Diminishing importance of elaiosomes for acacia seed removal in non-native ranges

Myrmecochorous plants produce seeds with lipid-rich appendages (elaiosomes) which act as a reward for seed-dispersing ants. Seed dispersal is important for exotic species, which often need to establish new mutualistic interactions in order to colonize new non-native habitats. However, little is known about the importance of elaiosomes for seed removal in many of their non-native ranges. We studied ant–seed interactions of elaiosome-bearing and elaiosome-removed seeds of the Australian trees Acacia dealbata and Acacia longifolia in order to assess the relative importance of elaiosomes for seed removal between their native (Australia) and non-native (Portugal) ranges. In Portugal, we also studied the co-occurring native plant species with myrmecochorous seeds, Pterospartum tridentatum and Ulex europaeus, across three contiguous levels of acacia invasion: control (i.e. no acacia), low, and high acacia tree density. Acacia seeds were successfully removed by ants in their non-native region by a diversified assemblage of ant species, even in sites where native plants interacted with only one specialized ant species. In the invaded range, diminishing relative importance of elaiosomes was associated with changes in the ant community due to acacia invasion, and for A. dealbata, ant species richness decreased with increasing acacia tree density. Although seed removal was high for both acacia species, the importance of elaiosomes was proportionally lower for A. dealbata in the non-native region. Native plant species experienced significant reductions in seed removal in areas highly invaded by acacia, identifying another mechanism of displacement of native plants by acacias.     

Indirect facilitation of an anuran invasion by non-native fishes

Positive interactions among non‐native species could greatly exacerbate the problem of invasions, but are poorly studied and our knowledge of their occurrence is mostly limited to plant‐pollinator and dispersal interactions. We found that invasion of bullfrogs is facilitated by the presence of co‐evolved non‐native fish, which increase tadpole survival by reducing predatory macroinvertebrate densities. Native dragonfly nymphs in Oregon, USA caused zero survival of bullfrog tadpoles in a replicated field experiment unless a non‐native sunfish was present to reduce dragonfly density. This pattern was also evident in pond surveys where the best predictors of bullfrog abundance were the presence of non‐native fish and bathymetry. This is the first experimental evidence of facilitation between two non‐native vertebrates and supports the invasional meltdown hypothesis. Such positive interactions among non‐native species have the potential to disrupt ecosystems by amplifying invasions, and our study shows they can occur via indirect mechanisms.     

Thermodynamics and kinetics of non-native interactions in protein folding: a single point mutant significantly stabilizes the N-terminal domain of L9 by modulating non-native interactions in the denatured state

Comparatively little is known about the role of non-native interactions in protein folding and their role in both folding and stability is controversial. We demonstrate that non-native electrostatic interactions involving specific residues in the denatured state can have a significant effect upon protein stability and can persist in the transition state for folding. Mutation of a single surface exposed residue, Lys12 to Met, in the N-terminal domain of the ribosomal protein L9 (NTL9), significantly increased the stability of the protein and led to faster folding. Structural and energetic studies of the wild-type and K12M mutant show that the 1.9 kcal mol(-1) increase in stability is not due to native state effects, but rather is caused by modulation of specific non-native electrostatic interactions in the denatured state. pH dependent stability measurements confirm that the increased stability of the K12M is due to the elimination of favorable non-native interactions in the denatured state. Kinetic studies show that the non-native electrostatic interactions involving K12 persist in the transition state. The analysis demonstrates that canonical Phi-values can arise from the disruption of non-native interactions as well as from the development of native interactions.     

Engineering a compact non-native state of intestinal fatty acid-binding protein

The last three C-terminal residues (129-131) of intestinal fatty acid-binding protein (IFABP) participate in four main-chain hydrogen bonds and two electrostatic interactions to sequentially distant backbone and side-chain atoms. To assess if these interactions are involved in the final adjustment of the tertiary structure during folding, we engineered an IFABP variant truncated at residue 128. An additional mutation, Trp-6-->Phe, was introduced to simplify the conformational analysis by optical methods. Although the changes were limited to a small region of the protein surface, they resulted in an IFABP with altered secondary and tertiary structure. Truncated IFABP retains some cooperativity, is monomeric, highly compact, and has the molecular dimensions and shape of the native protein. Our results indicated that residues 129-131 are part of a crucial conformational determinant in which several long-range interactions, essential for the acquisition of the native state, are established. This work suggests that carefully controlled truncation can populate equilibrium non-native states under physiological conditions. These non-native states hold a great promise as experimental models for protein folding.     

The role of non-native interactions in the folding of knotted proteins

Stochastic simulations of coarse-grained protein models are used to investigate the propensity to form knots in early stages of protein folding. The study is carried out comparatively for two homologous carbamoyltransferases, a natively-knotted N-acetylornithine carbamoyltransferase (AOTCase) and an unknotted ornithine carbamoyltransferase (OTCase). In addition, two different sets of pairwise amino acid interactions are considered: one promoting exclusively native interactions, and the other additionally including non-native quasi-chemical and electrostatic interactions. With the former model neither protein shows a propensity to form knots. With the additional non-native interactions, knotting propensity remains negligible for the natively-unknotted OTCase while for AOTCase it is much enhanced. Analysis of the trajectories suggests that the different entanglement of the two transcarbamylases follows from the tendency of the C-terminal to point away from (for OTCase) or approach and eventually thread (for AOTCase) other regions of partly-folded protein. The analysis of the OTCase/AOTCase pair clarifies that natively-knotted proteins can spontaneously knot during early folding stages and that non-native sequence-dependent interactions are important for promoting and disfavouring early knotting events.     

Energetics of the native and non-native states of the glycophorin transmembrane helix dimer

Using an implicit membrane model (IMM1), we examine whether the structure of the transmembrane domain of Glycophorin A (GpA) could be predicted based on energetic considerations alone. The energetics of native GpA shows that van der Waals interactions make the largest contribution to stability. Although specific electrostatic interactions are stabilizing, the overall electrostatic contribution is close to zero. The GXXXG motif contributes significantly to stability, but residues outside this motif contribute almost twice as much. To generate non-native states a global conformational search was done on two segments of GpA: an 18-residue peptide (GpA74-91) that is embedded in the membrane and a 29-residue peptide (GpA70-98) that has additional polar residues flanking the transmembrane region. Simulated annealing was done on a large number of conformations generated from parallel, antiparallel, left- and right-handed starting structures by rotating each helix at 20 degrees intervals around its helical axis. Several crossing points along the helix dimer were considered. For 18-residue parallel topology, an ensemble of native-like structures was found at the lowest effective energy region; the effective energy is lowest for a right-handed structure with an RMSD of 1.0 A from the solid-state NMR structure with correct orientation of the helices. For the 29-residue peptide, the effective energies of several left-handed structures were lower than that of the native, right-handed structure. This could be due to deficiencies in modeling the interactions between charged sidechains and/or omission of the sidechain entropy contribution to the free energy. For 18-residue antiparallel topology, both IMM1 and a Generalized Born model give effective energies that are lower than that of the native structure. In contrast, the Poisson-Boltzmann solvation model gives lower effective energy for the parallel topology, largely because the electrostatic solvation energy is more favorable for the parallel structure. IMM1 seems to underestimate the solvation free energy advantage when the CO and NH dipoles just outside the membrane are parallel. This highlights the importance of electrostatic interactions even when these are not obvious by looking at the structures.     

Increased abundance of native and non-native spiders with habitat fragmentation

Habitat fragmentation and invasive species often contribute to the decline of native taxa. Since the penetration of non‐native species into natural habitat may be facilitated by habitat fragmentation, it is important to examine how these two factors interact. Previous research documented that, in contrast to most other arthropod taxa, spiders increased in density and morphospecies richness with decreasing fragment area and increasing fragment age (time since insularization) in urban habitat fragments in San Diego County, California, USA. We tested whether a specific mechanism, an increase in non‐native species with fragmentation, is responsible for this pattern. We found that both native and non‐native taxa contributed to the pattern. Abundance of native spiders per pitfall trap sample increased significantly with decreasing fragment size (i.e. a negative density–area relationship) and abundance of non‐natives increased significantly with increasing fragment age. The proportion of non‐native individuals also increased significantly with age. One non‐native species, Oecobius navus, comprised the majority of non‐native individuals (82.2%) and a significant proportion of total individuals (25.1%). Richness of spider families per sample (family density) increased with fragment age due to an increase in the occurrence of non‐natives in older fragments, however, native family richness did not vary with age or area. Due to increasing dominance by non‐native and some native families, family evenness declined with decreasing fragment size and increasing fragment age. Native and non‐native abundance covaried positively arguing against strong negative interactions between the two groups. O. navus had a strong positive association with another common non‐native arthropod, the Argentine ant (Linepitheme humile), suggesting a possible direct interaction. In contrast, abundance of native spiders was negatively correlated with Argentine ant abundance. We hypothesize that fragmentation in this semiarid habitat increases productivity in smaller and older fragments enhancing the density of both native and non‐native taxa.     

The effect of increasing the stability of non-native interactions on the folding landscape of the bacterial immunity protein Im9

How stabilising non-native interactions influence protein folding energy landscapes is currently not well understood: such interactions could speed folding by reducing the conformational search to the native state, or could slow folding by increasing ruggedness. Here, we examine the influence of non-native interactions in the folding process of the bacterial immunity protein Im9, by exploiting our ability to manipulate the stability of the intermediate and rate-limiting transition state (TS) in the folding of this protein by minor alteration of its sequence or changes in solvent conditions. By analysing the properties of these species using Phi-value analysis, and exploration of the structural properties of the TS ensemble using molecular dynamics simulations, we demonstrate the importance of non-native interactions in immunity protein folding and demonstrate that the rate-limiting step involves partial reorganisation of these interactions as the TS ensemble is traversed. Moreover, we show that increasing the contribution to stability made by non-native interactions results in an increase in Phi-values of the TS ensemble without altering its structural properties or solvent-accessible surface area. The data suggest that the immunity proteins fold on multiple, but closely related, micropathways, resulting in a heterogeneous TS ensemble that responds subtly to mutation or changes in the solvent conditions. Thus, altering the relative strength of native and non-native interactions influences the search to the native state by restricting the pathways through the folding energy landscape.     

Interguild interactions between a non-native phloem feeder and a native outbreaking defoliator

1. Competitive and synergistic interactions directly or indirectly drive community dynamics of herbivorous insects. Novel interactions between non-native and native insects are unpredictable and not fully understood. 2. We used manipulative experiments on mature red spruce trees to test interactions between a non-native phloem feeding insect, the brown spruce longhorn beetle (BSLB), and an outbreaking native defoliator, the spruce budworm. We subjected treatment trees to defoliation by three densities of spruce budworm larvae. Treatment trees were: stressed by (i) girdling (to mimic beetle feeding) or (ii) girdling + BSLB before spruce budworm larvae were introduced on branches in sleeve cages. Budworm larvae then fed on foliage and developed to pupation. We assessed all branches for budworm performance, defoliation, shoot production and shoot growth. 3. Shoot length did not differ in response to stress from girdling or BSLB infestation. Neither stress from girdling, nor interactions with BSLB feeding affected spruce budworm performance or defoliation. Intraspecific impacts on performance and defoliation in relation to budworm density were stronger than the effects of tree stress. 4. Prior infestation of red spruce by BSLB in our experimental set-up did not influence spruce budworm performance. BSLB is a successful invader that has blended into its novel ecological niche because of ecological and phylogenetic similarities with a native congener, Tetropium cinnamopterum. 5. Outbreaks by BSLB will not likely impede or facilitate spruce budworm outbreaks if they co-occur. It would be useful to evaluate the reverse scenario of BSLB success after defoliation stress by spruce budworm.     

Dominance of an invasive earthworm in native and non-native grassland ecosystems

More attention is currently being focused on earthworm invasions; however, in many ecosystems the relative abundance of native and invasive earthworm species is unknown. We characterized earthworm populations of two grassland types within the Palouse region: native prairie remnants and Conservation Reserve Program (CRP) set asides planted with exotic grasses. The earthworm community in both grassland types was completely dominated by the exotic-invasive Aporrectodea trapezoides. Only one individual of a native species, Driloleirus americanus (the giant Palouse earthworm), was found in a prairie remnant. No differences were found between prairie remnants and CRP sites for mean earthworm density (24–106 individuals m⁻²) or fresh weight (12–45 g m⁻²). Our results suggest that the combined effects of land-use change, habitat fragmentation and competitive interactions have resulted in the decimation of native earthworm populations and dominance of invasive earthworms in native and non-native grasslands of the Palouse region.     

Coupling long-term studies with meta-analysis to investigate impacts of non-native crayfish on zoobenthic communities

1. Biological invasions are widely recognised as a significant component of human‐caused environmental change and a primary threat to native biodiversity. The negative impacts of species invasions are particularly evident for freshwater crayfish faunas. 2. This study provides novel insight into the ecological effects of native and non‐native crayfish on zoobenthic communities (with emphasis on the non‐native rusty crayfish, Orconectes rusticus) across broad scales by combining a meta‐analysis of small‐scale experimental studies with a long‐term observational study conducted over a 24 year period in Sparkling Lake, Wisconsin, U.S.A. (46°00′N, 89°42′W). 3. The meta‐analysis summarised quantitatively the results of cage experiments for seven species of crayfish spanning four continents. We found that total zoobenthos densities (primarily Gastropoda and Diptera) were significantly lower in treatments containing crayfish relative to controls; a result that was significant for non‐native crayfish but not for crayfish in their native range, perhaps owing to a small sample size. In contrast to other species, rusty crayfish were also negatively associated with Ephemeroptera. 4. Results from the time series analysis comparing temporal trends in rusty crayfish and invertebrate abundances from Sparkling Lake were consistent with the findings from the meta‐analysis. Rusty crayfish were negatively correlated with the abundance of total zoobenthos, Diptera, Ephemeroptera and Odonata, as well as families of Trichoptera. 5. By coupling the results from short and long‐term research, our study offers greater insight into the nature of crayfish‐invertebrate interactions in aquatic systems, revealing consistent effects of invasive crayfish on native fauna. The control and management of invasive species is facilitated by the knowledge that well executed small‐scale studies may be extrapolated to understand larger‐scale ecological interactions.     

Ecosystem effects of non-native earthworms in Mid-Atlantic deciduous forests

In many mid-Atlantic forests where both native and non-native earthworms exist, it is the non-native species that are the dominant component of the soil macrofauna. Few earthworm ecology studies, however, focus attention on these forest systems in order to determine the relative ecological roles and potential interactions of the native and non-native earthworms. In a series of field samplings and experimental manipulations we collected data on the effects of earthworms on below-and aboveground ecosystem processes. Earthworm abundance and the ecological processes measured were dynamic in space and time across the range of study sites. Leaf litter decay rates doubled at sites that had abundant non-native earthworms. Earthworms also altered the abundance of soil fungi, the activity of extracellular enzymes, soil respiration, and the growth of tree seedlings but the effects varied among sites depending on differences in land-use history and forest age. Red oak seedling growth was less at sites that had abundant earthworms but tulip poplar and red maple seedlings grew equally well with and without abundant earthworms. These preliminary results suggest that non-native earthworms have significant ecosystem effects, even in forests where native earthworms still occur. Land use history, however, plays an important role in determining what those effects will be, and these effects are likely to be dynamic, depending on the abundance of non-native earthworms.     

Trophic ecology of the Atlantic blue crab Callinectes sapidus as an invasive non-native species in the Aegean Sea

Invasive non-native crustaceans are a biodiversity and management concern in the Mediterranean Sea. The Atlantic blue crab (Callinectes sapidus) was first recorded in the Mediterranean Sea in 1949, but may have arrived as early as in the 1930’s. Blue crabs in the Mediterranean Sea are of concern due to their presumed potential for negative consumptive and competitive interactions with native fauna. The aim of this study was to provide a first assessment of the trophic ecology of non-native blue crab in the Northern Aegean Sea using stable carbon (δ¹³C) and nitrogen isotope (δ¹⁵N) analysis. We found limited isotopic niche overlap between blue crabs and seven native species examined at Gökçeada Island in April, June, and August of 2017. In addition, the range of calculated trophic positions of blue crabs at Gökçeada Island (2.0 to 4.4), while broad, is in general agreement with prior studies in both native and non-native ranges. We also observe that trophic position declined and the relative importance of pelagic carbon sources to blue crabs increased from April to August. However, we also found that differing assumptions as to the number and type of food web baselines and trophic discrimination factors led to differing estimates of trophic position in blue crabs at Gökçeada Island by as much as one to two trophic levels. These methodical differences make it challenging to directly compare results within and between studies, and thus limit our ability to assess negative consumptive and competitive interactions of invasive blue crab with native coastal species in the Mediterranean Sea.

     

Susceptibility of native and non-native common reed to the non-native mealy plum aphid (Homoptera: Aphididae) in North America

An aggressive, non-native haplotype (distinct genetic lineage within a species) of Phragmites australis is invading brackish and freshwater systems in the eastern United States, potentially displacing native haplotypes. We studied the differential susceptibility of native and non-native populations collected from sites throughout North America to the non-native aphid, Hyalopterus pruni. In a greenhouse study, we found significantly higher aphid populations on native haplotypes than on the non-native haplotype 2 mo after infestation. Aphid feeding caused chlorosis and death of native stems, and in some cases, killed whole native genets. The non-native plants remained relatively undamaged. In a field study, non-native plants had significantly lower aphid densities than native plants or remained aphid free. There was an interactive effect in which aphid populations increased on the native plants over the 1-mo study period but remained low on non-native plants over the same period. The susceptibility of native North American populations of P. australis to non-native aphid infestation may indirectly affect the ability of these native plants to compete with non-native plant populations, ultimately contributing to the decline of native haplotypes.     

Response of semi-desert grasslands invaded by non-native grasses to altered disturbance regimes

Aim Using a long‐term data set we investigated the response of semi‐desert grasslands to altered disturbance regimes in conjunction with climate patterns. Specifically, we were interested in the response of a non‐native grass (Eragrostis lehmanniana), mesquite (Prosopis velutina), and native species to the reintroduction of fire and removal of livestock. Location The study site is located on the 45,360‐ha Buenos Aires National Wildlife Refuge (31°32′ N, 110°30′ W) in southern Arizona, USA. In 1985, livestock were removed and prescribed fires were reintroduced to this semi‐desert grassland dominated by non‐native grasses and encroaching mesquite trees. Methods Plant species cover was monitored along 38, 30‐m transects five times over a period of 15 years. Data were analysed using principal components analysis on the variance–covariance and correlation matrix, multivariate analysis of variance for changes over time in relation to environmental data, and analysis of variance for altered disturbance regimes. Results Reintroduction of fire and removal of livestock have not led to an increase in native species diversity or a decrease in non‐native grasses or mesquite. The cover of non‐native grass was influenced by soil type in 1993. Main conclusions Variability of plant community richness, diversity, and cover over time appear to be most closely linked to fluctuations in precipitation rather than human‐altered disturbance regimes. The effects of altered grazing and fire regimes are likely confounded by complex interactions with climatic factors in systems significantly altered from their original physiognomy.     

The establishment success of native versus non-native herbaceous seed mixes on a revegetated roadside in Central Texas

Revegetation is an essential component of roadside and building site construction and improvement. In the southern United States, non-native grass species are frequently included in revegetation seed mixes used by highway authorities. Non-native species are frequently selected for aggressive growth characteristics; however, these same traits also render them potentially invasive, and subsequently hazardous to adjacent plant communities. Although the use of pure native seed mixes has been rejected in the past due to perceived inferior establishment characteristics, there have been few comparative quantitative field studies that justify this belief. The establishment characteristics of three seed mixes—one containing non-native species and two with native grass and forb species only—were compared in a randomized-block design along a Texas roadside following spring and summer sowing. After 60 days following the spring sowing, the two native-only seed mixes demonstrated 180 and 560% (F = 10.18; P < 0.0001) higher seed densities than the recommended native/non-native mix. The summer sowing results were similar with seedling densities 180 and 330% (F = 9.20; P < 0.01) greater than the standard non-native seeding. Although an aggressive colonizer from vegetative tissue such as stolons and rhizomes, the non-native Bermudagrass (Cynodon dactylon) had a lower than expected establishment rate thought to be due to high water demand during the first week following sowing. Given the invasive characteristics of this common component of many recommended revegetation seed mixes, these results call into question the widespread recommended use of Bermudagrass for such projects. Although gathered during 1 year only, these data indicate that examination of suites of early- and late-successional native species can provide a highly effective mix for revegetation projects. Furthermore, this reduces the potential for negative ecological consequences and provides added benefits associated with wholly native plant communities.     

Environmental niche patterns of native and non-native fishes within an invaded African river system

To test ecological niche theory, this study investigated the spatial patterns and the environmental niches of native and non-native fishes within the invaded Great Fish River system, South Africa. For the native fishes, there were contrasting environmental niche breadths that varied from being small to being large and overlapped for most species, except minnows that were restricted to headwater tributaries. In addition, there was high niche overlap in habitat association among fishes with similar distribution. It was therefore inferred that habitat filtering-driven spatial organisation was important in explaining native species distribution patterns. In comparison, most non-native fishes were found to have broad environmental niches and these fishes showed high tolerance to environmental conditions, which generally supported the niche opportunity hypothesis. The proliferation of multiple non-native fishes in the mainstem section suggest that they form a functional assemblage that is probably facilitated by the anthropogenic modification of flow regimes through inter-basin water transfer. Based on the distribution patterns observed in the study, it was inferred that there was a likelihood of negative interactions between native and non-native fishes. Such effects are likely to be exacerbated by altered flow regime that was likely to have negative implications for native ichthyofauna.     

Atmospheric invasion of non-native pollen in the Mediterranean region

Most research on the impacts of plant invasion focuses on native plant performance, community structure, and ecosystem functioning. Some non-native species can also pose a risk to human health. One such risk is the allergenic nature of the pollen of some introduced plants. We examined whether patterns of airborne pollen differed between non-native and native taxa by summarizing data from seven Spanish Mediterranean localities monitored over 13 yr. The pollen spectra contained 27 native pollen taxa and 18 non-native taxa. Even though pollen from native taxa were more diverse and were present longer in the atmosphere than the non-native, in some years neither the prevalence of the two nor their weekly maximum pollen values differed significantly. However, maximum values for non-native taxa were found earlier in the season than for native pollen. A small percentage of non-native pollen includes pollen from introduced taxa that have not invaded natural habitats (e.g., ornamental plants). Non-native pollen has a larger proportion of allergenic pollen than native pollen. Therefore, the results reveal that the presence of non-native airborne pollen from naturalized and non-naturalized plant species increases the total amount of airborne allergenic pollen grains and the period of allergenic susceptibility.     

The biodiversity impacts of non-native species should not be extrapolated from biased single-species studies

The presence, diversity and abundance of non-native plant species in natural vegetation are common condition indicators used to determine conservation status, with consequences for management strategies and investment. The rationale behind non-native species metrics as condition indicators is the assumption that non-natives have negative consequences on native biodiversity and habitat condition. The case against non-native species is not so clear-cut, with some studies reporting neutral or even facilitative interactions, often depending on spatial scale. Observational and experimental evaluations of the impact of particular non-native species on biodiversity provide a vital evidence-base for general conservation management strategies. Unintentionally though, many studies that quantify the impacts of non-native species have resulted in a publication bias in which species with known impacts are selected for investigation far more often than benign species. Here we argue that meta-analyses of the impacts of individual non-native species on natives, no matter how meticulous or objective, should not be generalized beyond the set of ‘training’ species. The likelihood of such extrapolation is increased when meta-analyses are reported with little qualification as to the skewed sampling towards problematic species, and because alternative findings such as non-native assemblages having positive interactions with native biodiversity, are under-reported. To illustrate, we discuss two meta-analyses that make general conclusions from impact studies skewed towards ‘transformers’, the most extreme invaders. We warn that if generic non-native species management strategies were to be based on these conclusions, they could not only fail to meet objectives but in some instances harm native biodiversity.