Trait divergence and the ecosystem impacts of invading species

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

During community assembly, early arriving exotic species might suppress other species to a greater extent than do native species. Because most exotics were intentionally introduced, we hypothesize there was human selection on regeneration traits during introduction. This could have occurred at the across- or within-species level (e.g. during cultivar development). We tested these predictions by seeding a single species that was either native, exotic 'wild-type' (from their native range), or exotic 'cultivated' using 28 grassland species in a glasshouse experiment. Priority effects were assessed by measuring species' effect on establishment of species from a seed mix added 21 d later. Exotic species had higher germination and earlier emergence dates than native species, and differences were found in both 'wild' and 'cultivated' exotics. Exotic species reduced biomass and species diversity of later arriving species much more than native species, regardless of seed source. Results indicate that in situations in which priority effects are likely to be strong, effects will be greater when an exotic species arrives first than when a native species arrives first; and this difference is not merely a result of exotic species cultivation, but might be a general native-exotic difference that deserves further study.

Invited article: The impacts of non‐native species: a review of the British Ornithologists’ Union's Autumn 2008 Scientific Meeting

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

Invasive species have been identified as a major threat to native biodiversity and ecosystem functioning worldwide due to their superiority in spread and growth. Such superiority is explained by the invasional meltdown phenomena, which suggests that invasive species facilitate the establishment of more invasive species rather than native species by modifying the plant-soil feedback (PSF). We conducted a two-phase plant-soil feedback experiment using the native Prosopis cineraria and the invasive Prosopis juliflora in Oman. Firstly, we conditioned the soil by planting seedlings of native species, invasive species, native and invasive species "mixed", and unconditioned soil served as a control. Secondly, we tested the feedback of these four conditioned soil on the two species separately by measuring the productivity (total biomass) and the performance in the form of plant functional traits (plant height, specific leaf area (SLA), leaf nitrogen content (Nmass), leaf carbon content (Cmass) and specific root length (SRL) of native and invasive species as well as the nutrient availability in soil (soil organic carbon (SOC) and soil total nitrogen (STN)). We found that the native species produced more biomass, best performance, and higher SOC and STN when grown in soil conditioned by native species, additionally, it gave lower biomass, reduced performance, and lower SOC and STN when grown in the soil conditioned by invasive and mixed species. These results suggest negative PSF for native species and positive PSF for invasive species in the soil conditioned by invasive species, which can be considered as red flag concerning the restoration of P. cineraria as an important native species in Oman, as such positive PSF of the invasive species P. juliflora will inhibit the regeneration of P. cineraria.

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.

Human influence on the environment is so extensive that virtually all ecosystems on the planet are now affected by biological invasions. And, often, ecosystems are invaded by multiple co‐occurring non‐native species. Hence, it is important to understand the impacts these invasions are producing on biodiversity and ecosystem processes.Here, we present results of a 2‐year long field experiment where we tested the effects of co‐occurring invasive C4African grasses in a Cerrado area in central Brazil. We compared plant and arthropod communities, plant biomass, and soil nitrogen dynamics and soil chemical characteristics across five experimental treatments:Urochloa decumbensremoval;Melinis minutifloraremoval; bothU.decumbensandM.minutifloraremoval;U.decumbensandM.minutiflorainvaded plots; and uninvaded Cerrado. We hypothesized that selective removal of invasive grasses would have distinct effects on the native ecosystem structure and functioning. We expected that each invasive grass would produce a different type of impact on the native ecosystem and that their impacts would be synergistic when co‐occurring.Removal ofM.minutifloradoubled native plant diversity and biomass when compared to invaded plots, whereas removal ofU.decumbensdid not alter these parameters. Cerrado plots had four times more plant species than plots cleared of invasives. Removal of invasive grasses did not affect the species richness or community composition of soil epigeal fauna. Cerrado soils had lower fertility, organic matter content and pH than invaded soils. The effects were generally higher when both invasive grasses were removed, suggesting impacts were synergistic, butM.minutiflorahad greater effects on plants and soils thanU.decumbens. Both invasive species produced negative impacts, but a single species was the main driver. We also detected persistent effects of the invasive grass species on the ecosystem after 2 years of removal.We conclude that invasive species of the same functional group have similar types of effects in native ecosystems, but the magnitude of impact was largely dependent on invasive species biomass and cover. Where multiple invasive species are present, research and management of invaded ecosystems should tackle the interacting effects of co‐occurring invaders.

We compared 13 traits of invasive exotic, noninvasive exotic, and ecologically similar native species to determine if there are generalizable differences among these groups that relate to persistence and spread of exotic species in tallgrass prairie plant communities. When species were grouped as invasive (two species), noninvasive (five species), and native (six species), no differences were found for the suite of traits examined, likely because of the high variability within and between groups. However, when exotic species, regardless of invasiveness, were compared with the native species, specific leaf area was ca. 40% higher for the exotic species, a result that is consistent with that of other studies. This pattern was also observed for five of seven pairwise comparisons of exotic and native species with similar life history traits. In contrast, total end‐of‐season biomass was as much as three times higher for the native species in five of seven of the native‐exotic species pairs. For other traits, differences between exotic and native species were species‐specific and were generally more numerous for noninvasive than for invasive exotic species pair‐wise comparisons. Thus, contrary to predictions, exotic species capable of successfully invading tallgrass prairie did not differ considerably from native species in most traits related to resource utilization and carbon gain. Moreover, invasive exotic species, those capable of displacing native species and dominating a community, were not distinct for the observed traits from their native counterparts. These results indicate that other traits, such as the ability to respond to resource pulses or herbivory, may explain more effectively why certain invasive species are able to invade these communities aggressively.

Native plants do not benefit from arriving early, but invasives pay to arrive late

Taking the long view on the ecological effects of plant invasions.

ABSTRACTUnderstanding the ecological differences between native and invasive species is of considerable scientific and practical interest. We examined such differences between native and invasive inland fish species from the Iberian Peninsula in order to analyse the importance of phylogenetic correction and variability (in addition to central tendency). We collected 26 quantitative and qualitative variables on the ecology, life‐history traits and human use of the 69 inland fish species of the Iberian Peninsula, including native, invasive and migratory species. The taxonomic distribution of invasive fish species deviated significantly from world freshwater richness and in contrast to native species, invasive fish belongs to only five taxonomic orders but to a wide spectrum of families not native to the Iberian Peninsula. Because the life‐history traits were highly dependent on taxonomy, the results, with or without applying phylogenetic methods, differed and after accounting for phylogeny, invasive species displayed higher and wider latitude in general and a different reproductive season mainly among salmonids and cyprinids. Human use was also significantly different between native and invasive fish species and produced more variability in life‐history traits of invasive species and uneven taxonomic distribution because of the high diversity of species introduced. We show that accounting for taxonomy and studying variability in addition to central tendency is important in the comparison of life‐history traits between native and invasive species.

Invasive species may be released from consumption by their native herbivores in novel habitats and thereby experience higher fitness relative to native species. However, few studies have examined release from herbivory as a mechanism of invasion in oceanic island systems, which have experienced particularly high loss of native species due to the invasion of non-native animal and plant species. We surveyed putative defensive traits and leaf damage rates in 19 pairs of taxonomically related invasive and native species in Hawaii, representing a broad taxonomic diversity. Leaf damage by insects and pathogens was monitored in both wet and dry seasons. We found that native species had higher leaf damage rates than invasive species, but only during the dry season. However, damage rates across native and invasive species averaged only 2% of leaf area. Native species generally displayed high levels of structural defense (leaf toughness and leaf thickness, but not leaf trichome density) while native and invasive species displayed similar levels of chemical defenses (total phenolics). A defense index, which integrated all putative defense traits, was significantly higher for native species, suggesting that native species may allocate fewer resources to growth and reproduction than do invasive species. Thus, our data support the idea that invasive species allocate fewer resources to defense traits, allowing them to outperform native species through increased growth and reproduction. While strong impacts of herbivores on invasion are not supported by the low damage rates we observed on mature plants, population-level studies that monitor how herbivores influence recruitment, mortality, and competitive outcomes are needed to accurately address how herbivores influence invasion in Hawaii.

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.

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.

Diversified grasslands that contain native plant species can produce biofuels, support sustainable grazing systems, and produce other ecosystem services. However, ecosystem service production can be disrupted by invasion of exotic perennial plants, and these plants can have soil-microbial “legacies” that may interfere with establishment and maintenance of diversified grasslands even after effective management of the invasive species. The nature of such legacies is not well understood, but may involve suppression of mutualisms between native species and soil microbes. In this study, we tested the hypotheses that legacy effects of invasive species change colonization rates, diversity, and composition of arbuscular-mycorrhizal fungi (AMF) associated with seedlings of co-occurring invasive and native grassland species. In a glasshouse, experimental soils were conditioned by cultivating three invasive grassland perennials, three native grassland perennials, and a native perennial mixture. Each was grown separately through three cycles of growth, after which we used T-RFLP analysis to characterize AMF associations of seedlings of six native perennial and six invasive perennial species grown in these soils. Legacy effects of soil conditioning by invasive species did not affect AMF richness in seedling roots, but did affect AMF colonization rates and the taxonomic composition of mycorrhizal associations in seedling roots. Moreover, native species were more heavily colonized by AMF and roots of native species had greater AMF richness (number of AMF operational taxonomic units per seedling) than did invasive species. The invasive species used to condition soil in this experiment have been shown to have legacy effects on biomass of native seedlings, reducing their growth in this and a previous similar experiment. Therefore, our results suggest that successful plant invaders can have legacies that affect soil-microbial associations of native plants and that these effects can inhibit growth of native plant species in invaded communities.