Multiple Non-native Species Research Articles

Functional diversity does not explain the co-occurrence of non-native species within a flow-modified African river system.

Globally, there is growing concern on the occurrence of multiple non-native species within invaded habitats. Proliferation of multiple non-native species together with anthropogenic-driven habitat modifications raise questions on the mechanisms facilitating the co-occurrence of these species and their potential impact within the recipient systems. Using the Great Fish River system (South Africa) which is anthropogenically-modified by inter-basin water transfer (IBWT), as a case study, this research employed trait-based approaches to explore patterns associated with the co-occurrence of multiple non-native fish species. This was achieved by investigating the role of functional diversity of non-native and native fishes in relation to their composition, distribution and environmental relationships. Nineteen functional traits that defined two broad ecological attributes (habitat use and feeding) were determined for 13 fish species that comprised eight native and five non-native fishes. We used these data to, firstly, evaluate functional diversity patterns and to compare functional traits of native and non-native fishes in the Great Fish River system. Secondly, we employed multivariate ordination analyses (factor analysis, RLQ and fourth-corner analyses) to investigate interspecific trait variations and potential species-trait-environmental relationships. From a functional diversity perspective, there were no significant differences in most functional diversity indices between native and non-native species. Despite interspecific variation in body morphology-related traits, we also found no clear separation between native and non-native species based on the ordination analysis of the functional traits. Furthermore, while RLQ ordination showed broad spatial patterns, the fourth-corner analyses revealed no significant relationships among species distribution, functional traits and environmental variables. The weak species-trait-environment relationship observed in this study suggests that environmental filtering was likely a poor determinant of functional trait structure within the Great Fish River. Modification of the natural flow regime may have weakened the relationship between species traits and the environment as has been shown in other systems.

Timing of invasive species removal influences nonnative biotic resistance and trajectories of community reassembly

Abstract As biological invasions increasingly threaten biodiversity, the removal of invasive nonnative species emerges as a possibility to recover the structure and function of native communities. Yet, we have limited knowledge of how communities assemble after nonnative removals. Since most ecosystems are invaded by multiple nonnative species, the impact of their removal likely depends on the interactions among nonnative species which, in turn, are contingent on the environmental context in which they occur. We evaluated the community assembly after the targeted removal of two highly invasive shrubs, Sweetbriar rose (Rosa rubiginosa) and Scotch broom (Cytisus scoparius). The removal was performed at two different times in the growing season (early or late removal) in field and mesocosm communities. In search of general patterns across species, we modelled species responses as a function of their origin (i.e. native/nonnative) and functional traits. We found evidence for negative and asymmetric interactions between dominant invasive species that translated into changes in the abundances of the rest of the species in the community. Depending on the identity of the removed species, the removal of invasive species affected community assembly by promoting other nonnative species or hindering the performance of native species. These effects were modulated by the timing of removal and did not depend on leaf or seed traits. Synthesis. Accounting for nonnative interactions and their temporal dependency should improve our inferences about assembly processes and the effectiveness of nonnative removal aimed at reducing the accumulation of nonnatives.

Fluctuations in resource availability shape the competitive balance among non-native plant species.

Fluctuating resource availability plays a critical role in determining non-native plant invasions through mediating the competitive balance between non-native and native species. However, the impact of fluctuating resource availability on interactions among non-native species remains largely unknown. This represents a barrier to understanding invasion mechanisms, particularly in habitats that harbor multiple non-native species with different responses to fluctuating resource availability. To examine the responses of non-native plant species to nutrient fluctuations, we compared the growth of each of 12 non-native species found to be common in local natural areas to nutrients supplied at a constant rate or supplied as a single large pulse in a pot experiment. We found that seven species produced more biomass with pulsed nutrients compared to constant nutrients (hereafter "benefitting species"), while the other five species did not differ between nutrient enrichment treatments (hereafter "non-benefitting species"). To investigate how nutrient fluctuations influence the interactions among non-native plant species, we established experimental non-native communities in the field with two benefitting and two non-benefitting non-native species. Compared with constant nutrient supply, the single large pulse of nutrient did not influence community biomass, but strongly increased the biomass and cover of the benefitting species and decreased those of the non-benefitting species. Furthermore, the benefitting species had higher leaf N content and greater plant height when nutrients were supplied as a single large pulse than at a constant rate, whereas the non-benefitting species showed no differences in leaf N content and were shorter when nutrients were supplied as a single large pulse than at a constant rate. Our results add to the growing evidence that the individual responses of non-native species to nutrient fluctuation are species-specific. More importantly, benefitting species were favored by nutrients coming in a pulse, while non-benefitting ones were favored by nutrients coming constantly when they grew together. This suggests that nutrient fluctuations can mediate the competitive balance among non-native plants and may thus determine their invasion success in a community harboring multiple non-native plant species.

Synergistic impacts of co‐occurring invasive grasses cause persistent effects in the soil‐plant system after selective removal

AbstractHuman 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.

Invasional meltdown: an experimental test and a framework to distinguish synergistic, additive, and antagonistic effects

The potential role of positive interactions among co-invaders is at the core of the invasional meltdown hypothesis. The interaction of non-native species could result in an exacerbation of each other’s effects. Thus, the resulting effect of multiple non-native species on ecosystems can be greater than the sum of their individual effects. We designed an analytical framework and a set of mesocosm experiments to assess the potential synergistic effects of three non-native species (Limnoperna fortunei, Astronotus crassipinnis, and Hydrilla verticillata) in a highly invaded floodplain in southern Brazil. We analyzed ecosystem, community, and population attributes in scenarios with non-natives. Our hypothesis of a synergistic effect was not supported. Even though effects of the invasive species were detected for all ecological levels, evidence indicated that these effects were additive. In addition to adding to the statement that origin (i.e., native vs. non-native status) does matter, we provide a tool to differentiate additive, synergistic, and antagonistic effects in situations with multiple invasions, and experimentally demonstrate additive effects of non-native species at different ecological levels.

Invasive non-native plants have a greater effect on neighbouring natives than other non-natives.

Human activity is creating a global footprint by changing the climate, altering habitats and reshuffling the distribution of species. The movement of species around the globe has led to the naturalization and accumulation of multiple non-native species within ecosystems, which is frequently associated with habitat disturbance and changing environmental conditions. However, interactions among species will also influence community composition, but little is known about the full range of direct and indirect interactions among native and non-native species. Here, we show through a meta-analysis of 1,215 pairwise plant interactions between 274 vascular plant species in 21 major habitat types that interactions between non-native plants are asymmetrical with interactions between non-native and native plants. Non-native plants were always bad neighbours, but the negative effect of non-natives on natives was around two times greater than the effect of non-natives on other non-natives. In contrast, the performance of non-native plants was five times higher in the presence of a neighbouring native plant species than in the presence of a neighbouring non-native plant species. Together, these results demonstrate that invaded plant communities may accumulate additional non-native species even if direct interactions between non-natives species are negative. Put another way, invasions may be more likely to lead to more invasions, requiring more active management of ecosystems by promoting native species restoration to undermine invasive positive feedback and to assist native species recovery in invaded ecosystems.

ALS-linked misfolded SOD1 species have divergent impacts on mitochondria.

Approximately 20 % of familial Amyotrophic Lateral Sclerosis (ALS) is caused by mutations in superoxide dismutase (SOD1), which leads to misfolding of the SOD1 protein, resulting in a toxic gain of function. Several conformation-restricted antibodies have been generated that specifically recognize misfolded SOD1 protein, and have been used as therapeutics in pre-clinical models. Misfolded SOD1 selectively associates with spinal cord mitochondria in SOD1 rodent models. Using the SOD1G93A rat model, we find that SOD1 conformational specific antibodies AMF7-63 and DSE2-3H1 labeled a fibrillar network concentrated in the anterior horn; while A5C3, B8H10, C4F6 and D3H5 labeled motor neurons as well as puncta in the neuropil. There is a time-dependent accumulation of misfolded SOD1 at the surface of spinal cord mitochondria with AMF7-63-labeled mitochondria having increased volume in contrast to a mitochondrial subset labeled with B8H10. In spinal cord homogenates and isolated mitochondria, AMF7-63, DSE2-3H1 and B8H10 detect misfolded SOD1 aggregates. SOD1 that lacks its metal cofactors has an increased affinity for naïve mitochondria and misfolded SOD1 antibodies B8H10 and DSE2-3H1 readily detect demetalated mutant and wild-type SOD1. Together, these data suggest that multiple non-native species of misfolded SOD1 may exist, some of which are associated with mitochondrial damage. Conformational antibodies are invaluable tools to identify and characterize the variation in misfolded SOD1 species with regards to biochemical characteristics and toxicity. This information is highly relevant to the further development of these reagents as therapeutics.Electronic supplementary materialThe online version of this article (doi:10.1186/s40478-016-0313-8) contains supplementary material, which is available to authorized users.

Walleye Foraging Ecology in an Interconnected Chain of Lakes Influenced by Nonnative Species

AbstractThe Laurentian Great Lakes are a source of nonnative species for smaller inland lakes throughout the region; however, the foraging ecology of predators in smaller systems invaded by nonnative species has not been well studied. We used diet and stable isotope analyses to describe the contributions of native and nonnative forage species to the diets of Walleyes Sander vitreus within an inland lake chain invaded by multiple nonnative species. Our hypothesis was that the Walleyes' diet would be associated with the relative amount of prey available in littoral or offshore–pelagic habitats and would ultimately be linked to the dominant habitat type. We found that Walleyes in the study lakes exhibited flexibility in their dietary inclusion of nonnative prey species. Similar to predators in the Laurentian Great Lakes, Walleyes in smaller inland lakes integrated littoral nonnative species into their diets. However, Walleyes unexpectedly showed limited usage of accessible nonnative pelagic forage. We suggest that predator responses to species introductions are likely context dependent and warrant further investigation in multiple systems and with multiple predators to determine the full extent of nonnative species' integration into the food web and their influence on native populations.Received June 27, 2015; accepted November 2, 2015

Status and management of non-native plant invasion in three of the largest national parks in the United States

Globally, invasion by non-native plants threatens resources that nature reserves are designated to protect. We assessed the status of non-native plant invasion on 1,662, 0.1-ha plots in Death Valley National Park, Mojave National Preserve, and Lake Mead National Recreation Area. These parks comprise 2.5 million ha, 23% of the national park land in the contiguous USA. At least one non-native species inhabited 82% of plots. Thirty-one percent of plots contained one non-native species, 30% two, 17% three, and 4% four to ten non-native species. Red brome (Bromus rubens), an ‘ecosystem engineer’ that alters fire regimes, was most widespread, infesting 60% of plots. By identifying frequency of species through this assessment, early detection and treatment can target infrequent species or minimally invaded sites, while containment strategies could focus on established invaders. We further compared two existing systems for prioritizing species for management and found that a third of species on plots had no rankings available. Moreover, rankings did not always agree between ranking systems for species that were ranked. Presence of multiple non-native species complicates treatment, and while we found that 40% of plots contained both forb and grass invaders, exploiting accelerated phenology of non-natives (compared to native annuals) might help manage multi-species invasions. Large sizes of these parks and scale of invasion are formidable challenges for management. Yet, precisely because of their size, these reserves represent opportunities to conserve large landscapes of native species by managing non-native plant invasions.

Above- and below-ground effects of plant diversity depend on species origin: an experimental test with multiple invaders.

Although many plant communities are invaded by multiple nonnative species, we have limited information on how a species' origin affects ecosystem function. We tested how differences in species richness and origin affect productivity and seedling establishment. We created phylogenetically paired native and nonnative plant communities in a glasshouse experiment to test diversity-productivity relationships and responsible mechanisms (i.e. selection or complementarity effects). Additionally, we tested how productivity and associated mechanisms influenced seedling establishment. We used diversity-interaction models to describe how species' interactions influenced diversity-productivity relationships. Communities with more species had higher total biomass than did monoculture communities, but native and nonnative communities diverged in root:shoot ratios and the mechanism responsible for increased productivity: positive selection effect in nonnative communities and positive complementarity effect in native communities. Seedling establishment was 46% lower in nonnative than in native communities and was correlated with the average selection effect. Interspecific interactions contributed to productivity patterns, but the specific types of interactions differed between native and nonnative communities. These results reinforce findings that the diversity-productivity mechanisms in native and nonnative communities differ and are the first to show that these mechanisms can influence seedling establishment and that different types of interactions influence diversity-productivity relationships.

Predicted Redistribution of Ceratomyxa shasta Genotypes with Salmonid Passage in the Deschutes River, Oregon

A series of dams on the Deschutes River, Oregon, act as migration barriers that segregate the river system into upper and lower basins. Proposed fish passage between basins would reunite populations of native potamodromous fish and allow anadromous fish of Deschutes River origin access to the upper basin. We assessed the potential redistribution of host-species-specific genotypes (O, I, II, III) of the myxozoan parasite Ceratomyxa shasta that could occur with fish passage and examined the influence of nonnative fish on genotype composition. To determine the present distribution of the parasite genotypes, we exposed eight salmonid species-three native and five stocked for sport fishing-in present and predicted anadromous salmonid habitats. We monitored fish for infection by C. shasta and sequenced a section of the parasite ribosomal DNA gene from fish and water samples to determine parasite genotype. Genotype O was present in both upper and lower basins and detected only in steelhead Oncorhynchus mykiss. Genotype I was spatially limited to the lower basin, isolated predominantly from Chinook salmon O. tshawytscha, and lethal for this species only. Genotype II was detected in both basins and in multiple species, but only as a minor component of the infection. Genotype III was also present in both basins, had a wide host range, and caused mortality in native steelhead and multiple nonnative species. Atlantic salmon Salmo salar and kokanee O. nerka were the least susceptible to infection by any genotype of C. shasta. Our findings confirmed the host-specific patterns of C. shasta infections and indicated that passage of Chinook salmon would probably spread genotype I into the upper Deschutes River basin, but with little risk to native salmonid populations.

Community ecology of invasions: direct and indirect effects of multiple invasive species on aquatic communities

With many ecosystems now supporting multiple nonnative species from different trophic levels, it can be challenging to disentangle the net effects of invaders within a community context. Here, we combined wetland surveys with a mesocosm experiment to examine the individual and combined effects of nonnative fish predators and nonnative bullfrogs on aquatic communities. Among 139 wetlands, nonnative fish (bass, sunfish, and mosquitofish) negatively influenced the probability of occupancy of Pacific treefrogs (Pseudacris regilla), but neither invader correlated strongly with occupancy by California newts (Taricha torosa), western toads (Anaxyrus boreas), or California red-legged frogs (Rana draytonii). In mesocosms, mosquitofish dramatically reduced the abundance of zooplankton and palatable amphibian larvae (P. regilla and T. torosa), leading to increases in nutrient concentrations and phytoplankton (through loss of zooplankton), and rapid growth of unpalatable toad larvae (through competitive release). Bullfrog larvae reduced the growth of native anurans but had no effect on survival. Despite strong effects on natives, invaders did not negatively influence one another, and their combined effects were additive. Our results highlight how the net effects of multiple nonnative species depend on the trophic level of each invader, the form and magnitude of invader interactions, and the traits of native community members.

Characterization of Alkaline Transitions in Ferricytochrome c Using Carbon−Deuterium Infrared Probes

The alkaline-induced structural transitions of ferricytochrome c have been studied intensively as a model for how changes in metal ligation contribute to protein function and folding. Previous studies have demonstrated that multiple non-native species accumulate with increasing pH. Here, we used a combination of experiments and simulations to provide a high-resolution view of the changes associated with increasing alkaline conditions. Alkaline-induced transitions were characterized under equilibrium conditions by following changes in the IR absorptions of carbon-deuterium chromophores incorporated at Leu68, Lys72, Lys73, Lys79, and Met80. The data suggest that at least four intermediates are formed as the pH is increased prior to complete unfolding of the protein. The first alkaline transition observed appears to be driven by a single deprotonation and occurs with a midpoint of pH 8.8, but surprisingly, the intermediate formed does not appear to be one of the well-characterized lysine misligates. At higher pH, second and third deprotonations, with a combined apparent midpoint pH of 10.2, induce transitions to Lys73- or Lys79-misligated species. Interestingly, the lysine misligates appear to undergo iron reduction by the coordinated amine. A transition from the lysine misligates to another intermediate, likely a hydroxide-misligated species, is associated with a fourth deprotonation and a midpoint of pH 10.7. Finally, the protein loses tertiary structure with a fifth deprotonation that occurs with a midpoint of pH 12.7. Native topology-based models with enforced misligation are employed to help understand the structures of the observed intermediates.