Evaluation of non-opioid analgesic toxicity and their effects on native and invasive species of crustaceans (Gammaridae, Gammarus)

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

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.

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

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.

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.

1. Phenotypic plasticity is often cited as an important mechanism of plant invasion. However, few studies have evaluated the plasticity of a diverse set of traits among invasive and native species, particularly in low resource habitats, and none have examined the functional significance of these traits. 2. I explored trait plasticity in response to variation in light and nutrient availability in five phylogenetically related pairs of native and invasive species occurring in a nutrient-poor habitat. In addition to the magnitude of trait plasticity, I assessed the correlation between 16 leaf- and plant-level traits and plant performance, as measured by total plant biomass. Because plasticity for morphological and physiological traits is thought to be limited in low resource environments (where native species usually display traits associated with resource conservation), I predicted that native and invasive species would display similar, low levels of trait plasticity. 3. Across treatments, invasive and native species within pairs differed with respect to many of the traits measured; however, invasive species as a group did not show consistent patterns in the direction of trait values. Relative to native species, invasive species displayed high plasticity in traits pertaining to biomass partitioning and leaf-level nitrogen and light use, but only in response to nutrient availability. Invasive and native species showed similar levels of resource-use efficiency and there was no relationship between species plasticity and resource-use efficiency across species. 4. Traits associated with carbon fixation were strongly correlated with performance in invasive species while only a single resource conservation trait was strongly correlated with performance in multiple native species. Several highly plastic traits were not strongly correlated with performance which underscores the difficulty in assessing the functional significance of resource conservation traits over short timescales and calls into question the relevance of simple, quantitative assessments of trait plasticity. 5. Synthesis. My data support the idea that invasive species display high trait plasticity. The degree of plasticity observed here for species occurring in low resource systems corresponds with values observed in high resource systems, which contradicts the general paradigm that trait plasticity is constrained in low resource systems. Several traits were positively correlated with plant performance suggesting that trait plasticity will influence plant fitness.

Growth, biomass allocation, and photosynthetic characteristics of seedlings of five invasive non-indigenous and four native species grown under different light regimes were studied to help explain the success of invasive species in Hawaiian rainforests. Plants were grown under three greenhouse light levels representative of those found in the center and edge of gaps and in the understory of Hawaiian rainforests, and under an additional treatment with unaltered shade. Relative growth rates (RGRs) of invasive species grown in sun and partial shade were significantly higher than those for native species, averaging 0.25 and 0.17 g g-1 week-1, respectively, while native species averaged only 0.09 and 0.06 g g-1 week-1, respectively. The RGR of invasive species under the shade treatment was 40% higher than that of native species. Leaf area ratios (LARs) of sun and partial-shade-grown invasive and native species were similar but the LAR of invasive species in the shade was, on average, 20% higher than that of native species. There were no differences between invasive and native species in biomass allocation to shoots and roots, or in leaf mass per area across light environments. Light-saturated photosynthetic rates (Pmax) were higher for invasive species than for native species in all light treatments. Pmax of invasive species grown in the sun treatment, for example, ranged from 5.5 to 11.9 μmol m-2 s-1 as compared with 3.0-4.5 μmol m-2 s-1 for native species grown under similar light conditions. The slope of the linear relationship between Pmax and dark respiration was steeper for invasive than for native species, indicating that invasive species assimilate more CO2 at a lower respiratory cost than native species. These results suggest that the invasive species may have higher growth rates than the native species as a consequence of higher photosynthetic capacities under sun and partial shade, lower dark respiration under all light treatments, and higher LARs when growing under shade conditions. Overall, invasive species appear to be better suited than native species to capturing and utilizing light resources, particularly in high-light environments such as those characterized by relatively high levels of disturbance.

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.

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.

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

Invasive species are leading drivers of environmental change. Their impacts are often linked to their population size, but surprisingly little is known about how frequently they achieve high abundances. A nearly universal pattern in ecology is that species are rare in most locations and abundant in a few, generating right-skewed abundance distributions. Here, we use abundance data from over 24,000 populations of 17 invasive and 104 native aquatic species to test whether invasive species differ from native counterparts in statistical patterns of abundance across multiple sites. Invasive species on average reached significantly higher densities than native species and exhibited significantly higher variance. However, invasive and native species did not differ in terms of coefficient of variation, skewness, or kurtosis. Abundance distributions of all species were highly right skewed (skewness>0), meaning both invasive and native species occurred at low densities in most locations where they were present. The average abundance of invasive and native species was 6% and 2%, respectively, of the maximum abundance observed within a taxonomic group. The biological significance of the differences between invasive and native species depends on species-specific relationships between abundance and impact. Recognition of cross-site heterogeneity in population densities brings a new dimension to invasive species management, and may help to refine optimal prevention, containment, control, and eradication strategies.

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 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.

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.