across sites, testing for slope homogeneity and shifts in elevation (y-intercept) or along a common slope between species groups. Across the global dataset, native and invasive species had similar carbon capture strategies (i.e., similar slopes), with invasive species occupying a position of greater resource acquisition. However, these patterns did not hold when regions were analyzed individually. Regional differences may be driven by differences in life form between native and invasive species, and variation in soil resource availability among regions. Our context-dependent results reveal not only that management of invasive species will differ across regions but also that global comparisons of invasive and native species can be misleading.

Patterns of fruiting phenology in temperate ecosystems are poorly understood, despite the ecological importance of fruiting for animal nutrition and seed dispersal. Herbarium specimens represent an under-utilized resource for investigating geographical and climatic factors affecting fruiting times within species, patterns in fruiting times among species, and differences between native and non-native invasive species. We examined over 15,000 herbarium specimens, collected and housed across New England, and found 3159 specimens with ripe fruits, collected from 1849-2013. We examined patterns in fruiting phenology among 37 native and 18 invasive woody plant species common to New England. We compared fruiting dates between native and invasive species, and analyzed how fruiting phenology varies with temperature, space, and time. Spring temperature and year explained a small but significant amount of the variation in fruiting dates. Accounting for the moderate phylogenetic signal in fruiting phenology, invasive species fruited 26 days later on average than native species, with significantly greater standard deviations. Herbarium specimens can be used to detect patterns in fruiting times among species. However,the amount of intraspecific variation in fruiting times explained by temporal, geographic, and climatic predictors is small, due to a combination of low temporal resolution of fruiting specimens and the protracted nature of fruiting.Later fruiting times in invasive species, combined with delays in autumn bird migrations in New England, may increase the likelihood that migratory birds will consume and disperse invasive seeds in New England later into the year.

An invasive and native plant differ in their effects on the soil food-web and plant-soil phosphorus cycle

Invasive plants are a major cause of the global biodiversity crisis; it is therefore crucial to understand mechanisms that contribute to their success. South‐western Australia is a global biodiversity hotspot with extremely low soil phosphorus (P) concentrations. In this region, a large proportion of native plant species release carboxylates that mobilise soil P. Many widespread invasive plant species in south‐western Australia are arbuscular mycorrhizal (AM). We hypothesised that some of these invasive AM plant species exhibit similar P‐acquisition strategies as native carboxylate‐releasing P‐mobilising species which allows them to thrive in P‐impoverished soils. To test this hypothesis, we collected 23 common invasive species in the field and assessed their leaf manganese concentration [Mn], relative to that of native reference species at different locations, as a proxy for carboxylate release. In addition, we cultivated seven of the invasive species in hydroponics at different P supply to measure their root carboxylate exudation. Furthermore, we measured leaf P concentration and photosynthetic P‐use efficiency (PPUE) of five invasive species in the glasshouse. In the field investigation, almost all invasive species exhibited significantly higher leaf [Mn] than the negative references, which do not release carboxylates, indicating carboxylate release of the invasive plants. Leaf [Mn] of a few invasives even exceeded that of positive references, which exhibit significant carboxylate release, indicating substantial carboxylate release of these invasives. All glasshouse‐grown invasive species with high field leaf [Mn] released root carboxylates under low P supply. Most of the tested invasive plant species also exhibited greater PPUE than native plants under low P supply. Invasive AM plant species exhibited root exudation of carboxylates as a P‐acquisition strategy, which very likely allows their successful invasion of severely P‐impoverished habitats. Read the free Plain Language Summary for this article on the Journal blog.